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UBC Theses and Dissertations

Moose habitat selection in relation to forest cutting practices in northcentral British Columbia Schwab, Francis Edward 1985

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MOOSE HABITAT SELECTION IN RELATION TO FOREST CUTTING PRACTICES IN NORTHCENTRAL BRITISH COLUMBIA By FRANCIS EDWARD SCHWAB 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 , 1974 M.Sc. , 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 , 1979 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY i n THE FACULTY OF GRADUATE STUDIES (Dep a r t m e n t of P l a n t S c i e n c e ) We a c c e p t t h i s t h e s i s as c o n f o r m i n g 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 1985 © F r a n c i s Edward Schwab, 1985 7 8 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of Plant Scienr.p The University of British Columbia 1956 Main Mall Vancouver, Canada V6T 1Y3 Date 1 0 F p h n i a r y IQRfi DE-6(3/81) A b s t r a c t T h i s t h e s i s i s p r i m a r i l y d i r e c t e d a t d o c u m e n t i n g moose (AJLc_ £ j 3 a ^  c. £js a n d e r_^ o n± i ) h a b i t a t s e l e c t i o n i n an a r e a c o n t a i n i n g a d i v e r s i t y o f a g e s and s t y l e s o f l o g g i n g . The h a b i t a t o c c u p a t i o n p a t t e r n s o f r a d i o c o l l a r e d moose w e r e r e c o r d e d d u r i n g t h e p e r i o d A p r i l 1980 t o May 1983 i n an a r e a c e n t r e d 50 km n o r t h e a s t o f P r i n c e G e o r g e B.C. The h a b i t a t a s s o c i a t i o n s of t h e r a d i o c o l l a r e d moose were t e s t e d a g a i n s t t h e f o l l o w i n g a l t e r n a t e h y p o t h e s e s . F o r t h e w i n t e r p e r i o d : (1) Moose a r e n o t r a n d o m l y d i s t r i b u t e d b u t s e l e c t h a b i t a t s w i t h g r e a t e r amounts of f o r a g e e x c e p t when snow d e p t h e x c e e d s 90 cm. I f snow d e p t h e x c e e d s 90 cm, t h e n moose w i l l s e l e c t h a b i t a t s w i t h g r e a t e r f o r e s t c o v e r . (2) Moose a r e n o t r a n d o m l y d i s t r i b u t e d but s e l e c t t h e h a b i t a t w i t h t h e g r e a t e s t f o r e s t c o v e r a s p r o t e c t i o n f r o m c o l d s t r e s s when a i r t e m p e r a t u r e s a r e l o w e r t h a n -20°C. F o r t h e summer ( n o n w i n t e r ) p e r i o d : (1) Moose a r e n o t r a n d o m l y d i s t r i b u t e d b u t s e l e c t h a b i t a t s w i t h g r e a t e r a m o u n t s o f f o r a g e . (2) Moose a r e n o t r a n d o m l y d i s t r i b u t e d b u t w i l l s e l e c t h a b i t a t s w i t h g r e a t e r f o r e s t c o v e r when a i r t e m p e r a t u r e s e x c e e d 20°C. M o ose f o l l o w e d a g e n e r a l a n n u a l p a t t e r n t h a t v a r i e d a c c o r d i n g t o t h e c o n d i t i o n s p r e v a l e n t d u r i n g i n d i v i d u a l s e a s o n s . I n s p r i n g , moose w e r e f o u n d i n open h a b i t a t s . The s p r i n g t o summer change of s e a s o n saw moose move i n t o h a b i t a t i i i c h a r a c t e r i z e d by h i g h f o r e s t c o v e r v a l u e s . I n f a l l , moose w e r e a g a i n a s s o c i a t e d w i t h h a b i t a t s w i t h c l o s e d f o r e s t c a n o p i e s . E a r l y w i n t e r p r o d u c e d a d r a m a t i c s h i f t o f moose l o c a t i o n s i n t o open h a b i t a t s . In m i d w i n t e r and l a t e w i n t e r , moose r e t r e a t e d t o h a b i t a t s w i t h heavy f o r e s t c o v e r . Moose were u s u a l l y n o n r a n d o m l y d i s t r i b u t e d w i t h r e s p e c t t o h a b i t a t t y p e . However, the a l t e r n a t e h y p o t h e s e s g e n e r a l l y f a i l e d t o a c c o u n t f o r t h e d e p a r t u r e s f r o m r a n d o m n e s s . In s p r i n g , h a b i t a t a s s o c i a t i o n s s u g g e s t e d s e l e c t i o n f o r h a b i t a t s w i t h g r e a t e r f o r a g e b i o m a s s . H o w e v e r , t h e c o r r e l a t i o n s b e t w e e n moose h a b i t a t a s s o c i a t i o n s and a v a i l a b l e f o r a g e b i o m a s s w e r e n o t c o n s i s t e n t l y h i g h and one o f f o u r s p r i n g p e r i o d s p r o d u c e d a n e g a t i v e c o r r e l a t i o n b e t w e e n a v a i l a b l e f o r a g e and h a b i t a t a s s o c i a t i o n s . I n summer, moose h a b i t a t a s s o c i a t i o n s c o u l d be e x p l a i n e d by moose a v o i d i n g h e a t s t r e s s . F a l l h a b i t a t a s s o c i a t i o n s were nev e r e x p l i c a b l e by t h e t e s t e d a l t e r n a t e h y p o t h e s e s . In e a r l y w i n t e r , h a b i t a t a s s o c i a t i o n s once a g a i n s u g g e s t e d t h a t moose were s e l e c t i n g h a b i t a t w h i c h p r o d u c e d p l e n t i f u l f o r a g e . D u r i n g one e a r l y w i n t e r , however, t h e r e was a h i g h e r c o r r e l a t i o n b e t w e e n m o o s e h a b i t a t a s s o c i a t i o n s a n d f o r e s t c o v e r t h a n b e t w e e n h a b i t a t a s s o c i a t i o n s and a v a i l a b l e f o r a g e . I n one m i d w i n t e r s e a s o n moose h a b i t a t a s s o c i a t i o n s were c o n s i s t e n t w i t h th e h y p o t h e s i s t h a t moose s e e k h a b i t a t w i t h h e a v y f o r e s t c o v e r when snow d e p t h s e x c e e d 90 cm, b u t a v o i d a n c e o f c o l d s t r e s s w o u l d have i v p r o d u c e d t h e same p a t t e r n of h a b i t a t a s s o c i a t i o n . N e i t h e r of th e t e s t e d a l t e r n a t e h y p o t h e s e s c o u l d e x p l a i n moose h a b i t a t a s s o c i a t i o n s i n t h e o t h e r m i d w i n t e r s e a s o n , n o r i n two l a t e w i n t e r s e a s o n s . C o m p a r i s o n s o f c a l c u l a t e d e q u i v a l e n t b l a c k b o d y t e m p e r a t u r e s ( T g ) t o known u p p e r c r i t i c a l l i m i t s c o n f i r m e d t h a t h e a t s t r e s s i s an i m p o r t a n t f a c t o r f o r moose i n summer. F u r t h e r , T g e s t i m a t e s r e v e a l e d t h a t h e a t s t r e s s and not c o l d , was a f f e c t i n g moose i n w i n t e r and t h a t t h e o b s e r v e d p a t t e r n of w i n t e r h a b i t a t a s s o c i a t i o n s c o u l d be e x p l a i n e d by moose a v o i d i n g heat s t r e s s . T h e r e a r e h y p o t h e s e s w h i c h w e r e n o t e x p l i c i t l y t e s t e d w h i c h c o u l d e x p l a i n moose h a b i t a t a s s o c i a t i o n s u n e x p l a i n e d by th e a l t e r n a t e h y p o t h e s e s : moose may not s e l e c t h a b i t a t on t h e b a s i s o f p r e v a i l i n g c o n d i t i o n s , b u t r a t h e r ( 1 ) may be e x h i b i t i n g p a t t e r n s l e a r n e d f r o m t h e i r dams. (2) Moose a v o i d a n c e o f open a r e a s i n f a l l c o u l d be c a u s e d by t h e i r a v o i d a n c e o f h u m a n a c t i v i t y . F i n a l l y , ( 3 ) t h e l a r g e p r o p o r t i o n o f moose l o c a t i o n s i n h e a v y f o r e s t c o v e r s i n a l l s e a s o n s c o u l d be a p r e d a t o r a v o i d a n c e s t r a t e g y . The h y p o t h e s i s t h a t moose r e q u i r e f o r e s t c o v e r t o p r o t e c t them f r o m h e a t s t r e s s was much more s u c c e s s f u l a t p r e d i c t i n g moose h a b i t a t a s s o c i a t i o n s t h a n was t h e h y p o t h e s i s t h a t moose s h o u l d s e e k h a b i t a t p r o d u c i n g l a r g e a m o u n t s o f f o r a g e V c o n d i t i o n a l upon snow d e p t h b e i n g l e s s t h a n 90 cm. I n f a c t , f o r a g e p r o d u c t i o n c o n d i t i o n a l upon snow d e p t h c o u l d n o t u n e q u i v o c a l l y e x p l a i n the p a t t e r n of moose h a b i t a t a s s o c i a t i o n i n any s e a s o n . However, t h i s r e s u l t s h o u l d be i n t e r p r e t e d as s u p p o r t i n g the need f o r mature t r e e s on moose ra n g e , t o a v o i d h e a t s t r e s s , and n o t t o m i n i m i z e c o n c e r n f o r moose f o r a g e r e q u i r e m e n t s . I n any s e a s o n , good moose r a n g e w i l l c o n t a i n b o t h f o r a g e and mature t r e e s . On t h e s t u d y a r e a , t h e h e t e r o g e n o u s i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g s h o u l d be c o n s e r v e d as moose h a b i t a t , e s p e c i a l l y w h e r e i t o c c u r s on moose w i n t e r r a n g e . Moose h a b i t a t c a n be c o n s e r v e d o r i m p r o v e d w i t h 4 ha c l e a r c u t s . Where l o g g i n g e c o n o m i c s demand l a r g e c l e a r c u t s , c o v e r b l o c k s and t r a v e l c o r r i d o r s s h o u l d be p r o v i d e d t o c o n s e r v e moose h a b i t a t . v i T a b l e of C o n t e n t s A b s t r a c t i i T a b l e of C o n t e n t s v i L i s t of T a b l e s v i i i L i s t of F i g u r e s i x Acknowledgements x i I INTRODUCTION 1 1. L i t e r a t u r e Review 1 1.1 W i n t e r H a b i t a t S e l e c t i o n 2 1.2 Summer H a b i t a t S e l e c t i o n 7 2. Summary and H y p o t h e s e s 11 I I STUDY AREA 13 1. Geography 13 2. L o g g i n g H i s t o r y 18 I I I METHODS 19 1. H a b i t a t S e l e c t i o n 19 1.1 H a b i t a t Maps 19 1.2 Moose L o c a t i o n s 22 1.3 S t a t i s t i c a l A n a l y s i s 24 2. F o r a g e Biomass P r o d u c t i o n 28 2.1 Shrub Biomass P r o d u c t i o n 28 2.1.1 R e g r e s s i o n e q u a t i o n s 29 2.1.2 Shrub d e n s i t y e s t i m a t e s 33 2.1.3 Shrub biomass 35 2.1.4 V a r i a n c e of s h r u b biomass p r o d u c t i o n 36 2.2 Herb L a y e r B i omass E s t i m a t i o n 38 2.3 S e a s o n a l Biomass E s t i m a t e s 39 3. F o r e s t C o v e r 41 4. Snow Depth 42 5. A i r T e m p e r a t u r e 43 IV RESULTS 44 1. S p r i n g 48 2. Summer 62 3. F a l l 66 4. E a r l y W i n t e r 69 5. M i d w i n t e r 72 6. L a t e W i n t e r 81 7. Summary 84 7.1 R e l e v a n c e o f Working H y p o t h e s e s 85 v i i V DISCUSSION 87 1. F o r a g e P r o d u c t i o n C o n d i t i o n a l Upon Snow Depth 87 2. T h e r m a l S t r e s s 92 2.1 C a l c u l a t i o n of E q u i v a l e n t B l a c k b o d y T e m p e r a t u r e 93 2.1.1 A i r t e m p e r a t u r e 94 2.1.2 A b s o r b e d r a d i a t i o n 94 2.1.3 R e s i s t a n c e t o h e a t t r a n s f e r a n d T e 97 2.2 W i n t e r C o l d S t r e s s 99 2.3 W i n t e r Heat S t r e s s 101 2.4 Summer Heat S t r e s s 107 3. U n t e s t e d H y p o t h e s e s 115 4. F o r a g e Biomass E s t i m a t i o n T e c h n i q u e s 118 4.1 Shrub Biomass E s t i m a t i o n 118 4.2 Herb L a y e r B i omass E s t i m a t i o n 121 5. A s s o c i a t i o n I n d i c e s 122 6. N o n f o r e s t H a b i t a t s 126 6.1 U p l a n d Shrub 126 6.2 R i p a r i a n Shrub 135 6.3 W e t l a n d s 136 6.4 L a k e s 136 6.5 F r a s e r R i v e r 137 7. Moose H a b i t a t Management 138 7.1 Management I m p l i c a t i o n s of R e s u l t s 138 7.2 Management I m p o r t a n c e of S p e c i f i c H a b i t a t s 142 7.2.1 U n l o g g e d f o r e s t 142 7.2.2 D i a m e t e r l i m i t l o g g i n g 144 7.2.3 I n t e r m e d i a t e u t i l i z a t i o n l o g g i n g . . . 145 7.2.4 O l d c l e a r c u t s 147 7.2.5 R e c e n t c l e a r c u t s 149 7.2.6 N o n f o r e s t h a b i t a t s 150 7.3 F o r e s t C u t t i n g Recommendations 152 7.3.1 L i t e r a t u r e r e v i e w and p r a c t i c e s on and near t h e s t u d y a r e a 152 7.3.2 C u t t i n g recommendations f o r f o r e s t 156 7.3.3 N o n f o r e s t h a b i t a t s c u r r e n t p r a c t i c e s and r ecommendations 158 VI SUMMARY AND CONCLUSIONS 161 V I I LITERATURE CITED 167 v i i i L i s t of T a b l e s T a b l e 1. A r e a o c c u p i e d by h a b i t a t t y p e s on s p r i n g t o e a r l y w i n t e r and mid- t o l a t e w i n t e r moose r a n g e s 21 T a b l e 2. C o l e (Cy) i n d e x o f a s s o c i a t i o n 26 T a b l e 3. Number o f moose l o c a t i o n s i n f o r e s t / l o g g e d h a b i t a t s by y e a r and s e a s o n 47 T a b l e 4. C o l e i n d e x and s t a n d a r d e r r o r ( i n b r a c k e t s ) f o r f o r e s t / l o g g e d h a b i t a t s 56 T a b l e 5. Summer a v a i l a b l e f o r a g e (g/m ) 57 T a b l e 6. E s t i m a t e s of f o r e s t c o v e r as a d e c i m a l f r a c t i o n of t h e sky h e m i s p h e r e 58 T a b l e 7. C o r r e l a t i o n s between C o l e i n d i c e s and a v a i l a b l e f o r a g e and between C o l e i n d i c e s and f o r e s t c o v e r 59 T a b l e 8. S e a s o n a l mean minimum and mean maximum a i r t e m p e r a t u r e s (°C) 61 T a b l e 9. M o n t h l y mean minimum and mean maximum a i r t e m p e r a t u r e s (°C) 64 T a b l e 10. F a l l a v a i l a b l e f o r a g e (g/m 2) 68 T a b l e 11. E a r l y w i n t e r a v a i l a b l e f o r a g e (g/m ) 71 T a b l e 12. Mean snow d e p t h s i n f o r e s t / l o g g e d h a b i t a t s d u r i n g t h e w i n t e r of 1981-1982 77 T a b l e 13. Mean snow d e p t h s i n f o r e s t / l o g g e d h a b i t a t s d u r i n g t h e w i n t e r of 1982-1983 78 T a b l e 14. M i d w i n t e r a v a i l a b l e f o r a g e (g/m ) 80 T a b l e 15. L a t e w i n t e r a v a i l a b l e f o r a g e (g/m ) 83 T a b l e 16. Number of moose l o c a t i o n s i n n o n f o r e s t h a b i t a t s by y e a r and s e a s o n 134 i x L i s t of F i g u r e s F i g u r e 1. L o c a t i o n o f t h e s t u d y a r e a w i t h i n B r i t i s h C o l u m b i a 14 F i g u r e 2. B o u n d a r i e s o f t h e s t u d y a r e a , l o c a t i o n s o f p l o t s , and t h e r m o g r a p h s on a 1:250,000 s c a l e map 16 F i g u r e 3. P r o p o r t i o n of moose l o c a t i o n s i n f o r e s t /1 o g g ed h a b i t a t t y p e s 45 F i g u r e 4. The C o l e ( C y ) i n d e x of a s s o c i a t i o n between moose l o c a t i o n s and f o r e s t / l o g g e d h a b i t a t t y p e s f r o m s p r i n g 1980 t o f a l l 1980 50 F i g u r e 5. The C o l e ( C y ) i n d e x o f a s s o c i a t i o n between moose l o c a t i o n s and f o r e s t / l o g g e d h a b i t a t t y p e s f r o m s p r i n g 1981 t o l a t e w i n t e r 1982 52 F i g u r e 6. The C o l e ( C y ) i n d e x of a s s o c i a t i o n between moose l o c a t i o n s and f o r e s t / l o g g e d h a b i t a t t y p e s from s p r i n g 1982 t o s p r i n g 1983 54 F i g u r e 7. Mean snow d e p t h s i n f o r e s t / 1 o g g e d h a b i t a t s d u r i n g t h e w i n t e r of 1981-1982 73 F i g u r e 8. Mean snow d e p t h s i n f o r e s t / 1 o g g e d h a b i t a t s d u r i n g t h e w i n t e r of 1982-1983 75 F i g u r e 9. E q u i v a l e n t b l a c k b o d y t e m p e r a t u r e ( T e ) d u r i n g an a v e r a g e d a i l y c y c l e i n m i d w i n t e r 1982 ( t o p ) and m i d w i n t e r 1983 ( b o t t o m ) 103 F i g u r e 10. E q u i v a l e n t b l a c k b o d y t e m p e r a t u r e ( T e ) d u r i n g an a v e r a g e d a i l y c y c l e summer 1981 ( t o p ) and summer 1982 ( b o t t o m ) 110 F i g u r e 11. E q u i v a l e n t b l a c k b o d y t e m p e r a t u r e ( Te) d u r i n g an a v e r a g e d a i l y c y c l e J u l y and August 1981 ( t o p ) and J u l y and A u g u s t 1982 ( b o t t o m ) 112 F i g u r e 12. C o l e (Cy) i n d e x o f a s s o c i a t i o n between moose l o c a t i o n s and n o n f o r e s t h a b i t a t t y p e s from s p r i n g 1980 t o f a l l 1980 128 X F i g u r e 1 3 . C o l e ( C y ) i n d e x o f a s s o c i a t i o n b e t w e e n m o o s e l o c a t i o n s a n d n o n f o r e s t h a b i t a t t y p e s f r o m s p r i n g 1 9 8 1 t o l a t e w i n t e r 1 9 8 2 1 3 0 F i g u r e 1 4 . C o l e ( C y ) i n d e x o f a s s o c i a t i o n b e t w e e n m o o s e l o c a t i o n s a n d n o n f o r e s t h a b i t a t t y p e s f r o m s p r i n g 1 9 8 2 t o s p r i n g 1 9 8 3 132 x i A cknowledgements The c o n t r i b u t i o n s of t h e f o l l o w i n g p e r s o n s a r e a c k n o w l e d -ged f o r t h e i n d i c a t e d r e a s o n s : Dr. M. D. P i t t , f o r s u p e r v i s -i n g t h e r e s e a r c h ; Dr. V. C. R u n e c k l e s a s C h a i r m a n o f t h e r e s e a r c h c o m m i t t e e ; Dr. T. A. B l a c k , Dr. F. L. B u n n e l l , Dr. D. S. E a s t m a n , Dr. A. R. E. S i n c l a i r , and Dr. W. G. W e l l i n g t o n f o r s e r v i n g as members of t h e r e s e a r c h and c o m p r e h e n s i v e exam c o m m i t t e e s ; Mr. B i l l A r t h u r , Mr. Ken C h i l d , Mr. Doug J a n z , Mr. Ken F u j i n o , Dr. D a r r y l H e b e r t , Mr. Don M o r b e r g , and Mr. S t e v e W i l l e t t were members of t h e B.C. M i n i s t r y of E n v i r o n m e n t F i s h and W i l d l i f e B r a n c h who c o n t r i b u t e d t o t h e s t u d y ; Mr. G e r r y L y n c h of t h e A l b e r t a F i s h and W i l d l i f e p r o v i d e d h i s knowledge o f moose i m m o b i l i z a t i o n t o t h e s t u d y ; Dr. J o h n S m i t h o f t h e C a n a d i a n W i l d l i f e S e r v i c e p r o v i d e d a computer program w h i c h a n a l y s e s C o r r e c t e d P o i n t D i s t a n c e e s t i m a t e s o f d e n s i t y and i n s t r u c t i o n on i t s u s e ; Mr. Doug Cowen, Mr. G r aham F a r s t a d , Mrs. Leanna Head, Mr. S t e v e Head, and Mr. G r a n t Hazelwood were e s p e c i a l l y h e l p f u l m e m b e r s o f S p r u c e C i t y W i l d l i f e A s s o c i a t i o n ; Mr. Bob W e i n a r d o f N o r t h w o o d P u l p and T i m b e r L t d . ; M r s . S u s a n S t e v e n s o n and Mr. D a v i d S t e v e n s o n p r o v i d e d S u s a n and m y s e l f w i t h a p l a c e t o s t a y i n P r i n c e G e o r g e ; Mr. G a r y B e l l and Mr. G r e g C h e e s m a n o f t h e B r i t i s h C o l u m b i a M i n i s t r y o f E n v i r o n m e n t A i r S t u d i e s B r a n c h p r o v i d e d t h e r m o g r a p h s and i n s t r u c t i o n s on t h e i r o p e r a t i o n ; Mr. B i l l x i i G u r n e y and Mr. R o r y C o n r o y o f C r u i s e A i r L t d . ; Mr. B a r r y Wong w r o t e a c o m p u t e r p r o g r a m t o c a l c u l a t e t h e v a r i a n c e o f a m u l t i p l e r e g r e s s i o n . T h i s work w o u l d n o t h a v e been p o s s i b l e w i t h o u t t h e c o o p e r a t i o n and h a r d work o f my w i f e S u s a n Schwab n o r t h e m o r a l s u p p o r t of my f a t h e r and mother Wendel and Mary Schwab. F i n a n c i a l s u p p o r t f o r t h i s s t u d y came f r o m B r i t i s h C o l u m b i a M i n i s t r y of E n v i r o n m e n t , B r i t i s h C o l u m b i a R e s e a r c h C o u n c i l , Northwood P u l p and T i m b e r L t d . , S p r u c e C i t y W i l d l i f e A s s o c i a t i o n , and t h e 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 . 1 I INTRODUCTION H i s t o r i c a l l y , t h e m o s t i m p o r t a n t p r o d u c e r o f q u a l i t y moose ( A l c e s a l c e s a n d e r s o n i i ) h a b i t a t was w i l d f i r e ( P e t e r s o n 1955, K r e f t i n g 1974a, E a s t m a n 1 9 7 8 ) . S i n c e t h e r e d u c t i o n o f w i l d f i r e by more e f f i c i e n t f i r e f i g h t i n g t e c h n i q u e s , l o g g i n g h a s been t h e most i m p o r t a n t d i s t u r b a n c e i n moose h a b i t a t i n n o r t h c e n t r a l B r i t i s h C o l u m b i a ( E a s t m a n 1978). T h i s t h e s i s f o l l o w s t h e h a b i t a t o c c u p a t i o n by r a d i o c o l l a r e d moose, p r i n c i p a l l y w i t h r e s p e c t t o l o g g i n g p a t t e r n s , i n an a r e a c e n t e r e d 50 km n o r t h e a s t o f P r i n c e G e o r g e B.C., and i s g u i d e d by two c e n t r a l q u e s t i o n s : (1) Do moose s e l e c t l o g g e d a r e a s b e c a u s e t h e l o g g i n g h a s s t i m u l a t e d a b u n d a n t . g r o w t h o f f o o d p l a n t s ? (2) Does t h e e l i m i n a t i o n of the f o r e s t canopy c a u s e c o n d i t i o n s w h i c h w o u l d s u b j e c t moose t o t h e r m a l s t r e s s and c a u s e them t o a v o i d l o g g e d a r e a s ? 1. L i t e r a t u r e Review The f o l l o w i n g l i t e r a t u r e r e v i e w documents o b s e r v e d moose h a b i t a t s e l e c t i o n p a t t e r n s i n w i n t e r and i n summer. F o r t h e p u r p o s e s o f t h i s i n t r o d u c t i o n summer r e f e r s t o a l l s e a s o n s o t h e r t h a n w i n t e r . F u r t h e r , t h i s r e v i e w s e e k s t o e s t a b l i s h t h e p r o b a b l e c a u s e s o f moose h a b i t a t s e l e c t i o n , l e a d i n g t o moose h a b i t a t s e l e c t i o n h y p o t h e s e s . 2 1.1 W i n t e r H a b i t a t S e l e c t i o n One o f two h y p o t h e s e s i s u s u a l l y i m p l i c a t e d i n moose h a b i t a t s e l e c t i o n d u r i n g w i n t e r . The f i r s t of t h e s e , and t h e one most o f t e n u s e d t o e x p l a i n m o o s e / h a b i t a t r e l a t i o n s i n w i n t e r , i s t h a t moose w i l l o c c u p y o pen a r e a s w h i c h p r o d u c e l a r g e a m o u n t s o f b r o w s e e x c e p t when snow e x c e e d s a c r i t i c a l d e p t h . I f snow e x c e e d s a c r i t i c a l d e p t h , t h e n moose w i l l r e t r e a t t o a r e a s w i t h c l o s e d f o r e s t c a n o p i e s w h e r e t h e r e i s r e d u c e d snow de p t h . The se c o n d h y p o t h e s i s i s t h a t moose w i l l be found under c l o s e d f o r e s t c a n o p i e s i n w i n t e r t o a v o i d c o l d s t r e s s . Many a u t h o r s h a v e b e e n i n t e r e s t e d i n t h e r e l a t i o n s h i p b e t w e e n snow and moose h a b i t a t . T h e r e h a s b e e n p r o p o s e d a c r i t i c a l d e p t h of snow w h i c h c a u s e s w i n t e r h a b i t a t s h i f t s of moose. N a s i m o v i c h ( 1 9 5 5 ) f e l t t h a t snow d e e p e r t h a n 90 t o 100 cm r e p r e s e n t s a c r i t i c a l d e p t h f o r moose b e c a u s e movement b e c o m e s d i f f i c u l t and t h e number o f w i n t e r moose d e a t h s i n c r e a s e s when t h i s d e p t h i s e x c e e d e d . M o ose e x p e r i e n c e d d i f f i c u l t y r u n n i n g when snow d e p t h s e x c e e d e d 85 cm ( N a s i m o v i c h 1 9 5 5 ) . K e l s a l l ( 1 9 6 9 ) r e a c h e d t h e same c o n c l u s i o n s a s N a s i m o v i c h a f t e r o b s e r v i n g Nova S c o t i a n and New B r u n s w i c k m o o s e . C o a d y ( 1 9 7 4 ) s t a t e d t h a t m o o s e a r e p a r t i a l l y r e s t r i c t e d by snow dee p e r t h a n t w o - t h i r d s c h e s t h e i g h t (>70 cm) and s e v e r e l y r e s t r i c t e d by snow d e e p e r t h a n c h e s t h e i g h t (>100 cm). T h e s e o b s e r v a t i o n s p o i n t t o a " c r i t i c a l " d e p t h of snow i n t h e r e g i o n of 90 cm. I t h a s been c o m m o n l y o b s e r v e d t h a t i n w i n t e r moose s e l e c t h i g h f o o d p r o d u c t i o n a r e a s e x c e p t when snow d e p t h s e x c e e d t h e 90 cm c r i t i c a l d e p t h . When snow e x c e e d s t h e c r i t i c a l d e p t h moose w i l l move u n d e r f o r e s t c a n o p i e s w h i c h i n t e r c e p t snow and t h e r e b y make movement e a s i e r . K r e f t i n g (1974a) n o t e d t h a t i n g e n e r a l t h e moose of n o r t h c e n t r a l N o r t h A m e r i c a a r e fo u n d under dense c o v e r when snow i s d e e p e r t h a n 90 cm. T e l f e r (1970) o b s e r v e d New B r u n s w i c k moose move f r o m open t o dense f o r e s t s when snow d e p t h s e x c e e d e d 100 cm i n t h e open a r e a s . M o n t h e y ( 1 9 8 4 ) r e p o r t e d h e a v y moose u s e o f c l e a r c u t s i n e a r l y w i n t e r b u t a s h i f t t o s o f t w o o d f o r e s t s as snow d e p t h i n c r e a s e d and c r u s t s formed i n m i d w i n t e r . S e v e r a l M i n n e s o t a p a p e r s a l s o s u p p o r t t h e h y p o t h e s i s t h a t moose o c c u p y f o o d p r o d u c i n g a r e a s e x c e p t when snow i s deep. Van B a l l e n b e r g h e and P e e k ( 1 9 7 1 ) n o t e d t h a t moose r e t r e a t e d t o d e n s e f o r e s t s f r o m o pen a r e a s d u r i n g deep snow. B e r g and P h i l l i p s (1972) saw moose move from open w i l l o w a r e a s to more mature w i l l o w - a s p e n f o r e s t s i n l a t e w i n t e r . B e r g and P h i l l i p s ( 1 9 7 4 ) n o t e d t h a t t h e o c c u p a t i o n o f " T a l l - m a t u r e " h a b i t a t s was c o r r e l a t e d w i t h snow d e p t h ( r = 0.706). Peek et a l . (1976) o b s e r v e d moose t h a t s h i f t f r o m open s t a n d s t o dense c o v e r as snow d e p t h s i n c r e a s e d . 4 In Montana moose p r e f e r r e d open a r e a s but r e t r e a t e d t o s p r u c e ( P i cea e n g e l m a n n i / P . g l a u c a ) f o r e s t d u r i n g heavy snow p e r i o d s ( S i n g e r 1 9 7 9 ) . ( S c i e n t i f i c names o f p l a n t s f o l l o w T a y l o r and MacBryde 1977.) Idaho moose s e l e c t e d f o r e s t d u r i n g w i n t e r , p r e s u m a b l y f o r c e d t h e r e by deep snow ( P i e r c e and Peek 1984). S h i r a s moose, r e s i d e n t i n Idaho, Montana, and Wyoming, r e q u i r e f o r e s t c o v e r i n a r e a s where w i n t e r snow d e p t h s e x c e e d 90 cm ( P e e k e t a l . 1 9 82). T e l f e r ( 1 9 7 8 a ) n o t e d t h a t A l b e r t a n moose w i n t e r h a b i t a t s c o i n c i d e d w i t h a r e a s o f h i g h b r o w s e p r o d u c t i o n , b u t snow d e p t h s r a r e l y e x c e e d e d 60 cm. A l s o i n A l b e r t a , R o l l e y and K e i t h ( 1 9 8 0 ) saw a n o t i c e a b l e s h i f t f r o m open t o more d e n s e l y f o r e s t e d a r e a s i n l a t e w i n t e r , p r o b a b l y t r i g g e r e d by snow a c c u m u l a t i o n . I n g e n e r a l , A l a s k a n moose have used e a r l y s u c c e s s i o n a l d e c i d u o u s a r e a s e x c e p t when deep snow f o r c e d t hem t o s e e k s h e l t e r , u n d e r c o n i f e r o u s f o r e s t c a n o p i e s ( L e R e s c h e e t a l . 1 9 7 4 ) . B a n g s e t a l . ( 1 9 8 4 ) n o t e d heavy use of e a r l y s u c c e s s i o n a l f o r e s t by w i n t e r i n g b u l l moose on t h e K e n a i Refuge i n A l a s k a . I n B r i t i s h C o l u m b i a , Cowan e t a l . ( 1 9 5 0 ) b e l i e v e d t h a t t h e b e s t w i n t e r r a n g e s were e a r l y s u c c e s s i o n a l s t a g e s , b e a r i n g l a r g e q u a n t i t i e s o f d e c i d u o u s b r o w s e , w i t h an a d m i x t u r e o f o l d e r f o r e s t . The o l d e r f o r e s t p r o v i d e s f o r r e t r e a t d u r i n g d e e p s n o w s. E d w a r d s and R i t c e y ( 1 9 5 6 ) f o u n d moose i n open d e c i d u o u s g r o w t h a l l w i n t e r , but snow d e p t h s d i d not ex c e e d 64 5 cm (25 i n ) . E d w a r d s ( 1 9 5 6 ) d o c u m e n t e d i n c r e a s e s i n moose n u m b e r s i n s h a l l o w snow y e a r s and d e c r e a s e s i n d e e p snow y e a r s . T h e s e e f f e c t s can be o f f s e t i n deep snow y e a r s by t h e p r e s e n c e of abundant f o o d and c o v e r on w i n t e r r a n g e s (Edwards 1956). S i l v e r (1976), w o r k i n g a t the Cameron R i v e r R e s e r v e i n n o r t h e a s t e r n B.C., f o u n d t h a t m a t u r e c o n i f e r o u s f o r e s t p r o f o u n d l y r e d u c e d snow d e p t h b u t moose d i d n o t u s e t h e s e a r e a s b e c a u s e snow i n o p e n a r e a s r a r e l y e x c e e d e d 76 cm. E a s t m a n ( 1 9 7 8 ) , w o r k i n g n e a r P r i n c e G e o r g e , f o u n d moose s e l e c t e d c o n i f e r o u s f o r e s t , a s c o m p a r e d w i t h e a r l y s u c c e s s i o n a l s t a g e s , when snow deepened. N e v e r t h e l e s s , t h e r e l a t i o n s h i p b e t w e e n e a r l y s u c c e s -s i o n a l s t a g e s and f o r e s t s i n w i n t e r i s not as s t r a i g h t f o r w a r d as t h e a b o v e r e f e r e n c e s i n d i c a t e . I t may n o t be j u s t snow d e p t h w h i c h d r i v e s moose i n t o heavy c o v e r d u r i n g w i n t e r . Many a u t h o r s d i r e c t l y o r i n d i r e c t l y s u p p o r t t h e h y p o t h e s i s t h a t c o l d s t r e s s i s i m p o r t a n t i n moose s e e k i n g s h e l t e r e d a r e a s i n w i n t e r . T e l f e r ( 1 9 7 8 b ) l i s t e d moose a v o i d a n c e o f w i n d c h i l l as an i m p o r t a n t w i n t e r c o n s i d e r a t i o n . B r u s n y k and G i l b e r t (1983) i n O n t a r i o c i t e d t h e more f a v o u r a b l e t h e r m a l e n v i r o n m e n t i n f o r e s t r e s e r v e s i n w i n t e r f o r t h e i r g r e a t e r u s e by moose. I n d i r e c t s u p p o r t f o r t h e t h e r m a l r e g u l a t i o n h y p o t h e s i s comes f r o m C h a m b e r l i n ( 1 9 7 2 ) who f o u n d moose i n d e n s e f o r e s t when snow d e p t h s i n t h e open a r e a s w e r e l e s s t h a n 76 cm (30 i n ) , 6 e v e n t h o u g h P e t e r s o n ( 1 9 5 5 ) had o b s e r v e d t h a t 76 cm o f snow p r e s e n t e d l i t t l e h i n d e r a n c e t o moose. A d d i s o n e t a l . ( 1 9 8 0 ) s p e c u l a t e d t h a t r a p i d a c c u m u l a t i o n o f snow d u r i n g w i n t e r s t o r m s r a t h e r t h a n snow d e p t h d r i v e s moose t o dense h a b i t a t . T h e r e i s s u p p o r t f o r t h e r m a l s t r e s s a s an i m p o r t a n t f a c t o r i n w i n t e r from o t h e r p a r t s of N o r t h A m e r i c a . P h i l l i p s e t a l . ( 1 9 7 3 ) i n M i n n e s o t a n o t e d w i n t e r moose movement t o h e a v y c o v e r when snow d e p t h s w e r e l e s s t h a n 51 cm (20 i n ) . Van B a l l e n b e r g h e and Peek (1971), a l s o i n M i n n e s o t a , o b s e r v e d moose l e a v i n g heavy c o v e r a f t e r t h e p a s s i n g of w i n t e r s t o r m s . K n o w l t o n (1960) o b s e r v e d t h a t s t o r m s c a u s e d moose i n Montana t o w i t h d r a w t o c o n i f e r o u s t i m b e r b u t t h e y r e t u r n e d t o o p e n w i l l o w h a b i t a t w i t h t h e s t o r m ' s p a s s i n g . R o l l e y and K e i t h ( 1 9 8 0 ) w o r k i n g i n A l b e r t a s t a t e d : " I t i s a l s o p o s s i b l e t h a t c l o s e d c a n o p y u p l a n d s o f f e r moose a more f a v o u r a b l e t h e r m a l e n v i r o n m e n t . " I n B r i t i s h C o l u m b i a , H a t t e r ( 1 9 5 0 ) f e l t t h a t m a t u r e f o r e s t s p r o t e c t e d moose f r o m b o t h snow and w i n d . E a s t m a n (1978) f o u n d moose b e d d i n g w i t h i n i n c r e a s i n g l y p r o t e c t e d a r e a s as w i n t e r p r o g r e s s e d . B e d d i n g s e l e c t i o n would be r e l a t e d t o t h e r m a l p r o t e c t i o n r a t h e r t h a n t o ease of l o c o m o t i o n . R e n e c e k e r e t a l . ( 1 9 7 8 ) p r o v i d e d g u i d e l i n e s as t o what c o n s t i t u t e s c o l d w e a t h e r f o r moose w i t h t h e i r work on moose c a l v e s . T e m p e r a t u r e s o f l e s s t h a n - 2 0 ° w i t h w i n d p r o d u c e d 7 e l e v a t e d m e t a b o l i c r a t e s i n moose c a l v e s i n r e s p o n s e t o c o l d s t r e s s . 1.2 Summer H a b i t a t S e l e c t i o n A n a l y s i s of moose h a b i t a t s e l e c t i o n has u s u a l l y f o c u s e d on t h e w i n t e r s e a s o n . F o r e x a m p l e , T e l f e r ( 1 9 7 8 b ) d i s c u s s e d w i n t e r h a b i t a t r e q u i r e m e n t s a l m o s t e x c l u s i v e l y i n a p a p e r t i t l e d " H a b i t a t r e q u i r e m e n t o f moose - t h e p r i n c i p a l T a i g a r a n g e a n i m a l . " B r a s s a r d e t a l . ( 1 9 7 4 ) s t a t e d , " O n l y w i n t e r h a b i t a t h a s b e e n c o n s i d e r e d , a s l i m i t i n g f a c t o r s a r e most p r e d o m i n a n t d u r i n g t h i s season." O b s e r v a t i o n of and s p e c u l a -t i o n a b o u t summer h a b i t a t s e l e c t i o n h a v e s u f f e r e d f r o m t h e a t t e n t i o n g i v e n w i n t e r h a b i t a t . I n s p i t e o f t h e l a c k o f a t t e n t i o n g i v e n moose summer h a b i t a t , t h e l i t e r a t u r e c o n t a i n s two h y p o t h e s e s t h a t may e x p l a i n summer h a b i t a t s e l e c t i o n . The f i r s t , and most p o p u l a r , h y p o t h e s i s i s t h a t moose summer i n a r e a s w h i c h p r o d u c e l a r g e q u a n t i t i e s of f o r a g e . The c o m p e t i n g h y p o t h e s i s s t a t e s t h a t summer h a b i t a t n e e d s a r e more c o m p l i c a t e d t h a n s i m p l y f o o d and t h a t p r i m a r y among t h e o t h e r needs i s s h e l t e r f r o m heat s t r e s s . T h e r e i s a w i d e s p r e a d p e r c e p t i o n t h a t moose s e l e c t f o r a g e p r o d u c i n g a r e a s i n s u m m e r , p a r t i c u l a r l y e a r l y s u c c e s s i o n a l s t a g e s p r o d u c e d by w i l d f i r e or l o g g i n g . F u r t h e r , 8 mature f o r e s t i s c o n s i d e r e d poor moose summer h a b i t a t b e c a u s e t h e g r o w t h o f f o r a g e p r o d u c i n g p l a n t s i s s u p p r e s s e d by t h e f o r e s t canopy. P e t e r s o n (1955) e m p h a s i z e d t h e i m p o r t a n c e of f i r e and c l e a r c u t t i n g i n p r o d u c i n g summer r a n g e f o r moose. K r e f t i n g ( 1 9 7 4 a ) s t r e s s e d t h e i m p o r t a n c e o f d i s t u r b a n c e t o c r e a t e moose summer r a n g e by r e m o v i n g f o r e s t s and p r o d u c i n g abundant f o o d i n e a r l y s u c c e s s i o n a l s t a g e s . K r e f t i n g (1974b) d i s c u s s e d t h e r o l e o f s p r u c e budworm, b l o w d o w n s , f o r e s t c u t t i n g , and f i r e i n p r o d u c i n g moose r a n g e by r e m o v i n g o v e r s t o r y and s t i m u l a t i n g browse p r o d u c t i o n on I s l e R o y a l . In O n t a r i o u s e o f t i m b e r r e s e r v e s a p p e a r e d t o be g o v e r n e d by s u i t a b l e a q u a t i c f e e d i n g a r e a s and t e r r e s t r i a l b r o w s e i n a d j a c e n t c u t o v e r s ( B r u s n y k and G i l b e r t 1983). S e v e r a l a u t h o r s i n M i n n e s o t a o b s e r v e d t h a t m o o s e s e l e c t e d f o r a g e p r o d u c i n g a r e a s i n summer. P h i l l i p s e t a l . ( 1 9 7 3 ) f o u n d moose i n o p e n w i l l o w ( S a 1_i_x spp.) s t a n d s d u r i n g summer and f a l l . B e r g and P h i l l i p s ( 1 9 7 4 ) o b s e r v e d moose s e l e c t p r e d o m i n a n t l y "low open t y p e s " , of w i l l o w , i n summer. I r w i n ( 1 9 7 5 ) d o c u m e n t e d a r a p i d i n c r e a s e i n summer moose n u m b e r s due t o a f l u s h o f new p a l a t a b l e f o r a g e i n a b u r n e d a r e a . Moose p r e f e r e n c e f o r d e c i d u o u s s h r u b p r o d u c i n g a r e a s i n summer has a l s o b e e n n o t e d i n w e s t e r n N o r t h A m e r i c a . I n M o n t a n a , K n o w l t o n ( 1 9 6 0 ) o b s e r v e d moose f e e d i n g m o s t l y i n s u b a l p i n e meadows and a d j a c e n t w i l l o w b o t t o m s . Dorn (1970), 9 a l s o i n Montana, fo u n d moose p r e d o m i n a n t l y i n w i l l o w s t a n d s i n summer. P i e r c e and Peek ( 1 9 8 4 ) , w o r k i n g i n I d a h o , n o t e d a s i g n i f i c a n t s e l e c t i o n o f open a r e a s , c l e a r c u t s and l a k e s , d u r i n g t h e summer. I n A l a s k a L e R e s c h e e t a l . ( 1 9 7 4 ) n o t e d h i g h summer moose d e n s i t i e s i n h a b i t a t s w i t h abundant f o r a g e s u c h a s b i r c h (B e t u 1 a s.JP_.P_0 , w i l l o w , a n d b o g h a b i t a t s . L e R e s c h e e t a l . ( 1 9 7 4 ) a l s o n o t e d l o w moose d e n s i t i e s i n f o r e s t s . I n c e n t r a l B r i t i s h C o l u m b i a , C o w a n e t a l . ( 1 9 5 0 ) m a i n t a i n e d t h a t moose do n o t o c c u p y c l i m a x f o r e s t i n summer e x c e p t by c o i n c i d e n t o c c u p a t i o n w i t h swamps. B o n a r e t a l . (1975), e x a m i n i n g moose summer use of u n l o g g e d f o r e s t s i n t h e P r i n c e G e o r g e a r e a , c o n c l u d e d t h a t moose u s e o f f o r e s t s c o i n c i d e d w i t h w a t e r c o u r s e s . A l s o i n t h e P r i n c e George a r e a , Eastman (1978) b e l i e v e d moose used p a r t i a l c u t o v e r s and b u r n s i n p r e f e r e n c e t o c o n i f e r o u s f o r e s t i n summer. Many o b s e r v e r s , however, have r e p o r t e d moose i n f o r e s t d u r i n g summer w i t h o u t s t a t i n g or i m p l y i n g t h a t t h e r e were no u s e f u l r e s o u r c e s i n t h e f o r e s t s t r u c t u r e . Dodds (1974) n o t e d t h a t i n t h e n o r t h e a s t e r n r e g i o n of N o r t h A m e r i c a , moose occupy f o r e s t i n summer. Peek et a l . (1976) o b s e r v e d a summer s h i f t o f M i n n e s o t a moose i n t o h a b i t a t c h a r a c t e r i z e d by g r e a t e r c o v e r . K e a r n e y and G i l b e r t (1976) f o u n d more moose l o c a t i o n s t h a n e x p e c t e d a t r a n d o m i n f o r e s t i n O n t a r i o d u r i n g summer. 10 Hauge and K e i t h ( 1 9 8 1 ) , i n A l b e r t a , f o u n d moose i n f o r e s t i n summer. A l s o i n A l b e r t a , M y t t o n and K e i t h ( 1 9 8 1 ) f o u n d moose i n t a l l dense d e c i d u o u s f o r e s t s i n summer. Bangs e t a l . (1984) documented heavy summer use of f o r e s t by A l a s k a n B u l l moose. I n B r i t i s h C o l u m b i a , E d w a r d s and R i t c e y ( 1 9 5 6 ) n o t e d t h a t moose were i n s u b a l p i n e f o r e s t i n summer. K n o r r e ( 1 9 5 9 ) p r o v i d e d t h e b a s i s f o r an e x p l a n a t i o n o f moose u s e o f f o r e s t i n summer. K n o r r e ( 1 9 5 9 ) o b s e r v e d t h a t moose s u f f e r s e v e r e l y f r o m h e a t . They r e q u i r e d shade or wa t e r t o l i e i n on summer d a y s w i t h a i r t e m p e r a t u r e s g r e a t e r t h a n 20°C. F u r t h e r , moose d r a s t i c a l l y a l t e r e d t h e i r d a i l y p a t t e r n s d u r i n g h o t w e a t h e r and w e r e i n f a r b e t t e r c o n d i t i o n a f t e r a c o o l summer as c o m p a r e d w i t h a h o t summer. S i x month o l d c a l v e s a v e r a g e d 146 kg a f t e r a c o o l summer but 101 kg a f t e r a h o t summer ( K n o r r e 1 9 5 9 ) . K e l s a l l and T e l f e r ( 1 9 7 4 ) a l s o o b s e r v e d moose s e v e r e l y s t r e s s e d by summer heat and p r o p o s e d t h a t h e a t and t h e a b s e n c e o f s h a d e w e r e i m p o r t a n t f a c t o r s i n l i m i t i n g m o o s e d i s t r i b u t i o n i n w e s t e r n N o r t h A m e r i c a . B e l o v s k y and J o r d a n (1978) s u g g e s t e d t h a t t h e r m a l c o n d i t i o n s l i m i t moose a c t i v i t y on t h e b a s i s o f a n e g a t i v e c o r r e l a t i o n ( r = 0.92) b e t w e e n t i m e s p e n t f e e d i n g and a i r t e m p e r a t u r e . B e l o v s k y ( 1 9 8 1 ) s u c c e s s f u l l y p r e d i c t e d t h a t i n summer moose s h o u l d use c o n i f e r o u s f o r e s t f o r a l l of t h e i r r e s t i n g and f o r a l a r g e p a r t o f t h e i r f o r a g i n g b a s e d on t h e a s s u m p t i o n o f t h e r m o n e u t r a l i t y . 11 2. Summary and H y p o t h e s e s The f o r e g o i n g r e v i e w of moose w i n t e r and summer h a b i t a t s e l e c t i o n p r o d u c e s two g e n e r a l o b s e r v a t i o n s on t h e f a c t o r s w h i c h a f f e c t moose h a b i t a t s e l e c t i o n : t h e f i r s t i s t h a t moose a r e f o u n d i n a r e a s w h i c h p r o d u c e l a r g e amounts of f o r a g e . The s e c o n d i s t h a t moose r e q u i r e s h e l t e r f r o m t h e r m a l s t r e s s . T h e s e o b s e r v a t i o n s a r e c o n t r a d i c t o r y b e c a u s e t h e g r e a t e s t p r o d u c t i o n of f o r a g e i s e x p e c t e d i n open a r e a s whereas s h e l t e r f r o m t h e r m a l s t r e s s w i l l be f o u n d i n a r e a s o f h e a v y f o r e s t c o v e r . The i m p o r t a n c e o f t h i s c o n t r a d i c t i o n t o l o g g i n g i s t h a t one o b s e r v a t i o n s u g g e s t s t h a t l o g g i n g p r o d u c e s moose h a b i t a t by c r e a t i n g e a r l y s u c c e s s i o n a l s t a n d s o f d e c i d u o u s s h r u b s w h i l e t h e o t h e r a n t i c i p a t e s t h e d e s t r u c t i o n o f moose h a b i t a t t h r o u g h t h e r e m o v a l o f t h e m o d e r a t e t h e r m a l e n v i r o n m e n t due t o mature t r e e s . T h e s e g e n e r a l o b s e r v a t i o n s c a n be s t a t e d a s f o r m a l h y p o t h e s e s . F o r t h e w i n t e r p e r i o d t h r e e outcomes a r e p o s s i b l e : (1) Moose a r e r a n d o m l y d i s t r i b u t e d w i t h r e s p e c t t o h a b i t a t t y p e s . (2) Moose a r e n o t r a n d o m l y d i s t r i b u t e d b u t s e l e c t t h e h a b i t a t w i t h t h e most f o r a g e a v a i l a b l e e x c e p t when snow d e p t h e x c e e d s 90 cm. I f snow d e p t h e x c e e d s 90 cm, t h e n moose w i l l s e l e c t h a b i t a t s w i t h g r e a t e s t f o r e s t c o v e r . 12 (3) Moose a r e n o t r a n d o m l y d i s t r i b u t e d b u t s e l e c t t h e h a b i t a t w i t h t h e g r e a t e s t f o r e s t c o v e r as p r o t e c t i o n f r o m c o l d s t r e s s , when t e m p e r a t u r e s a r e l e s s t h a n -20° T h e r e a r e t h r e e p a r a l l e l h y p o t h e s e s a p p l i c a b l e d u r i n g the summer ( n o n w i n t e r ) p e r i o d : (1) Moose a r e r a n d o m l y d i s t r i b u t e d w i t h r e s p e c t t o h a b i t a t t y p e . (2) Moose a r e n o t r a n d o m l y d i s t r i b u t e d b u t s e l e c t t h e h a b i t a t w i t h t h e most a v a i l a b l e f o r a g e . (3) Moose a r e n o t r a n d o m l y d i s t r i b u t e d b u t w i l l s e l e c t h a b i t a t s w i t h g r e a t e s t f o r e s t c o v e r w h e n a i r t e m p e r a t u r e s e x c e e d 20°C. 13 I I STUDY AREA 1. Geography The s t u d y a r e a i s an i r r e g u l a r p o l y g o n n o r t h e a s t o f P r i n c e George, B r i t i s h C o l u m b i a ( F i g u r e 1 and F i g u r e 2). The e x t r e m e n o r t h e r l y p o i n t i s 54° 27' N o r t h L a t i t u d e , t h e e x t r e m e e a s t e r l y p o i n t i s 1 2 1 ° 44' West L o n g i t u d e , t h e e x t r e m e s o u t h e r l y p o i n t i s 5 4 ° 07' N o r t h L a t i t u d e , and t h e e x t r e m e w e s t e r l y p o i n t i s 122° 37' West L o n g i t u d e . The s t u d y a r e a was d e f i n e d by t h e l o c a t i o n s of r a d i o c o l l a r e d moose, but c o n f o r m s where p o s s i b l e t o t h e a d m i n i s t r a t i v e b o u n d a r i e s of Northwood P u l p and T i m b e r ' s TFL 30. Moose w i n t e r range was d e f i n e d as a b r o a d swath o v e r t h e a r e a o c c u p i e d by moose i n m i d w i n t e r and l a t e w i n t e r . The l a n d f o r m s i n c l u d e t h e M c G r e g o r P l a t e a u i n t h e n o r t h e a s t , t h e F r a s e r B a s i n i n t h e n o r t h w e s t , and t h e Nechako P l a i n i n t h e s o u t h ( H o l l a n d 1 9 7 6 ) . The S u b b o r e a l S p r u c e b i o g e o c l i m a t i c z o n e c o v e r s t h e e n t i r e s t u d y a r e a ( B r i t i s h C o l u m b i a M i n i s t r y o f F o r e s t s 1980). M a j o r f e a t u r e s o f t h e s t u d y a r e a i n c l u d e : t h e F r a s e r R i v e r , a t a p p r o x i m a t e l y 600 m i n e l e v a t i o n , f l o w s n o r t h w e s t t h e n s o u t h t h r o u g h t h e s o u t h e r l y p o r t i o n s o f t h e s t u d y a r e a . A number o f r i d g e s r u n n o r t h w e s t t o s o u t h e a s t r a n g i n g i n e l e v a t i o n f r o m 990 m i n t h e s o u t h w e s t t o 1490 m i n t h e 14 gure 1. L o c a t i o n o f t h e s t u d y a r e a w i t h i n B r i t i s h C o l u m b i a 16 F i g u r e 2. B o u n d a r i e s o f t h e s t u d y a r e a , l o c a t i o n s o f p l o t s , and t h e r m o g r a p h s on a 1:250,000 s c a l e map The s o l i d l i n e r e p r e s e n t s t h e s p r i n g t o e a r l y w i n t e r boundary and t h e dashed l i n e r e p r e s e n t s t h e m i d w i n t e r and l a t e w i n t e r b o u n d a r y . The m i d w i n t e r and l a t e w i n t e r boundary i s o m i t t e d where i t i s c o i n c i d e n t w i t h t h e s p r i n g t o e a r l y w i n t e r b o u n d a r y . P l o t s on w h i c h f o r a g e p r o d u c t i o n and f o r e s t c o v e r were r e c o r d e d a r e r e p r e s e n t e d by u p p e r c a s e l e t t e r s a s f o l l o w s : F u n l o g g e d f o r e s t , D d i a m e t e r l i m i t l o g g i n g , I i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g , 0 o l d c l e a r c u t , R r e c e n t c l e a r c u t , U u p l a n d s h r u b , S r i p a r i a n s h r u b , and W w e t l a n d . A b a r a b o v e a h a b i t a t d e s i g n a t i o n i n d i c a t e s a p l o t on w h i c h a s n o w c o u r s e was c o n s t r u c t e d . W h e r e t h e h a b i t a t d e s i g n a t i o n i s p r e c e d e d by a T, a t h e r m o g r a p h was p r e s e n t i n t h e h a b i t a t t y p e i n d i c a t e d . 18 n o r t h e a s t , s e p a r a t e d by v a l l e y s w i t h f l o o r s a t 630 m i n t h e s o u t h w e s t t o 850 m i n t h e n o r t h e a s t . 2. L o g g i n g H i s t o r y L o g g i n g i n t h e s t u d y a r e a h a s u n d e r g o n e t h r e e p h a s e s : d i a m e t e r l i m i t l o g g i n g , i n t e r m e d i a t e u t i l i z a t i o n , a n d c 1 e a r c u t t i n g . I n d i a m e t e r l i m i t l o g g i n g , t r e e s w i t h a d i a m e t e r a t b r e a s t h e i g h t (DBH) o f g r e a t e r t h a n 30.5 cm (12 i n ) w e r e r e m o v e d . I n t e r m e d i a t e u t i l i z a t i o n f e a t u r e d t h e r e m o v a l of a l l c o m m e r c i a l l y v a l u a b l e t r e e s l e a v i n g noncommer-c i a l s p e c i e s , u s u a l l y At>i_e£3, o r c l e a r i n g s m a l l p a t c h e s o f a b o u t 4 ha and r e m o v i n g o n l y c o m m e r c i a l t r e e s f r o m a d j a c e n t s m a l l p a t c h e s . In c l e a r c u t t i n g a l l t r e e s a r e removed and t h e c u t may be s c a r i f i e d or burned. T h r e e t r e n d s a r e e v i d e n t t h r o u g h t i m e . (1) L o g g i n g has become more i n t e n s e . D i a m e t e r l i m i t l o g g i n g b e g a n i n t h e e a r l y 1950s and e n d e d i n 1959. I n t e r m e d i a t e u t i l i z a t i o n l o g g i n g was u s e d i n t h e 1 9 6 0 s . C l e a r c u t t i n g h a s b e e n t h e m e t h o d u s e d i n t h e 1970s and 1980s. (2) L o g g i n g h a s s p r e a d f r o m l o w e l e v a t i o n (<760 m) i n t h e 1950s up t o 1460 m i n t h e 1 9 7 0 s . ( 3 ) L o g g i n g h a s s p r e a d n o r t h a n d e a s t f r o m t h e v i c i n i t y of t h e F r a s e r R i v e r . 19 I I I METHODS To t e s t t h e h y p o t h e s e s s t a t e d i n t h e I n t r o d u c t i o n d a t a i n f i v e c a t e g o r i e s were c o l l e c t e d : (1) t o d e t e r m i n e w hether moose were s e l e c t i n g h a b i t a t s , and w h i c h h a b i t a t s were s e l e c t -ed, (2) t o e s t a b l i s h c r e d i b l e e s t i m a t e s o f s e a s o n a l f o r a g e p r o d u c t i o n , (3) t o e s t i m a t e t h e f o r e s t c o v e r on each h a b i t a t , (4) t o e s t i m a t e t h e snow d e p t h on e a c h h a b i t a t d u r i n g t h e w i n t e r , (5) t o e s t i m a t e t h e a i r t e m p e r a t u r e i n e a c h h a b i t a t d u r i n g a l l s e a s o n s . F o l l o w i n g i s a d e s c r i p t i o n o f t h e method used t o e s t i m a t e each o f t h e s e f a c t o r s . 1. H a b i t a t S e l e c t i o n 1.1 H a b i t a t Maps The s t u d y a r e a was s t r a t i f i e d i n t o 10 h a b i t a t t y p e s , 5 f o r e s t / l o g g e d h a b i t a t s and 5 n o n f o r e s t h a b i t a t s . The f o r e s t / l o g g e d h a b i t a t s i n c l u d e d : (1) r e c e n t c l e a r c u t s l o g g e d l a t e r t h a n summer 1973; (2) o l d c l e a r c u t s l o g g e d d u r i n g t h e summer o f 1973 o r e a r l i e r ; (3) i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g c o n d u c t e d d u r i n g t h e 1960s when a l l c o m m e r c i a l l y d e s i r a b l e t r e e s w e r e r e m o v e d and u n d e s i r a b l e t r e e s l e f t g r o w i n g , o r c o m m e r c i a l l y d e s i r a b l e t r e e s w e r e r e m o v e d f r o m 4 ha p a t c h e s and a d j a c e n t s m a l l p a t c h e s were c l e a r c u t ; (4) d i a m e t e r l i m i t l o g g i n g i n w h i c h a l l c o m m e r c i a l l y v a l u a b l e t r e e s w i t h a dbh 20 g r e a t e r t h a n 30 cm were h a r v e s t e d i n the 1950s; and (5) u n l o g -ged f o r e s t . The n o n f o r e s t h a b i t a t t y p e s i n c l u d e : (6) u p l a n d s h r u b , a l d e r (Aljiu^s v.irj^d_u^); (7) r i p a r i a n s h r u b ; (8) w e t -l a n d s ; (9) t h e F r a s e r R i v e r ; and (10) l a k e s . A h a b i t a t map was made on a 1:50,000 f o r e s t c o v e r map w i t h l o g g i n g b o u n d a r i e s . The 1:50,000 map was c o l o u r e d w i t h r e f e r e n c e t o 1:10,000 s c a l e f o r e s t c o v e r maps, 1:10,000 s c a l e l o g g i n g h i s t o r y maps, and 1:50,000 s c a l e a e r i a l p h o t o g r a p h s . The t o t a l a r e a o f t h e s t u d y s i t e e q u a l l e d 142,054.8 ha, as d e t e r m i n e d by r e d u c i n g t h e mapped a r e a t o s i m p l e r g e o m e t r i c s h a p e s , c a l c u l a t i n g t h e a r e a o f t h e s m a l l u n i t s , and a d d i n g t h e a r e a o f t h e s m a l l u n i t s t o g e t h e r . The c o n t r i b u t i o n o f each h a b i t a t t o the t o t a l a r e a was d e t e r m i n e d by c o u n t i n g t h e number o f d o t s on e a c h h a b i t a t u s i n g a s y s t e m a t i c d o t g r i d . T h e r e w e r e 1,333 r e c o r d s f o r e a c h p a s s o f t h e d o t g r i d . The p r o c e d u r e was r e p e a t e d u n t i l t h e b i n o m i a l v a r i a n c e of u n l o g g e d f o r e s t , t h e h a b i t a t w i t h t h e g r e a t e s t v a r i a n c e b e c a u s e i t was c l o s e s t t o 0.5 o f t h e t o t a l a r e a , w o u l d n o t be d e c r e a s e d by a f u r t h e r p a s s o f t h e g r i d . T h e b i n o m i a l v a r i a n c e was c a l c u l a t e d a c c o r d i n g t o M e n d e n h a l l (1967, p. 163). The a r e a of each h a b i t a t and p e r c e n t a g e of t h e t o t a l a r e a a r e i n T a b l e 1. H a b i t a t a r e a s and p e r c e n t a g e o f t h e t o t a l a r e a were a l s o d e t e r m i n e d f o r t h e s m a l l e r m i d w i n t e r and l a t e w i n t e r r a n g e s . The r e s u l t s a r e p r e s e n t e d i n T a b l e 1. 21 T a b l e 1. A r e a o c c u p i e d by each h a b i t a t t y p e on s p r i n g t h r o u g h e a r l y w i n t e r and mid- t o l a t e w i n t e r moose r a n g e s . S p r i n g t o Mid t o E a r l y W i n t e r L a t e W i n t e r Range Range H a b i t a t A r e a P e r c e n t a g e A r e a P e r c e n t a g e u n l o g g e d f o r e s t 78,017 ha 54. .9% 17,763 ha 34, .4% d i a m e t e r l i m i t 10,441 7. .4 7, 177 13. .9 i n t e r m e d i a t e 14,958 10, .5 12,083 23. .4 u t i l i z a t i o n o l d c l e a r c u t 5, 185 3, .7 3, 253 6, .3 r e c e n t c l e a r c u t 10,313 7. .3 4,905 9. ,5 u p l a n d s h r u b 8,438 5. ,9 1,446 2. ,8 r i p a r i a n s h r u b 4,816 3. ,4 1,291 2. ,5 w e t l a n d s 7,841 5. ,5 2,324 4. .5 F r a s e r R i v e r 1 ,122 0. ,8 981 1. ,9 l a k e s 909 0. .6 413 0. .7 T o t a l 142,055 ha 100.0% 51,636 ha 99.9% 22 1.2 Moose L o c a t i o n s F o r t y moose were c h e m i c a l l y i m m o b i l i z e d and f i t t e d w i t h r a d i o t r a n s m i t t e r s d u r i n g t h r e e i m m o b i l i z a t i o n s e s s i o n s : 8 moose, 24 and 25 March 1980; 12 moose, 30 and 31 J a n u a r y 1981; and 20 moose, 27 J a n u a r y t h r o u g h 13 F e b r u a r y 1982 ( S c h w a b e t a l . 1 9 8 4 ) . T h e s e m o o s e p r o v i d e d 1,265 r a d i o t e l e m e t r y l o c a t i o n s . T w e n t y - f o u r moose p r o v i d e d a l m o s t a l l o f t h e l o c a t i o n s b e c a u s e of d e a t h s and t r a n s m i t t e r f a i l u r e . F o u r of t h e s e 24 moose were male. R a d i o t e l e m e t r y l o c a t i o n began on 8 A p r i l 1980 and ended 31 May 1983 w i t h a b r e a k b e t w e e n 12 D e c e m b e r 1980 and 2 May 1981. From s p r i n g t o e a r l y w i n t e r w e e k l y l o c a t i o n s were made u s i n g a C e s s n a 172 a i r c r a f t w i t h a 4 e l e m e n t Y a g i a n t e n n a on e a c h w i n g s t r u t . T h e a i r c r a f t was s t e e r e d t o w a r d t h e s t r o n g e s t s i g n a l u s i n g a l e f t - r i g h t s w i t c h box a t t a c h e d b e t w e e n t h e r e c e i v e r and t h e a n t e n n a s . C o n f i r m a t i o n o f a t e n t a t i v e l o c a t i o n was made by f l y i n g t o w a r d t h e t e n t a t i v e l o c a t i o n f r o m a n u m b e r o f a n g l e s . V i s u a l c o n t a c t was f r e q u e n t l y made w i t h a n i m a l s i n open h a b i t a t and a l m o s t n e v e r made w i t h a n i m a l s i n dense h a b i t a t . From 15 J a n u a r y u n t i l t h e i n i t i a t i o n of s p r i n g m i g r a t i o n a e r i a l l o c a t i o n s w e r e s u p p l e m e n t e d w i t h g r o u n d l o c a t i o n s , u s i n g a hand h e l d 3 e l e m e n t Y a g i a n t e n n a . G r o u n d l o c a t i o n s w e r e e s t a b l i s h e d by p l o t t i n g t h e d i r e c t i o n s o f t r a n s m i t t e r s 23 f r o m a number o f t e l e m e t r y s t a t i o n s . G r o u n d l o c a t i o n s w e r e ea s y t o e s t a b l i s h b e c a u s e a l m o s t a l l moose w i n t e r e d w i t h i n 1 km of a f o r e s t h a u l r o a d and l o c a t i o n s c o u l d be compared w i t h c o n c u r r e n t a e r i a l t e l e m e t r y . L o c a t i o n s were made t w i c e w e e k l y w h i l e moose were on t h e i r w i n t e r range. L o c a t i o n s were p l o t t e d on a 1:50,000 t o p o g r a p h i c map and s u b m i t t e d t o H a r e s t a d ' s (1981) "Home" program g i v i n g t h e a n g l e and d i s t a n c e f r o m r e f e r e n c e p o i n t s . C o m p u t e r p l o t s on a 1:50,000 s c a l e were made c o n s i s t i n g of r e f e r e n c e p o i n t s and a c i r c l e r e p r e s e n t i n g an a r e a 200 m i n d i a m e t e r f o r e a c h l o c a t i o n . L y n c h ( 1 9 7 9 ) s u g g e s t e d t h a t a t r a n s m i t t e r was w i t h i n a c i r c l e w i t h a r a d i u s o f 100 m o f a map l o c a t i o n f o r a e r i a l t e l e m e t r y . The 3.142 ha c i r c l e w i t h a r a d i u s 100 m i s a b o u t h a l f t h e d i f f e r e n c e b e t w e e n t h e m i n i m u m and maximum e r r o r p o l y g o n s f o r t h e a e r i a l l o c a t i o n s of moose d e t e r m i n e d by P i e r c e and Peek ( 1 9 8 4 ) . V i s u a l c o n t a c t w i t h c o l l a r e d moose s u g g e s t e d t h a t t h e 200 m d i a m e t e r c i r c l e was a c o n s e r v a t i v e e s t i m a t e o f e r r o r . G r e a t e r p r e c i s i o n t h a n e x p e c t e d i n t h e l o c a t i o n s o f moose was c o m p e n s a t e d f o r by e r r o r s i n m a p p i n g l o c a t i o n s . G u i d e d by r e f e r e n c e p o i n t s , c o m p u t e r p l o t s w e r e o v e r l a i d on t h e h a b i t a t map. Any h a b i t a t t y p e w i t h i n a l o c a t i o n c i r c l e was s c o r e d an e q u a l p a r t of t h e l o c a t i o n . F o r e x a m p l e , i n t h e c a s e o f two h a b i t a t s w i t h i n t h e c i r c l e , e a c h s c o r e d h a l f a l o c a t i o n and i n t h e c a s e o f t h r e e h a b i t a t s w i t h i n t h e c i r c l e , e a c h s c o r e d o n e - t h i r d o f a l o c a t i o n . A l l 24 h a b i t a t s w i t h i n a l o c a t i o n a r e a were g i v e n e q u a l w e i g h t b e c a u s e a l l h a b i t a t i n t h e v i c i n i t y o f a moose may h a v e a f f e c t e d t h e a n i m a l ' s d e c i s i o n t o occupy t h e g e n e r a l a r e a . 1.3 S t a t i s t i c a l A n a l y s i s S e a s o n s w e r e d e f i n e d a s f o l l o w s : (1) s p r i n g , f r o m t h e o n s e t o f s p r i n g m i g r a t i o n u n t i l 15 J u n e ; (2) summer, f r o m 16 J u n e t o 15 S e p t e m b e r ; (3) f a l l , f r o m 16 S e p t e m b e r u n t i l t h e o n s e t of f a l l m i g r a t i o n ; (4) e a r l y w i n t e r , f r o m f a l l m i g r a t i o n u n t i l 15 J a n u a r y ; (5) m i d w i n t e r , f r o m 16 J a n u a r y u n t i l 15 M a r c h ; and (6) l a t e w i n t e r , f r o m 16 M a r c h u n t i l t h e o n s e t o f s p r i n g m i g r a t i o n . The G s t a t i s t i c , i n c o r p o r a t i n g t h e W i l l i a m s C o r r e c t i o n F a c t o r f o r d i s c o n t i n u i t y , i s t h e p r e f e r r e d t e s t o f w h e t h e r an a n i m a l i s d i s t r i b u t e d r a n d o m l y w i t h r e s p e c t t o t h e a v a i l a b i l i t y o f h a b i t a t t y p e s ; when t h e n u l l h y p o t h e s i s i s t r u e t h e G C Q r s t a t i s t i c i s a b e t t e r a p p r o x i m a t i o n of t h e C h i -s q u a r e d i s t r i b u t i o n t h a n x ( S o k a l and R o h l f 1981). To u s e G C Q r , t h e e x p e c t e d f r e q u e n c y of o b s e r v a t i o n s i n each c a t e g o r y must be a t l e a s t 5 ( S o k a l and R o h l f 1981). The x 2 t e s t c a n f u n c t i o n p r o p e r l y when t h e e x p e c t e d f r e q u e n c y i n each c a t e g o r y i s a s l o w as 1 ( S n e d e c o r and C o c h r a n 1980). S i n c e t h e r u l e f o r t h e a p p l i c a t i o n o f t h e G s t a t i s t i c was o f t e n v i o l a t e d by 25 t h e d a t a and t h e r u l e f o r t h e a p p l i c a t i o n o f x w a s n e v e r 9 v i o l a t e d , X was c h o s e n t o t e s t w h e t h e r moose d i s t r i b u t e d t h e m s e l v e s r a n d o m l y w i t h r e s p e c t t o t h e a v a i l a b i l i t y o f h a b i t a t t y p e s . The s i g n i f i c a n t l e v e l was s e t a t 0.1 f o r t h e X t e s t . M o o se d i s t r i b u t i o n was t e s t e d o v e r 6 c a t e g o r i e s : f i v e f o r e s t / l o g g e d h a b i t a t s p l u s n o n f o r e s t d a t a lumped i n t o a s i n g l e c a t e g o r y . I f the d i s t r i b u t i o n of moose was nonrandom w i t h r e s p e c t t o h a b i t a t , t h e h a b i t a t t y p e s r e s p o n s i b l e f o r n o n r a n d o m n e s s w e r e i d e n t i f i e d w i t h an a d a p t a t i o n o f t h e C o l e ( 1 9 4 9 ) i n d e x (Cy) o f a s s o c i a t i o n ( S i n c l a i r 1977). U s i n g a 2 by 2 c o n t i n g e n c y t a b l e ( T a b l e 2 ) , t h e C o l e i n d e x compares t h e number of l o c a t i o n s i n w h i c h moose and t h e h a b i t a t t y p e w e r e p r e s e n t ( a ) , t h e number o f l o c a t i o n s i n w h i c h moose were p r e s e n t and t h e h a b i t a t t y p e was a b s e n t ( b ) , t h e number o f l o c a t i o n s i n w h i c h moose w e r e a b s e n t and t h e h a b i t a t t y p e was p r e s e n t ( c ) , and t h e number o f l o c a t i o n s i n w h i c h n e i t h e r moose n o r t h e h a b i t a t t y p e w e r e p r e s e n t ( d ) . The number of p o s s i b l e l o c a t i o n s i n each h a b i t a t t y p e e q u a l l e d t h e a r e a o f t h e h a b i t a t t y p e d i v i d e d by 3.142 ha, i . e . t h e a r e a of c i r c l e 200 m i n d i a m e t e r . When ad >bc, (1) C y = ( a d - b c ) / [ ( a + d ) ( b + d ) ] . 26 T a b l e 2 . C o l e ( C y ) i n d e x o f a s s o c i a t i o n H a b i t a t A n i m a l S p e c i e s p r e s e n t a b s e n t p r e s e n t a b a b s e n t c d When a d > b e , Cy = ( a d - b c ) / [ ( a + d ) ( b + d ) ] + c c , a\ = ( a + c ) ( c + d ) / [ n ( a + b ) ( b + d ) ] . When be > a d a n d d ^ a , Cy = ( a d - b c ) / [ ( a + b ) ( a + c ) ] + aQ, a\ = ( b + d ) ( c + d ) / [ n ( a + b ) ( a + c ) ] . I n a l l c a s e s , n = a + b + c + d . 27 When be > a d a n d d _> a , ( 2 ) C y = ( a d - b c ) / [ ( a + b ) ( a + c ) ] . T h e r e i s a t h i r d e q u a t i o n a p p r o p r i a t e w h e n be > a d a n d a > d , b u t t h a t s i t u a t i o n n e v e r o c c u r r e d . T h e v a l u e o f t h e C o l e i n d e x r a n g e s f r o m - 1 . 0 t o + 1 . 0 . I f t h e n u m b e r o f l o c a t i o n s e q u a l s t h e n u m b e r e x p e c t e d a t r a n d o m , t h e n a v a l u e o f 0 . 0 i s r e c o r d e d . A s i g n i f i c a n t a s s o c i a t i o n i s f a r t h e r t h a n o n e s t a n d a r d e r r o r f r o m a n a s s o c i a t i o n o f 0 . 0 . When a d > b e , ( 3 ) 02c = ( a + c ) ( c + d ) / [ n ( a + b ) ( b + d ) ] . When be > a d a n d d >^  a , ( 4 ) o 2 . = ( b + d ) ( c + d ) / [ n ( a + b ) ( a + c ) ] . F o r e q u a t i o n s ( 3 ) a n d ( 4 ) , n i s a + b + c + d . P a r a m e t r i c c o r r e l a t i o n s w e r e u s e d t o d e s c r i b e t h e r e l a t i o n s h i p b e t w e e n C o l e i n d i c e s a n d e s t i m a t e s o f a v a i l a b l e f o r a g e a n d b e t w e e n C o l e i n d i c e s a n d e s t i m a t e s o f f o r e s t c o v e r . 28 2. F o r a g e Biomass P r o d u c t i o n T h e r e w e r e f i v e f o r a g e g r o u p s r e c o g n i z e d f o r t h e p u r p o s e s o f t h i s t h e s i s : t h e l e a v e s and t w i g s o f p a l a t a b l e s h r u b s and f o u r g r o u p s i n t h e h e r b l a y e r . The f o u r g r o u p s i n t h e h e r b l a y e r w e r e f e r n s and E^udjBeJtum spp., f o r b s , g r a s s , and s e d g e s and r u s h e s . The f o l l o w i n g d e s c r i b e s how d r y w e i g h t s were e s t i m a t e d f o r t h e p r o d u c t i o n i n each f o r a g e group and t o t a l f o r a g e a v a i l a b l e f o r e a c h s e a s o n . E s t i m a t e s o f v a r i a n c e o f e a c h g r o u p and o v e r a l l s e a s o n a l e s t i m a t e s o f v a r i a n c e a r e a l s o i n c l u d e d . 2.1 Shrub Biomass P r o d u c t i o n The f o l l o w i n g s e c t i o n s d e s c r i b e t h e t h r e e s t e p s i n v o l v e d i n e s t i m a t i n g t h e b i o m a s s o f s h r u b s . F i r s t , r e g r e s s i o n e q u a t i o n s , e s t i m a t i n g t h e p r o d u c t i o n of i n d i v i d u a l s t e ms, were e s t a b l i s h e d . Second, stem d e n s i t i e s were e s t i m a t e d c o n c u r r e n t w i t h t h e m e a s u r e m e n t o f i n d e p e n d e n t v a r i a b l e s f o r t h e r e g r e s s i o n e s t i m a t e s . T h i r d , t h e mean p r o d u c t i o n per stem was m u l t i p l i e d by mean stem d e n s i t y t o p r o d u c e a b i o m a s s e s t i m a t e per u n i t a r e a f o r a h a b i t a t t y p e . A f o u r t h s e c t i o n d e a l s w i t h t h e e s t i m a t i o n of v a r i a n c e f o r sh r u b b i o m a s s e s t i m a t e s . 29 2.1.1 R e g r e s s i o n e q u a t i o n s S a m p l e s o f s h r u b s o f 12 g r o u p s w e r e t a k e n b e t w e e n 21 J u l y and 13 A u g u s t 1981: A£i£JL JL £ £ i £ £ a r _p_a , A c e r S l a b r u m , A m e l a n c h i e r a l n i f o l i a , B e t u l a p a p y r i f e r a , B e t u l a g l a n d u l o s a , Co.J_n£££!.££.§_, Po_p__u^IJbja^s^mi_f_e_ra_, P o ^ u l u s _tre_m_u3L£iLd_e_s^, S.£ii2£ spp., S a i b u c u s Za.c_£i!l£££» S£j__b_u£ spp., and V.!££££££m e d u l e . S a m p l e s o f an a d d i t i o n a l f i v e g r o u p s w e r e t a k e n between 21 J u l y and 7 August 1982: C o r y l u s c o r n u t a , L o n i c e r a djivo 1 ucr_aJta , R.£s.<a spp., S__p_i_jrae_a spp., and Va^cjiiii^_u m spp. ( S c i e n t i f i c names of p l a n t s f o l l o w T a y l o r and MacBryde, 1977.) F o r e a c h s h r u b , s t e m d i a m e t e r s w e r e s t r a t i f i e d i n t o c l a s s e s s p a n n i n g 5 mm: 5-10 mm, 10-15 mm, 15-20 mm, and c o n t i n u i n g i n t h i s manner t o as l a r g e as d i a m e t e r s had b e e n r e c o r d e d f o r t h a t s p e c i e s d u r i n g p r e l i m i n a r y s a m p l i n g . Whole s h r u b s were s a m p l e d f r o m t h e r a n g e of h a b i t a t t y p e s o c c u p i e d by a s p e c i e s . A number of s h r u b s i n each stem d i a m e t e r c l a s s was a s s i g n e d t o each h a b i t a t . S amples were c l u s t e r e d a t t h e l o w e r and upper ends of t h e r a n g e of stem d i a m e t e r s o c c u p i e d by a p a r t i c u l a r g r o u p , e x c e p t f o r A b_ j i e__s J L £ £ i £ £ £ J__p_a_, A_c_e__r S l ^ I i E » a n d P o_p__u ^ ujs b a l s a m i f e r a i n w h i c h o n l y a few s t e m s w i t h v e r y l a r g e d i a m e t e r s w e r e s a m p l e d . R e p r e s e n t a t i v e s a m p l e s w e r e o b t a i n e d by s e l e c t i n g f r o m t h e w h o l e r a n g e o f v i g o u r , p a r t i c u l a r l y w i t h r e s p e c t t o t h e e f f e c t s of b r o w s i n g , d i s p l a y e d by a s p e c i e s on a s i t e . 30 M e a s u r e m e n t s t a k e n on s i t e i n c l u d e d stem d i a m e t e r 2.5 cm a b o v e g r o u n d , maximum h e i g h t o f f o l i a g e , d e p t h o f f o l i a g e , maximum f o l i a g e d i a m e t e r , and t h e d i a m e t e r p e r p e n d i c u l a r t o maximum f o l i a g e d i a m e t e r . The d i a m e t e r o f t h e stem was t a k e n a b o v e any u n u s u a l s w e l l i n g a t t h e b a s e o f t h e s t e m o r 2.5 cm fr o m t h e o l d g r o w t h i n t h e c a s e of a s u c k e r . The c o n d i t i o n of e a c h s h r u b was s u b j e c t i v e l y r e c o r d e d a s one o f u n b r o w s e d , browsed, or h e a v i l y browsed. Unbrowsed p l a n t s had no browsed t w i g s , e x c e p t t h a t up t o t h r e e browsed p o i n t s were t o l e r a t e d on p l a n t s j u d g e d l a r g e e n o u g h so as t o be u n a f f e c t e d by s u c h b r o w s i n g . A browsed p l a n t had m u l t i p l e browse p o i n t s , but t h e g r o w t h f o r m was u n a f f e c t e d . A h e a v i l y browsed p l a n t was one w h e r e b r o w s i n g had a f f e c t e d t h e g r o w t h f o r m o f t h e s h r u b , p r o d u c i n g a h e d g e d a p p e a r a n c e i n d e c i d u o u s p l a n t s and an h o u r g l a s s g r o w t h f o r m i n A b i e s l a s i o c a r p a . S a m p l e d s h r u b s were t a k e n t o t h e O l s s o n Camp f i e l d s t a t i o n w h e r e c u r r e n t a n n u a l g r o w t h o f t w i g s and l e a v e s was c l i p p e d , s e p a r a t e d , and bagged. N e e d l e s were not removed f r o m A b i e s l a s i o c a r p a t w i g s but were bagged w i t h t h e t w i g s t o w h i c h t h e y w e r e a t t a c h e d . The age o f t w i g s was i d e n t i f i e d by bud s c a r s , c o l o u r changes and b r a n c h i n g p a t t e r n s . The s a m p l e s were t a k e n t o t h e A g r i c u l t u r e Canada s t a t i o n i n P r i n c e G e o r g e i n D e c e m b e r 1981 and N o v e m b e r 1982. The s a m p l e s w e r e o v e n d r i e d f o r 48 h o u r s a t 5 0 ° C and w e i g h e d t o t h e n e a r e s t 0.1 g. 31 The s h r u b s w e r e d i v i d e d i n t o t h r e e c l a s s e s : c o n i f e r ( A JKL e_j3 1_ a_£dL££ a r__p_ a_) , t a l l g r o w i n g d e c i d u o u s ( A c e r _g, 3_ a_b_r_ii ni, A l l l J L L i l l l ^ I l i i L L f.£ii£» Be__t_u_la g l a n d u l o s a , B e t u l a p a p y r i f e r a , C o r y l u s c o r n u t a , C o r n u s s e r i c e a , P o p u l u s b a l s a m i f e r a , P o p u l u s _t£ e_m 11 ch; j>, S a 1 i x spp., and S o r b u s s p p . ) , and l o w g r o w i n g d e c i d u o u s ( L o n i c e r a i n v o l u c r a t a , Rosa spp., Sambucus r a c e m o s a , S p i r e a spp., V a c c i n i u m spp., and V i r b u r n u m e d u l e ) . R e g r e s s i o n e q u a t i o n s f o r s h r u b b i o m a s s e s t i m a t i o n were d e v e l o p e d f o r t h e t h r e e c l a s s e s . The f o l l o w i n g v a r i a b l e s were i n t r o d u c e d t o t h e r e g r e s s i o n a n a l y s i s as p r e d i c t o r v a r i a b l e s : - stem d i a m e t e r - canopy h e i g h t - canopy d e p t h - d i a m e t e r s q u a r e d - d i a m e t e r m u l t i p l i e d by h e i g h t - d i a m e t e r s q u a r e d m u l t i p l i e d by h e i g h t - d i a m e t e r m u l t i p l i e d by d e p t h - d i a m e t e r s q u a r e d m u l t i p l i e d by d e p t h - c a n o p y a r e a ( T T / A m u l t i p l i e d by maximum f o l i a g e w i d t h m u l t i p l i e d by w i d t h p e r p e n d i c u l a r t o maximum w i d t h ) - canopy volume (canopy a r e a m u l t i p l i e d by d e p t h , e x c e p t f o r A b i e s 1_ as^l o c a r__p_ a , w h i c h u s e s t h e f o r m u l a f o r c o n e v o l u m e , ( l / 3 ) i r r 2 m u l t i p l i e d by depth) - t h e n a t u r a l l o g s of each o f t h e p r e c e d i n g v a r i a b l e s 32 - b r o w s e c o n d i t i o n (0 f o r u n b r o w s e d , 1 f o r b r o w s e d , 2 f o r h e a v i l y browsed) - browse c o n d i t i o n s q u a r e d The r e s p o n s e v a r i a b l e s were t h e n a t u r a l l o g s of t w i g and l e a f w e i g h t s f o r t h e two d e c i d u o u s c l a s s e s and t h e n a t u r a l l o g o f c u r r e n t a n n u a l g r o w t h f o r A b i e s l a s i o c a r p a p r o d u c t i o n . The n a t u r a l l o g t r a n s f o r m a t i o n s w e r e r e q u i r e d b e c a u s e t h e v a r i a n c e s o f r e s p o n s e v a r i a b l e s w e r e p r o p o r t i o n a l t o t h e m a g n i t u d e o f p r e d i c t o r v a r i a b l e s and t h e r e was a t r e n d t o c u r v i l i n e a r i t y i n p l o t t e d d a t a . The s e t of p r e d i c t o r v a r i a b l e s w h i c h gave t h e h i g h e s t R f o r e a c h r e s p o n s e v a r i a b l e was d e t e r m i n e d w i t h t h e BMDP:9R " a l l p o s s i b l e s u b s e t s " r e g r e s s i o n ( F r a n e 1981). T h i s a n a l y s i s p r o d u c e d the f o l l o w i n g r e s u l t s : (5) In Y l = -4.0844 - 0.33605E-2(DTH) - 0.18700(BRSQ) + 0.81191(DSDL), A w h e r e I n Y l i s t h e n a t u r a l l o g o f A b i e s l _ a s i o c a r j > a p r o d u c t i o n , (6) In Y2 = 3.7015 + 0.31851E-l(DM) - 0.12855(BRSQ) - 0.31807(DSHL) + 0.83722(DSDL), where In Y2 i s t h e n a t u r a l l o g o f t a l l g r o w i n g d e c i d u o u s Twjs <' t r e e s , (7) In Y3 = -4.3546 - 0.79352E-1(BRSQ) + 0.48348(DSDL) + 0.23269(V0LNL), A w h e r e I n Y3 i s t h e n a t u r a l l o g o f t a l l g r o w i n g d e c i d u o u s l e a v e s , 33 (8) In Y4 = 2.1789 + 0.27834E-1(DTH) - 0.22498(BRSQ) + 0.20430(DSDL), A where In Y4 i s t h e n a t u r a l l o g of low g r o w i n g d e c i d u o u s t w i g s , (9) In Y5 = -5.1732 + 0.27929(DSDL) + 0.45939(V0LNL), A where In Y5 i s t h e n a t u r a l l o g of low g r o w i n g d e c i d u o u s l e a v e s . The p r e d i c t o r v a r i a b l e s were d e f i n e d as f o l l o w s : DM d i a m e t e r of t h e stem 2.5 cm above ground, DTH d e p t h of t h e canopy, DMLN n a t u r a l l o g o f DM, BRSQ t h e s q u a r e of t h e i n t e g e r r e p r e s e n t i n g browse c o n d i t i o n , DSHL n a t u r a l l o g of t h e s q u a r e of DM m u l t i p l i e d by canopy h e i g h t , DSDL n a t u r a l l o g o f the s q u a r e of DM m u l t i p l i e d by DTH, VOLNL n a t u r a l l o g o f canopy volume. 2.1.2. Shrub d e n s i t y e s t i m a t e s T h e d e n s i t y o f p a l a t a b l e s h r u b s 50 cm o r t a l l e r was e s t i m a t e d on 31 p l o t s , d u r i n g t h e p e r i o d f r o m l a t e J u n e t o l a t e A u g u s t 1982. T h e s e p l o t s i n c l u d e d 9 u n l o g g e d f o r e s t , 2 d i a m e t e r l i m i t l o g g e d s i t e s , 4 i n t e r m e d i a t e u t i l i z a t i o n l o g g e d s i t e s , 4 o l d c l e a r c u t s , 4 y o u n g c l e a r c u t s , 3 w e t l a n d s , 3 r i p a r i a n s h r u b , and 2 u p l a n d s h r u b s i t e s ( s e e F i g u r e 2 ) . The number and l o c a t i o n o f p l o t s w e r e s e l e c t e d t o r e p r e s e n t t h e 34 r a n g e of h a b i t a t w i t h i n e a c h t y p e based on v i s u a l e s t i m a t e s of t h e d i v e r s i t y of each h a b i t a t t y p e . A l l of t h e d a t a f r o m each h a b i t a t t y p e were combined t o p r o d u c e a s i n g l e e s t i m a t e i n t h e f o l l o w i n g a n a l y s e s . B a t c h e l e r ' s (1971) C o r r e c t e d P o i n t D i s t a n c e (CPD) method was used t o e s t i m a t e d e n s i t y . Most p l o t s were 70 m by 120 m. F i v e p a r a l l e l t r a n s e c t s 14 m a p a r t e x t e n d e d t h e l e n g t h of the p l o t . Twenty sample p o i n t s were l o c a t e d 6 m a p a r t a l o n g each t r a n s e c t . The d i s t a n c e b e t w e e n p o i n t s was s h o r t e n e d t o 4 m and t h e l e n g t h o f t h e t r a n s e c t t o 80 m i n 8 c a s e s w h e r e t h e h o m o g e n o u s v e g e t a t i o n t y p e was t o o s m a l l t o a c c o m m o d a t e t h e l o n g e r t r a n s e c t s o r w h e r e t h e g r o w t h was so d e n s e as t o r e s t r i c t movement. At e a c h p o i n t t h e d i s t a n c e t o t h e n e a r e s t p a l a t a b l e s h r u b , t h e d i s t a n c e f r o m t h e n e a r e s t s h r u b t o i t s n e a r e s t n e i g h b o u r , and t h e d i s t a n c e f r o m th e n e a r e s t n e i g h b o u r t o i t s n e a r e s t n e i g h b o u r were r e c o r d e d . P a l a t a b l e s h r u b s were t h o s e i n any o f t h e 17 s h r u b g r o u p s n o t e d a b o v e . The s p e c i e s o f e ach o f t h e t h r e e s h r u b s was r e c o r d e d . F o r e a c h o f t h e t h r e e s h r u b c l a s s e s , c o n i f e r , t a l l g r o w i n g d e c i d u o u s , and low g r o w i n g d e c i d u o u s , t h e d e n s i t y and v a r i a n c e o f t h e d e n s i t y o f s t e m s w e r e o b t a i n e d by a n a l y z i n g t h e CPD measurements w i t h t h e C a n a d i a n W i l d l i f e S e r v i c e (CWS) p r o g r a m w h i c h a n a l y s e s t h e s e d a t a a c c o r d i n g t o B a t c h e l e r 35 (1975). The p r o c e d u r e f o l l o w e d i n t h e a n a l y s i s of d e n s i t y and p r o b a b l e l i m i t s of e r r o r were d e s c r i b e d i n Boyd ( 1 9 7 9 ) . 2.1.3 Shrub biomass F o r e a c h s h r u b n e a r e s t t o t h e s y s t e m a t i c a l l y p l a c e d p o i n t , t h e h e i g h t , d e p t h , maximum w i d t h o f f o l i a g e , w i d t h p e r p e n d i c u l a r t o maximum w i d t h , stem d i a m e t e r 2.5 cm above t h e g r o u n d , and a b r o w s e c l a s s ( u n b r o w s e d , b r o w s e d , o r h e a v i l y b r o w s e d ) w e r e r e c o r d e d . F o r s u c h s h r u b s , p r o d u c t i o n was c a l c u l a t e d a s : (10) Y i j = a n t i l o g ( l n Y i j ) + e m s e / 2 , w h e r e Y i j i s t h e p r o d u c t i o n o f an i n d i v i d u a l s h r u b i n c l a s s i , In Y i j i s t h e p r o d u c t of t h e r e g r e s s i o n e q u a t i o n f o r c l a s s i (see e q u a t i o n s (5) t o ( 9 ) , mse i s t h e mean s q u a r e r e s i d u a l from t h e r e g r e s s i o n a n a l y s i s , and emse/Z ^ g a D i a s c o r r e c t i o n f a c t o r p r o p o s e d by M o u n t f o r d and Bunce ( 1 9 7 3 ) . O n l y b r o w s e 3.0 m o r l e s s i n h e i g h t was i n c l u d e d i n c a l c u l a -t i o n s , s i n c e T e l f e r ( 1 9 7 4 ) i n d i c a t e d t h a t moose r a r e l y t a k e b r o w s e f r o m a b o v e t h a t h e i g h t . The p r o p o r t i o n o f a s h r u b ' s p r o d u c t i o n t a l l e r t h a n 3.0 m was s u b t r a c t e d f r o m t h e s h r u b ' s t o t a l p r o d u c t i o n , a s s u m i n g t h a t b r o w s e was e v e n l y s p a c e d i n the s h r u b canopy. 36 The p r o d u c t i o n o f each s h r u b c l a s s was computed a s : (11) Mi = Y i j x D i , w h e r e M i i s b i o m a s s p e r m e t e r s q u a r e d f o r s h r u b s i n t h e i t h c l a s s , Y i j i s t h e mean p r o d u c t i o n of t h e i t h c l a s s , and Di i s t h e d e n s i t y of t h e s h r u b c l a s s as e s t i m a t e d by t h e CPD method. 2.1.4 V a r i a n c e of s h r u b biomass p r o d u c t i o n The v a r i a n c e o f t h e mean p r o d u c t i o n p e r s h r u b i n e a c h 2 2 c l a s s , S Y i , h a s two e l e m e n t s : s a m p l i n g e r r o r , S s a m i , and r e g r e s s i o n e r r o r , S 2 r e g i . (12) S 2 Y i = S 2 s a m i + S 2 r e g i , where (13) S 2 s a m i , = ( E ( Y i j - Y i j ) 2 1 ( n i - 1 ) ) / n i , (14) S z r e g i j = [ X l i j - X B l i X 2 i j - X B 2 i . . . ] c 11 c 12... c21 c 2 2 . . . X l i j - X B l i | (mse), X 2 i j - X B 2 i ' w h e r e X k i j i s t h e v a l u e o f t h e k t h i n d e p e n d e n t v a r i a b l e o f t h e j t h s h r u b i n t h e i t h s h r u b c l a s s , X B k i i s t h e mean o f t h e k t h i n d e p e n d e n t v a r i a b l e i n t h e i t h s h r u b c l a s s i n t h e d a t a w h i c h c r e a t e d t h e r e g r e s s i o n , c i j i s t h e i n v e r s e o f sums o f s q u a r e s and p r o d u c t s m a t r i x from t h e r e g r e s s i o n , and mse i s t h e mean s q u a r e r e s i d u a l f r o m t h e r e g r e s s i o n . E q u a t i o n (14) was a d a p t e d from Kozak ( 1 9 6 9 ) . (15) S 2 r e g i = Z ( S 2 r e g i j ) / n i 37 T h e r e i s no v a r i a n c e e s t i m a t e i n t h e u n t r a n s f o r m e d s c a l e w h i c h c o r r e c t l y r e p r e s e n t s t h e v a r i a n c e e s t i m a t e i n t h e n a t u r a l l o g s c a l e ( K o z a k 198A). S i n c e a r e g r e s s i o n v a r i a n c e i n t h e u n t r a n s f o r m e d s c a l e was n e c e s s a r y f o r f u r t h e r c a l c u l a -t i o n s , a v a r i a n c e o f a p p r o x i m a t e l y t h e c o r r e c t s i z e was e s t i m a t e d . T h i s e m p i r i c a l l y c o n s t r u c t e d v a r i a n c e was c a l c u -l a t e d by e s t a b l i s h i n g upper and l o w e r c o n f i d e n c e l i m i t s i n t h e n a t u r a l l o g s c a l e u s i n g S r e g i as e s t a b l i s h e d i n e q u a t i o n (15) and t h e n , (16) S 2 r e g i = { [ a n t i l o g ( u c l - l c l ) ] / 4 ) 2 . E q u a t i o n (16) a p p r o x i m a t e s a v a r i a n c e by r e v e r s i n g t h e c a l c u -l a t i o n of c o n f i d e n c e i n t e r v a l s i n t h e u n t r a n s f o r m e d s c a l e . (17) S 2 D i = ( P L E i ) 2 , w h e r e P L E i i s t h e p r o b a b l e l i m i t o f e r r o r f o r t h e s h r u b c l a s s g e n e r a t e d by t h e CPD a n a l y s i s ( B a t c h e l e r 1975). The v a r i a n c e o f t h e b i o m a s s p e r m e t e r s q u a r e d e s t i m a t e ( M i ) f o r a p a r t i c u l a r c l a s s i s : (18) S 2 M i = ( D i 2 ) ( S 2 Y i ) + ( Y i 2 ) ( S 2 D i ) - ( S 2 D i ) ( S 2 Y i ) w h e r e D i i s t h e d e n s i t y o f stems, and Y i i s t h e mean p r o d u c t i o n o f s t e m s . The v a r i a n c e o f t h e t o t a l b i o m a s s p e r m e t e r s q u a r e d f o r ea c h h a b i t a t t y p e (M) i s t h e sum o f t h e v a r i a n c e s of t h e sh r u b c l a s s e s w e i g h t e d by t h e number o f s h r u b s i n each c l a s s . E q u a t i o n (18) was t a k e n f r o m Goodman (1960, e q u a t i o n 5). 38 2.2 Herb L a y e r B i o m a s s E s t i m a t i o n C o n c u r r e n t w i t h t h e CPD e s t i m a t e s o f s h r u b d e n s i t y i n 1982 on e v e r y s e c o n d p o i n t t h e p r o d u c t i o n o f f o u r p l a n t c l a s s e s i n t h e h e r b l a y e r , (1) f e r n s and E ^ u i s e t u i spp., (2) f o r b s , (3) g r a s s e s , and (4) s e d g e s and r u s h e s , w e r e o c u l a r l y e s t i m a t e d w i t h a 0.25 m c i r c u l a r f r a m e . At e v e r y t e n t h p o i n t t h e s e f o u r c l a s s e s w e r e c l i p p e d t o g r o u n d l e v e l and b a g g e d s e p a r a t e l y . T h e s e s a m p l e s were t a k e n t o A g r i c u l t u r a l Canada R e s e a r c h S t a t i o n a t P r i n c e G e o r g e and o v e n d r i e d a t 5 0 ° C f o r 48 h o u r s i n N o v e m b e r 1982. The m e a s u r e d d r y w e i g h t s w e r e e n t e r e d as r e s p o n s e v a r i a b l e s and t h e e s t i m a t e d w e i g h t s as p r e d i c t o r v a r i a b l e s f o r a l l of t h e c l i p p e d f r a m e s f o r each o f t h e f o u r c l a s s e s , p r o d u c i n g t h e f o l l o w i n g r e g r e s s i o n e q u a t i o n s : (19) I n Y l = -1.7259 + 1.0098 l n ( o c u l a r e s t i m a t e Y l ) , w h e r e Y l i s t h e p r e d i c t e d d r y w e i g h t o f f e r n s a n d E q u i s e t u m spp. per 0.25 m , (20) In Y2 = -2.5713 + 1.2701 l n ( o c u l a r e s t i m a t e Y 2 ) , where Y2 i s t h e p r e d i c t e d d r y w e i g h t of f o r b s per 0.25 mZ, (21) In Y3 o -2.3199 + 1.3477 l n ( o c u l a r e s t i m a t e Y 3 ) , where Y3 i s t h e p r e d i c t e d d r y w e i g h t of g r a s s e s per 0.25 m1, (22) In Y4 = -0.37481 + 0.80965 l n ( o c u l a r e s t i m a t e Y 4 ) , w h e r e Y4 i s t h e p r e d i c t e d d r y w e i g h t o f s e d g e s and r u s h e s per 0.25 m . 39 The mean p r e d i c t e d d r y w e i g h t p r o d u c t i o n e s t i m a t e s r e s u l t i n g f r o m s o l v i n g t h e s e e q u a t i o n s f o r t h e o c u l a r e s t i m a t e s a t each f r a m e w i t h i n a h a b i t a t t y p e m u l t i p l i e d by 4 was t h e p r o d u c t i o n e s t i m a t e per m f o r each herb l a y e r c l a s s i n e a c h h a b i t a t . The v a r i a n c e f o r e a c h e s t i m a t e was c a l c u -l a t e d u s i n g e q u a t i o n s ( 1 2 ) , ( 1 3 ) , ( 1 4 ) , ( 1 5 ) , and ( 1 6 ) . 2.3 S e a s o n a l Biomass E s t i m a t e s I n summer t h e t o t a l b i o m a s s on a h a b i t a t t y p e was t h e sum of t h e p r o d u c t i o n of d e c i d u o u s l e a v e s and a l l f o u r h e r b l a y e r c l a s s e s . In f a l l , t o t a l b i o m a s s a v a i l a b l e was t h e sum of t h e p r o d u c t i o n of d e c i d u o u s t w i g s , g r a s s e s , s e d g e s and r u s h e s . I n i t i a l l y , t h e s t u d y assumed t h a t w i n t e r browse a v a i l a -b i l i t y c o u l d be u s e f u l l y a p p r o x i m a t e d by s u b t r a c t i n g t h e p r o d u c t i o n , measured i n summer, below the snow l i n e f r o m t h e t o t a l p r o d u c t i o n . The w i n t e r o f 1981-1982 p r o d u c e d v e r y deep snow, as much as 143 cm i n u n l o g g e d f o r e s t and 155 cm i n c l e a r c u t s . I t was o b v i o u s f r o m t h e damage t o browse t h a t t h e s i m p l e e s t i m a t e o f b r o w s e p r o d u c t i o n a v a i l a b i l i t y o u t l i n e d above was f a r f r o m a d e q u a t e . F i r s t , browse much h i g h e r t h a n maximum snow d e p t h s was b u r i e d . Second, i t a p p e a r e d p r o b a b l e t h a t more browse was b u r i e d i n some h a b i t a t s t h a n i n o t h e r s . To overcome t h i s p r o b l e m a s u r v e y of browse a v a i l a b i l i t y was appended to t h e snow s u r v e y s ( S e c t i o n 4). T h e r e were snow AO c o u r s e s on e a c h o f t h e f o r e s t - 1 o g g e d h a b i t a t t y p e s . A t e a c h o f t e n s t a t i o n s on e a c h snow c o u r s e , b r o w s e was m a r k e d w i t h c o l o u r coded t a p e i n each of f i v e h e i g h t z o n e s (0.0 t o <1.0 m, 1.0 t o <1.5 m, 1.5 t o <2.0 m, 2.0 t o <2.5 m, 2.5 t o 3.0 m). T h e r e w e r e f i v e f l a g s p e r h e i g h t z o n e a t e a c h o f t h e t e n s t a t i o n s f o r a t o t a l of 50 f l a g s per h e i g h t zone on e a c h snow c o u r s e . The number o f f l a g s v i s i b l e was c o u n t e d e a c h t i m e a snow c o u r s e was v i s i t e d d u r i n g t h e w i n t e r of 1982-1983 (Schwab e t a l . 1 9 8 5 ) . F o r t h e w i n t e r o f 1 9 8 2 - 1 9 8 3 t h e f o r a g e a v a i l a b l e was c a l c u l a t e d as f o l l o w s . In each h a b i t a t t y p e t h e c o n t r i b u t i o n o f t w i g s o f e a c h s h r u b t o e a c h h e i g h t z o n e was d e t e r m i n e d by a s s u m i n g t h a t t h e s h r u b ' s p r o d u c t i o n was e v e n l y s p r e a d o v e r t h e d e p t h o f i t s c a n o p y . I n e a r l y w i n t e r , f o r a g e a v a i l a b l e was t h e sum o f t h e p r o d u c t i o n o f d e c i d u o u s t w i g s i n e a c h h e i g h t z o n e m u l t i p l i e d by t h e p r o p o r t i o n v i s i b l e i n t h a t h e i g h t z o n e . I n m i d w i n t e r and l a t e w i n t e r f o r a g e a v a i l a b l e was t h e sum o f t h e p r o d u c t i o n o f d e c i d u o u s t w i g s and A b i e s 1 a s^ioc a ir j> £ f o l i a g e i n e a c h h e i g h t z o n e m u l t i p l i e d by t h e p r o p o r t i o n of browse v i s i b l e i n t h a t h e i g h t zone. The p r o p o r -t i o n o f b r o w s e v i s i b l e i n e a c h h e i g h t z o n e r e s u l t e d f r o m t h e d a t a g a t h e r e d n e a r e s t the median d a t e f o r each w i n t e r p e r i o d . Snow d e p t h s i n t h e e a r l y w i n t e r o f 1 9 8 2 - 1 9 8 3 w e r e a b o u t t h e same as t h o s e i n e a r l y w i n t e r 1 9 8 1 - 1 9 8 2 , b u t i n m i d - and l a t e w i n t e r snow d e p t h s were o n l y about h a l f t h o s e a t t a i n e d i n 41 t h e w i n t e r of 1981-1982. The p r o p o r t i o n s of browse v i s i b l e i n e ach h e i g h t zone i n e a r l y w i n t e r 1981-1982 were a s s i g n e d t h e same v a l u e s r e c o r d e d i n t h e w i n t e r of 1982-1983. In m i d - and l a t e w i n t e r 1981-1982, however, i t was assumed t h a t a l l o f t h e b r o w s e i n t h e 0.0 t o <1.0 m and 1.0 m t o <1.5 m z o n e s was u n a v a i l a b l e ^ F u r t h e r , t h e browse a v a i l a b l e i n t h e 1.5 t o <2.0 m, 2.0 t o <2.5 m, and 2.5 t o 3.0 m h e i g h t z o n e s was a s s i g n e d t h e a v a i l a b i l i t y o f b r o w s e i n t h e 0.0 t o <1.0 m, 1.0 t o <1.5 m, and 1.5 t o <2.0 m h e i g h t zones, i n t h e w i n t e r of 1982-1983, r e s p e c t i v e l y . The v a r i a n c e o f t h e b i o m a s s e s t i m a t e i n e a c h s e a s o n was t h e sum of t h e v a r i a n c e of each f o r a g e c l a s s d e t e r m i n e d t o be a v a i l a b l e d u r i n g t h a t s e a s o n . The f o r a g e c l a s s e s s e l e c t e d f o r i n c l u s i o n i n t h e f o r a g e a v a i l a b l e i n e a c h s e a s o n r e f l e c t t h e work o f Eastman (1978) and t h e r e v i e w by Peek ( 1 9 7 4 ) . 3. F o r e s t C over F o r e s t c o v e r s were q u a n t i f i e d u s i n g a method d e v e l o p e d by S t e y n (1980) t o c a l c u l a t e v i e w f a c t o r s i n u r b a n s e t t i n g s . In t h e c u r r e n t p r o j e c t 180° f i s h - e y e l e n s p h o t o g r a p h s were t a k e n f r o m u n d e r what w e r e j u d g e d t o be a v e r a g e o p e n i n g s i n t h e c a n o p i e s . Sky had t o be v i s i b l e a t t h e a p e x o f t h e s k y h e m i s p h e r e . T a k i n g t h e f i s h - e y e p h o t o g r a p h under an o p e n i n g i n t h e canopy does n o t b i a s t h e method t o w a r d p r o d u c i n g a low 42 f o r e s t c o v e r e s t i m a t e b e c a u s e t h e method i s most s e n s i t i v e t o t h e z o n e midway b e t w e e n t h e a p e x and h o r i z o n ( S t e y n 1 982). P h o t o g r a p h s w e r e t a k e n a t one s i t e on e a c h p l o t i n a l l h a b i t a t t y p e s e x c e p t i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g i n w h i c h p h o t o -g r a p h s w e r e t a k e n a t t h r e e s i t e s : an o p en s i t e , a p a r t i a l c a n o p y , and a c l o s e d c a n o p y s i t e on e a c h p l o t . P h o t o g r a p h s w e r e t a k e n b e t w e e n 25 A u g u s t and 30 A u g u s t 1982. F i g u r e 2 s h o w s t h e l o c a t i o n s o f p l o t s on w h i c h f i s h - e y e p h o t o g r a p h s were t a k e n . The p h o t o g r a p h s w e r e d e v e l o p e d on 20.3 cm by 25.4 cm (8 i n by 10 i n ) p a p e r and c o v e r c a l c u l a t e d u s i n g t h e a l g o r i t h m p r o v i d e d by S t e y n ( 1 9 8 0 ) . The f o r e s t c o v e r f o r e a c h h a b i t a t e q u a l l e d t h e mean c o v e r o f t h e p l o t s w i t h i n t h a t h a b i t a t . The f o r e s t c o v e r f o r i n t e r m e d i a t e u t i l i z a t i o n p l o t s was t h e mean o f t h e t h r e e c o v e r s c a l c u l a t e d f o r e a c h p l o t . Summer f o r e s t c o v e r was a l l c o v e r a p p e a r i n g on a p h o t o g r a p h . F a l l and w i n t e r c o v e r was c o n i f e r o u s f o l i a g e p l u s t h e s t e m s and l a r g e b r a n c h e s of d e c i d u o u s t r e e s on a p h o t o g r a p h . 4. Snow Depth I n t h e f a l l s o f 1981 and 1982, snow c o u r s e s w e r e l a i d o u t on e a c h o f a r e c e n t c l e a r c u t , an o l d c l e a r c u t , an i n t e r m e d i a t e u t i l i z a t i o n l o g g e d s i t e , a d i a m e t e r l i m i t l o g g e d s i t e , and an u n l o g g e d f o r e s t . A l l snow c o u r s e s w e r e on t h e moose w i n t e r 43 r a n g e on t h e n o r t h b a n k o f t h e F r a s e r R i v e r . F i g u r e 2 i n d i c a t e s t h e p l o t s w h i c h c o n t a i n e d snow c o u r s e s . E ach snow c o u r s e c o n s i s t e d of t e n s t a t i o n s a r r a n g e d 30 m a p a r t i n an a p p r o x i m a t e c i r c l e . The l o c a t i o n s o f s t a t i o n s w e r e c h o s e n t o r e p r e s e n t t h e r a n g e o f c o n d i t i o n s p r e s e n t on e a c h s i t e . E a ch s t a t i o n had an i d e n t i f y i n g marker, u s u a l l y a s m a l l c o n i f e r l i m b e d and c u t o f f a b o u t 2.5 m above t h e ground. A l l v e g e t a t i o n and d e b r i s w e r e c l e a r e d away t o g r o u n d l e v e l w i t h i n 1 m o f t h e m a r k e r . Snow d e p t h and d e n s i t y w e r e t o be measured once e v e r y two weeks as l o n g as snow p e r s i s t e d . T h i s s c h e d u l e was met d u r i n g t h e w i n t e r o f 1 9 8 2 - 1 9 8 3 , b u t deep l i g h t snow made t r a v e l d i f f i c u l t d u r i n g t h e w i n t e r o f 1981-1982 and snow measurement was e r r a t i c . 5. A i r T e m p e r a t u r e T h e r m o g r a p h s w e r e l o c a t e d a t f i v e s i t e s on t h e s t u d y a r e a : (1) a d i a m e t e r l i m i t l o g g e d s i t e a t 611 m i n e l e v a t i o n , (2) an u n l o g g e d f o r e s t a t 772 m, (3) an i n t e r m e d i a t e u t i l i z a -t i o n s i t e a t 778 m, (4) a r e c e n t c l e a r c u t a t 853 m, (5) an o l d c l e a r c u t a t 871 m. F i g u r e 2 shows w h e r e t h e r m o g r a p h s w e r e l o c a t e d . T h e r m o g r a p h s r e c o r d e d d a i l y m i n i m u m and maximum t e m p e r a t u r e s . I n s t a l l a t i o n o f t h e t h e r m o g r a p h s b e g a n on 6 N o v e m b e r 1 9 8 0 a n d was c o m p l e t e d on 5 D e c e m b e r 1 9 8 0 . T h e r m o g r a p h r e a d i n g s were t e r m i n a t e d on 1 June 1983. 44 IV RESULTS F i g u r e 3 i l l u s t r a t e s t h e p a t t e r n o f m o o s e h a b i t a t o c c u p a t i o n i n f o r e s t / 1 o g g e d h a b i t a t s d u r i n g t h e s i x s e a s o n s r e c o g n i z e d i n t h i s s t u d y ( T a b l e 3). F i g u r e 3 i n c l u d e s a l l d a t a g a t h e r e d d u r i n g each s e a s o n . In s p r i n g u n l o g g e d f o r e s t had f e w e r moose l o c a t i o n s t h a n e x p e c t e d and d i a m e t e r l i m i t l o g g i n g a p p r o x i m a t e l y t h e same number of l o c a t i o n s e x p e c t e d , i f l o c a t i o n s o c c u r r e d a t random. The more o p e n l o g g e d h a b i t a t s , i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g , o l d c l e a r c u t s , and r e c e n t c l e a r c u t s had more moose l o c a t i o n s t h a n e x p e c t e d , i f l o c a t i o n s o c c u r r e d a t r a n d o m . However, t h r o u g h summer, moose moved i n t o u n l o g g e d f o r e s t f r o m l o g g e d a r e a s , e s p e c i a l l y c l e a r c u t s . T he s p r i n g t o summer t r e n d c o n t i n u e d i n t o f a l l when moose o c c u p a t i o n o f u n l o g g e d f o r e s t p e a k e d and t h e r e w e r e r e l a t i v e l y few l o c a t i o n s i n l o g g e d h a b i t a t s . T h e r e was a d r a m a t i c r e d u c t i o n i n u n l o g g e d f o r e s t u s e i n e a r l y w i n t e r t o a b o u t h a l f o f i t s f a l l l e v e l b e c a u s e moose moved i n t o l o g g e d h a b i t a t s , p a r t i c u l a r l y i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g . I n m i d w i n t e r t h e number o f l o c a t i o n s i n d i a m e t e r l i m i t l o g g i n g and u n l o g g e d f o r e s t i n c r e a s e d as moose moved t o t h o s e h a b i t a t s f r o m i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g . T h e r e w e r e a l s o n o t i c e a b l e e a r l y t o m i d w i n t e r r e d u c t i o n s o f moose 4 5 F i g u r e 3. P r o p o r t i o n of moose l o c a t i o n s i n f o r e s t / l o g g e d h a b i t a t types Table 3. Number of moose locations i n forest/logged habitats by year and season Total a l l Season FR DL IU OC RC NF Habitats p a 1980 21.2(37)b 2.5(4) 16.7(29) 3.8(7) 3.2( 6) 9.7(17: 57.0 p<.001 Spring 1981 20.1(48) 0.0 5.5(13) 2.3(5) 4.8(12) 9.3(22: 42.0 p=.316 1982 32.2(43) 10.8(14) 8.0(11) 3.8(5) 7.2(10) 13.0(17: 75.0 p=.163 1983 26.8(40) 6.3( 9) 12.8(19) 0.0 9.0(13) 12.0(18: 67.0 .01<p<.05 (42) c ( 7) (18) (4) (10) (19: 1980 23.3(75) 0.0 1.3( 4) 0.5(2) 0.0 5.8(18: 31.0 p=.139 Summer 1981 43.7(68) 0.0 4.5( 7) 0.0 0.0 15.8(25: 64.0 .001<p<.01 1982 76.7(55) 11.3( 8) 22.7(16) 1.0(1) 3.3( 2) 25.0(18: 140.0 .01<p<.05 (61) ( 5) (12) (1) ( 1) (20: 1980 23.8(77) 0.0 0.5( 2) 0.0 0.0 6.7(22: 31.0 .01<p<.05 F a l l 1981 23.0(74) 0.0 0.0 0.0 0.0 8.0(26: 31.0 .01<p<.05 1982 47.8(58) 5.5( 7) 6.3( 8) 0.0 1.0( 1) 21.3(26: 82.0 .01<p<.05 (70) ( 2) ( 3) ( 1) (24: Early 1 31-82 17.7(45) 3.0( 8) 9.8(25) 0.5(1) 1.3( 3) 6.7(17: 39.0 p=.078 Winter 82-83 16.0(24) 0.0 33.8(51) 5.5(8) 4.0( 6) 6.7(10: 66.0 p<.001 (32) ( 3) (42) (6) ( 5) (14: Mid- 1982 42.5(42) 18.0(18) 26.8(26) 3.3(3) 0.3( 0) 11.0(11) 102.0 .01<p<.05 winter 1983 69.4(41) 14.1( 8) 64.2(38) 7.7(5) 1.5( 1) i 2 . u 7: 169.0 p<.001 (38) (12) (34) (4) ( 1) ( 9 : Late 1982 59.5(37) 29.2(18) 42.3(26) 5.8(4) 7.0( 4) 18.2(11) 162.0 p=.096 Winter 1983 31.0(41) 5.5( 7) 24.0(32) 5.0(7) 3.0( 4) 6.5( 9: 75.0 p=.lll (38) (15) (28) (5) ( 4) (10) FR unlogged forest 0C old clearcut DL diameter limit logging RC recent clearcut IU intermediate u t i l i z a t i o n logging NF nonforest a probability of random moose distribution with respect to habitat type b bracketed figure i s the percentage of locations i n the habitat type c bracketed figure at the bottom of the year/season/habitat column i s the percentage of locations i n that season/habitat for a l l years 4 8 l o c a t i o n s i n c l e a r c u t s . I n l a t e w i n t e r t h e n u m b e r o f l o c a t i o n s i n u n l o g g e d f o r e s t and d i a m e t e r l i m i t l o g g i n g s t a y e d r e l a t i v e l y even, but t h e r e was some movement t o c l e a r c u t s f r o m i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g . From l a t e w i n t e r t o s p r i n g r e c e n t c l e a r c u t s and u n l o g g e d f o r e s t g a i n e d l o c a t i o n s w h i l e i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g and d i a m e t e r l i m i t l o g g i n g l o s t moose l o c a t i o n s . The p r o p o r t i o n o f l o c a t i o n s i n o l d c l e a r c u t s was t h e same i n l a t e w i n t e r and s p r i n g . However, t h i s g e n e r a l t r e n d d i s g u i s e s i m p o r t a n t y e a r t o y e a r d i f f e r e n c e s i n moose h a b i t a t a s s o c i a t i o n s . A l s o , t h e r e w e r e s e a s o n t o s e a s o n c h a n g e s i n h a b i t a t a v a i l a b i l i t y t h a t a f f e c t t h e c a l c u l a t i o n of h a b i t a t a s s o c i a t i o n s . The f o l l o w i n g i s a s e a s o n a l a c c o u n t of moose h a b i t a t o c c u p a t i o n . 1. S p r i n g The s p r i n g s e a s o n e x t e n d e d f r o m t h e o n s e t o f s p r i n g moose m i g r a t i o n u n t i l 15 J u n e . The o n s e t o f s p r i n g m i g r a t i o n was e s t a b l i s h e d by t h r e e c r i t e r i a : F i r s t , i n l a t e w i n t e r , w e e k l y moose m o v e m e n t s a p p e a r r a n d o m and were no more t h a n a few hu n d r e d m e t e r s p e r week. W i t h the o n s e t of m i g r a t i o n , moose make w e e k l y movements of as much as 20 km, l e a d i n g d i r e c t l y t o th e a r e a i n w h i c h t h e y summer. Second, t h e l a r g e d i r e c t i o n a l s p r i n g movements r e s u l t i n an i n c r e a s e i n t h e e l e v a t i o n on t h e 49 a r e a s o c c u p i e d . T h i r d , t h e r e i s a n o t i c e a b l e s h i f t t o more open h a b i t a t w h i c h i s c o n c o m i t a n t w i t h s p r i n g m o v e m e n t s . M o o s e w e r e l o c a t e d d u r i n g f o u r s p r i n g p e r i o d s : 1980, 1981, 1982, and 1983. The f i r s t i n d i c a t i o n s o f s p r i n g m i g r a t i o n w e r e on 14 A p r i l , 8 May, 14 May, a n d 22 A p r i l i n t h e s e f o u r y e a r s r e s p e c t i v e l y . A t i m e s e r i e s o f t h e C o l e i n d e x o f a s s o c i a t i o n f o r e a c h h a b i t a t t y p e i s p r e s e n t e d i n F i g u r e s 4, 5, 6, and T a b l e 4. S p r i n g e s t i m a t e s o f a v a i l a b l e f o r a g e w e r e t h e same a s t h o s e c a l c u l a t e d f o r summer ( T a b l e 5). F o r e s t c o v e r e s t i m a t e s a r e i n T a b l e 6. The s p r i n g s of 1980 and 1983 p r o d u c e d 2 s i g n i f i c a n t l y nonrandom d i s t r i b u t i o n s , p ( X 5 d f ) < 0.1, of moose w i t h r e s p e c t t o h a b i t a t t y p e s . W i t h t h e e x c e p t i o n o f s p r i n g 1983, t h e c o r r e l a t i o n b e t w e e n a v a i l a b l e f o r a g e and moose h a b i t a t a s s o c i a t i o n s was g r e a t e r t h a n t h a t b e t w e e n f o r e s t c o v e r and h a b i t a t a s s o c i a t i o n s ( T a b l e 7). The n e g a t i v e c o r r e l a t i o n between a v a i l a b l e f o r a g e and h a b i t a t a s s o c i a t i o n s i n 1983 and l o w p r o b a b i l i t y o f t h e c o r r e l a t i o n i n 1981 i n d i c a t e t h a t t h e r e l a t i o n s h i p i s not s t r o n g . T h e r e was a p p a r e n t l y l i t t l e p r e s s u r e f o r moose t o occupy any p a r t i c u l a r h a b i t a t d u r i n g s p r i n g . Two o f f o u r s p r i n g s e a s o n s p r o d u c e d r a n d o m d i s t r i b u t i o n s o f moose l o c a t i o n s . A l s o , t h e s p r i n g s e a s o n , more so t h a n any o t h e r , t e n d e d t o p r o d u c e a c l u s t e r of h a b i t a t a s s o c i a t i o n s n e a r z e r o ( F i g u r e s 4 , 5, and 6 ). 50 F i g u r e 4. The C o l e ( C y ) i n d e x o f a s s o c i a t i o n b e t w e e n moose l o c a t i o n s and f o r e s t / l o g g e d h a b i t a t t y p e s f r o m s p r i n g 1980 t o f a l l 1980. O p e n a n d c l o s e d c i r c l e s r e p r e s e n t n o n s i g n i f i c a n t a s s o c i a t i o n s and s i g n i f i c a n t a s s o c i a t i o n s r e s p e c t i v e l y . The a s t e r i s k i n d i c a t e s s e a s o n s i n w h i c h t h e d i s t r i b u t i o n 2 o f moose l o c a t i o n s was nonrandom, p (X5<jf) < 0.1, w i t h r e s p e c t t o t h e a v a i l a b i l i t y of h a b i t a t t y p e s . • • - _ • \ y X ^ :% N V * * 0 \ 'spring summer * fall early winter midwinter recent c learcut old c lea rcu t in te rmed iate u t i l i za t ion d i a m e t e r l imit un logged forest 52 F i g u r e 5. C o l e ( C y ) i n d e x o f a s s o c i a t i o n b e t w e e n moose l o c a t i o n s and f o r e s t / l o g g e d h a b i t a t t y p e s f r o m s p r i n g 1981 t o l a t e w i n t e r 1982. Open and c l o s e d c i r c l e s r e p r e s e n t n o n s i g n i f i c a n t a s s o c i a -t i o n s and s i g n i f i c a n t a s s o c i a t i o n s r e s p e c t i v e l y . The a s t e r i s k i n d i c a t e s s e a s o n s i n w h i c h t h e d i s t r i b u t i o n of 2 moose l o c a t i o n s was nonrandom, p ( X 5 d f ) < 0.1, w i t h r e s p e c t t o t h e a v a i l a b i l i t y of h a b i t a t t y p e s . 0 . 6 0 . 3 X <D "D o o O - 0 . 6 - 0 . 9 -1 .2 - • ° \ " \ * I / A * " I * // \ / / • \ x \ <• / I i 1 1 — 1 spring •summer 'fall •early winter •midwinter 'late winter recent c lea rcu t old c learcut i n te rmed ia te u t i l i za t ion d i a m e t e r l imit 54 F i g u r e 6. C o l e ( C y ) i n d e x o f a s s o c i a t i o n b e t w e e n moose l o c a t i o n s and f o r e s t / l o g g e d h a b i t a t t y p e s f r o m s p r i n g 1982 t o s p r i n g 1983. Open and c l o s e d c i r c l e s r e p r e s e n t n o n s i g n i f i c a n t a s s o c i a t i o n s and s i g n i f i c a n t a s s o c i a t i o n s r e s p e c t i v e l y . The a s t e r i s k i n d i c a t e s s e a s o n s i n w h i c h t h e d i s t r i b u t i o n of moose l o c a -2 t i o n s was nonrandom, p ( X 5 d f ) < 0.1, w i t h r e s p e c t t o t h e a v a i l a b i l i t y of h a b i t a t t y p e s . 0.6 0.3 X <D "O c o o o •0.3H • 0 . 6 H - 0 . 9 H -1.2 • * - - *• *^^^  * * * t • •• m • /P ° v • \ ^^^^  \ \ \ • ^ ^ ^ \ t > / V \ / \ / • • / V ' r \ \ \ \ \ • recent c lea rcu t old c lea rcu t i n te rmed ia te ut i l i zat ion d i a m e t e r l imit 56 Table 4. Cole index and standard error (in brackets) for forest/logged habitats Season Year FR DL -0.40(0.47) -1.00(0.55)b 0.08(0.04) 0.02(0.03) -1.00(0.64)b -1.00(0.44)b 0.01(0.02) -1.00(0.64)b -1.00(0.44)b -0.09(0.39) 0.00(0.04) -1.00(0.44)b 0.04(0.04)° -0.40(0.19)b 0.05(0.03)° -0.47(0.29)b IU 0.21(0.50)c 0.03(0.05) 0.00(0.04) 0.10(0.04)° -0.59(0.52)b -0.33(0.37) 0.06(0.03)° -0.85(0.53)b -1.00(0.52)b -0.27(0.31) 0.16(0.05)° 0.46(0.04)° 0.04(0.05) 0.19(0.04)° 0.04(0.04) 0.11(0.06)° 0C 0.03(0.02)° 0.02(0.03) 0.02(0.03) -1.00(0.63)b -0.56(0.92) -1.00(0.63)b 0.80(0.43)b -1.00(0.92)b -1.00(0.92)b -1.00(0.57)b -0.65(0.82) 0.05(0.02) -0.48(0.38)b -0.28(0.30) -0.43(0.30)b 0.00(0.03) RC -0.23(0.47) 0.05(0.04)° 0.02(0.03) 0.07(0.03)° -1.00(0.64)b -1.00(0.45)° -0.67(0.30)b -1.00(0.64)b -1.00(0.64)b -0.83(0.39)b -0.53(0.57) -0.17(0.44) -0.97(0.31)b -0.91(0.23)b -0.55(0.24)b -0.58(0.36)b Spring 1980a -0.32(0.12)b 1981 -0.13(0.14) 1982 -0.22(0.10)b 1983a -0.27(0.ll) b Summer 1980 0.45(0.20)° 1981a 0.30(0.14)° 1982a 0.00(0.08) F a l l 1980a 0.49(0.20)° 1981a 0.43(0.23)° 1982a 0.08(0.12) Early 1981-823 -0.18(0.14)b Winter 1982-833 -0.56(0.ll) b Mid- 1982a 0.11(0.07)° Winter 1983a 0.10(0.05)° Late 1982a 0.04(0.05) Winter 1983 0.11(0.06)° FR, unlogged forest; DL, diameter l i m i t logging; IU, intermediate utilization; 0C, old clearcut; RC, recent clearcut a p<0.1 that moose were randomly distributed with respect to habitat type i n the year and season indicated b significant negative association of moose with habitat type ° significant positive association of moose with habitat type 57 Table 5. Summer available forage (g/m ) Ferns Sedges Deciduous and and Total Habitat Leaves Equisteum Forbs Grasses Rushes Available Forest 14,46(0.696)a 1.98(0.114) 3.79(0.135) 0.10(0.116) .26(0.216) 20.60(1.032) D l 16 r Limit 35.24(4.034) 0.80(0.108) 4.16(0.266) 0.09(0.166) 0.0 (0.0) 40.29(4.098) Limit 35.24(4.034) 0.80(0.108) 4.16(0.266) 0.09(0.166) 0.0 (0.0) 40.29(4.098) Inter-mediate U t i l i z a -tion 45.56(3.508) 3.77(0.609) 13.75(1.388) 0.88(0.159) 0.14(0.217) 64.11(3.831) Old Clearcut 34.37(2.828) 1.44(0.161) 13.64(1.092) 1.24(0.198) 0.88(0.249) 51.57(3.114) Recent Clearcut 11.72(1.746) 4.44(0.579) 13.31(1.604) 2.21(0.206) 1.34(0.304) 33.03(2.396) Upland Shrub 38.35(5.464) 18.47(3.297) 10.15(1.106) 0.11(0.117) 0.0 (0.0) 67.08(5.863) Riparian Shrub 165.66(8.712) 5.47(0.704) 4.53(0.284) 1.44(0.180) 0.96(0.353) 178.05(8.798) Wetland 9.24(0.967) 0.0 (0.0) 1.45(0.110) 0.07(0.116) 6.36(0.352) 17.13(1.230) bracketed figures are standard deviations 58 Table 6. Estimates of forest cover as a decimal fraction of the sky hemisphere Habitat n Spring and Summer mean + s.d. n F a l l and Early Winter mean + s.d. n Mid and Late Winter 3 mean + s.d. Forest 8 0.772 + 0.046b 8 0.772 + 0.046 5 0.768 + 0.058 Diameter Limit 2 0.706 + 0.001 2 0.701 + 0.001 2 0.701 + 0.001 Intermediate Utili z a t i o n 4 0.366 + 0.059 4 0.355 + 0.080 4 0.355 + 0.080 Old Clearcut 4 0.145 + 0.140 4 0.060 + 0.029 2 0.059 + 0.020 Recent Clearcut 4 0.032 + 0.029 4 0.026 + 0.028 2 0.040 + 0.037 a contains data from winter range plots only b estimates are the fraction of the sky hemisphere obscured by vegetation 59 Table 7. Correlations between Cole indices and available forage and between Cole indices and forest cover Season Year Correlation Coefficient Cy and forage Correlation Coefficient Cy and forest cover P C Spring 1980d 0.835 0.045 -0.499 0.782 0.218 1981 0.160 0.409 -0.636 0.855 0.145 1982 0.574 0.179 -0.470 0.764 0.236 1983d -0.075 0.559 0.226 0.371 0.629 Summer 1980 -0.442 0.749 0.512 0.212 0.788 1981d -0.329 0.684 0.585 0.171 0.829 1982d -0.016 0.508 0.859 0.034 0.966 F a l l 1980d -0.677 0.877 0.631 0.147 0.853 1981d -0.727 0.903 0.623 0.151 0.849 1982d -0.267 0.652 0.961 0.003 0.997 Early 1981-82d -0.252 0.641 0.682 0.119 0.881 Winter 1982-83d 0.614 0.156 -0.650 0.864 0.136 Mid- 1982d -0.033 0.519 0.852 0.037 0.963 Winter 1983d 0.185 0.397 0.519 0.219 0.791 Late 1982d -0.012 0.508 0.851 0.007 0.963 Winter 1983 0.202 0.386 0.278 0.335 0.665 probability of observing an r value > r calculated probability of observing an r value > r calculated probability of observing an r value < r calculated p<0.1 that moose were distributed randomly with respect to habitat type i n the year and season indicated 60 T h e l a c k o f s t r o n g h a b i t a t a s s o c i a t i o n s i n s p r i n g i n d i c a t e s t h a t t h e r e was l i t t l e p r e s s u r e t o o c c u p y any p a r t i c u l a r h a b i t a t due t o f o r a g e p r o d u c t i o n o r t o e s c a p e t h e r m a l c o n d i t i o n s . T h e r e i s l i t t l e p r e s s u r e f r o m h e a t s t r e s s a s a l l h a b i t a t s a r e w e l l b e l o w K n o r r e ' s ( 1 9 5 9 ) 2 0 ° C g u i d e l i n e , d u r i n g a l l s p r i n g p e r i o d s when t e m p e r a t u r e d a t a were g a t h e r e d ( T a b l e 8). The mean maximum t e m p e r a t u r e a t t h e P r i n c e G e o r g e a i r p o r t was 16.9° d u r i n g t h e s p r i n g o f 1980 ( C a n a d a , A t m o s p h e r i c E n v i r o n m e n t S e r v i c e 1 9 8 0 ) , i n d i c a t i n g t h a t t h e r e was l i t t l e h e a t s t r e s s i n t h e s p r i n g o f 1980. A l s o , a s i n g l e h i g h c o r r e l a t i o n between a v a i l a b l e f o r a g e and C o l e i n d i c e s i n f o u r s p r i n g s e a s o n s ( T a b l e 7) i n d i c a t e s l i t t l e p r e s s u r e t o s e l e c t f o r h a b i t a t s w h i c h c o n t a i n e d l a r g e amounts of f o r a g e . S i n c e h a b i t a t a s s i g n m e n t s of a v a i l a b l e f o r a g e were based on summer f o r a g e , t h e a v a i l a b l e s p r i n g f o r a g e r e p r e s e n t t h e f o r a g e on t h e p l o t s i n t h e l a s t d a y s o f s p r i n g r a t h e r t h a n d u r i n g t h e whole s e a s o n . In e a r l y t o m i d s p r i n g th e amount of a v a i l a b l e f o r a g e was more l i k e l y r e l a t e d t o t h e amount of sun w h i c h r e a c h e d t h e g r o u n d t o m e l t snow and s t i m u l a t e e a r l y g r o w t h . N e g a t i v e c o r r e l a t i o n s b e t w e e n m o o s e h a b i t a t a s s o c i a t i o n s and f o r e s t c o v e r w e r e c o n s i s t e n t l y h i g h e r t h a n c o r r e l a t i o n s between h a b i t a t a s s o c i a t i o n s and a v a i l a b l e f o r a g e ( T a b l e 7 ) . S e l e c t i o n f o r o p e n h a b i t a t s a p p e a r s t o e x p l a i n moose s p r i n g b e h a v i o u r b e t t e r t h a n a v a i l a b l e f o r a g e based on 61 Table 8. Seasonal mean minimum and mean maximum a i r temperatures (°C) Year 1981 1982 1983 Season FR DL IU OC RC Spring 5.2 a 2.5 b 4.0 3.9 5.4 15.9 a 18.0 b 16.6 15.9 17.1 Summer 8.2 b 5.4 6.8 6.5 9.3 2 0 . l b 23.1 20.0 20.8 22.3 F a l l 2.3 0.4 0.9 0.7 2.2 8.8 11.2 10.0 9.7 10.9 Early -8.8 b - 1 0 . l b -10.1 -10.8 -9.0 Winter -4.2 b - 4.9 b - 2.0 - 4.3 -3.5 Midwinter -13.7 b -17.9 b -15.6 b -16.6 -13.1 - 8.0 b - 7.8 b - 7.4 b - 8.0 - 6.8 Late -3.8 b -6.3 b -4.9 b -5.2 -3.0 Winter 5.4 b 7.0 b 6.7 b 5.9 7.2 Spring 3.5 0.4 2.8 b 3.0 5.9 16.9 19.4 17.9 b 17.5 18.8 Summer 8.2 6.2 7.3 8.9 18.9 20.7 18.4 b 19.1 19.5 F a l l 1.3 b -1.2 0.5 b -0.4 2.9 7.1 b 8.8 8.3 b 7.8 8.6 Early -7.8 -11.1 -8.9 -9.7 -7.5 Winter -3.4 -4.0 -3.0 -4.0 -3.3 Midwinter -5.0 -9.0 -5.8 -6.6 -4.4 0.8 0.5 3.5 1.0 1.0 Late -3.8 -6.9 -4.0 -4.3 -2.5 Winter 4.7 7.3 6.9 6.4 6.4 Spring 4.1 1.4 2.1 2.2 4.8 15.8 17.5 16.4 15.3 16.9 FR, unlogged forest; DL, diameter l i m i t logging; IU, intermediate u t i l i z a t i o n logging; 0C, old clearcut; RC, recent clearcut a i n the data pairs, the mean minimum temperature i s l i s t e d on top and the mean maximum temperature i s on the bottom contains regression estimates f o r missing data 62 summer abundance. T h i s o b s e r v a t i o n s h o u l d be t e m p e r e d by t h e f a c t t h a t o n l y two o f f o u r s p r i n g s p r o d u c e d a p r o b a b i l i t y o f a random d i s t r i b u t i o n of l e s s t h a n 0.1. 2. Summer The summer s e a s o n b e g a n on 16 J u n e and e n d e d on 15 S e p t e m b e r e a c h y e a r . M o o s e l o c a t i o n s w e r e g a t h e r e d d u r i n g t h r e e summer p e r i o d s : 1980, 1981, and 1982. A summary o f moose l o c a t i o n s w i t h r e s p e c t t o h a b i t a t t y p e i s i n T a b l e 3 and a summary o f C o l e i n d i c e s i s i n F i g u r e s 4, 5, 6, and T a b l e 4. 2 Moose were s i g n i f i c a n t l y n o n r a n d o m l y d i s t r i b u t e d p ( X 5 d f ) < 0.1, w i t h r e s p e c t t o h a b i t a t t y p e d u r i n g t h e summers o f 1981 a n d 1 9 8 2 . I n t h e s u m m e r o f 1 9 8 0 m o o s e w e r e r a n d o m l y d i s t r i b u t e d , b u t t h i s may h a v e b e e n a r e s u l t o f a l a c k o f o b s e r v a t i o n s i n t h a t s e a s o n . The summers o f 1980 and 1981 w e r e v e r y s i m i l a r t o e a c h o t h e r i n t e r m s o f m o o s e d i s t r i b u t i o n , but b o t h c o n t r a s t w i t h moose d i s t r i b u t i o n d u r i n g t h e summer o f 1982. D a t a f r o m 1980 and 1981 b o t h p r o d u c e d a s t r o n g p o s i t i v e a s s o c i a t i o n b e t w e e n moose l o c a t i o n s and u n l o g g e d f o r e s t h a b i t a t , b u t n e g a t i v e a s s o c i a t i o n s b e t w e e n moose and a l l l o g g e d h a b i t a t s . I n 1982 t h e r e was a n e u t r a l a s s o c i a t i o n b e t w e e n m o o s e a n d u n l o g g g e d f o r e s t . T h e a s s o c i a t i o n between moose and d i a m e t e r l i m i t l o g g i n g became n e u t r a l . T h e r e was a p o s i t i v e a s s o c i a t i o n between moose and 63 i n t e r m e d i a t e u t i l i z a t i o n . The r e l a t i o n s h i p between moose and a l l c l e a r c u t s r e m a i n e d n e g a t i v e . The c o r r e l a t i o n s b e t w e e n f o r e s t c o v e r and C o l e i n d i c e s were a l w a y s much h i g h e r t h a n c o r r e l a t i o n s between a v a i l a b l e f o r a g e and C o l e i n d i c e s ( T a b l e 7). The h e a v y f o r a g e b e a r i n g o l d c l e a r c u t s , 51.57 g/m ( T a b l e 5), had s i g n i f i c a n t n e g a t i v e a s s o c i a t i o n s w i t h moose i n b o t h 1981 and 1982. On t h e o t h e r e x t r e m e t h e p o o r e s t f o r a g e p r o d u c e r , u n l o g g e d f o r e s t 20.60 g/m^ ( T a b l e 5), a l w a y s had l a r g e numbers of moose l o c a t i o n s i n summer. P a r t o f t h e e x p l a n a t i o n f o r t h e c h a n g e i n p a t t e r n o f moose a s s o c i a t i o n s b e t w e e n summer 1981 and 1982 was t h e d i f f e r e n c e i n t e m p e r a t u r e s b e t w e e n t h e two s e a s o n s . The summer o f 1981 p r o d u c e d a mean maximum o f 20.0° w h i l e t h e s u m m e r o f 1 982 p r o d u c e d a mean m a x i m u m o f 1 8 . 4 ° i n i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g ( T a b l e 8), t h e h a b i t a t w h i c h g a i n e d t h e g r e a t e s t p r o p o r t i o n o f moose l o c a t i o n s i n summer 1982 as c o m p a r e d w i t h summer 1981. Maximum t e m p e r a t u r e s e x c e e d e d 2 0 ° on 60 d a y s i n 1981 v e r s u s 36 d a y s i n 1982. The f i n a l 15 d a y s o f J u n e a c c o u n t e d f o r 15 d a y s o f 2 0 ° + w e a t h e r i n 1982. The summer o f 1982 was v e r y c o o l d u r i n g J u l y , mean maximum of 19.2°, and Aug u s t , mean maximum of 16.4° ( T a b l e 9), a f t e r two weeks o f v e r y warm w e a t h e r e n d e d on 1 J u l y . A l l t e m p e r a t u r e s l i s t e d w e r e r e c o r d e d i n an o p e n i n g i n i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g . 64 Table 9. Monthly mean minimum and mean maximum a i r temperatures (°C) Month FR DL IU OC RC May 5.5 a 2.7 a 4.1 4.2 5.6 16.2 a 18.4 a 17.0 16.1 17.4 June 4.8 2.6 3.7 3.5 5.5 15.1 17.4 15.6 15.2 16.4 July 9.2 7.2 7.8 7.1 10.2 20.7 24.1 20.4 21.3 22.7 August 9 - ° K 5.4 7.3 7.9 10.5 23.2 b 26.3 22.9 24.5 25.9 September 6.6 b 3.4 5.6 4.6 7.0 14.8 17.6 16.0 16.0 17.5 October 0.7 -0.7 -0.9 -0.9 0.4 7.0 9.2 8.0 7.7 8.9 November -1.9 -0.2 -2.7 -3.5 -3.0 2.0 2.1 3.2 2.2 2.3 December -10.5 -15.2 b -12.1 -12.6 b -10.1 -6.2 -7.1 b -5.4 -6.4 -5.1 January -18.9 b -21.0 b -21.0 -21.2 -18.3 -12.6 b -13.9 b -11.9 -13.0 -11.3 February -13.5 b -18.0 b -15.4 b -17.1 -12.9 -8.5 b -8.1 b -8.1 b -8.4 -7.3 March -9.3 b -8.2 b -9.6 -6.7 1.0b 1.9 b 1.9 b 1.1 1.9 A p r i l -4.7 -7.9 b -5.9 b -6.0 -3.8 4.5 6.3 b 5.8 b 5.0 6.5 May 1.3 -2.4 0.2 b 0.7 2.8 11.6 13.4 13. l b 12.2 13.7 Table 9 continued next page 65 T a b l e 9. c o n t i n u e d Year Month FR DL IU , OC RC June 7.6 4.1 7.0 10.0 22.5 25.7 22.9 b 23.3 24.1 July 9.4 7.2 8.7 8.7 10.1 19.0 21.2 19.2 20.2 20.3 August 7.1 5.8 6.7 6.3 7.7 15.9 19.2 16.4 17.2 17.7 September 6.2b 3.6 5.3 b 4.8 6.2 17.2 b 16.5 15.5 b 15.7 16.2 October 1.5 -0.4 0.8 0.2 4.6 6.5 8.6 7.9 7.3 8.0 November -7.4 -9.3 -8.1 -9.9 -7.7 -3.3 -2.8 -2.5 -3.8 -2.6 December -7.5 -11.5 -8.7 -9.3 -7.3 -3.3 -4.0 -3.1 -4.0 -3.4 January -6.4 -11.1 -7.0 -8.2 -5.4 -1.5 -2.8 -0.4 -1.4 -0.9 February -3.5 -6.8 -4.4 -4.8 -3.5 1.1 0.8 1.7 1.1 0.8 March -4.9 -8.5 -5.6 -6.1 -4.0 3.5 5.4 5.1 4.6 4.6 A p r i l -0.6 -2.6 -1.8 -1.8 0.6 10.0 12.0 11.5 10.2 11.8 May 5.5 2.2 3.6 b 3.7 6.2 17.0 18.9 17.2 b 16.7 17.8 1982 1983 FR, unlogged fo r e s t ; DL, diameter l i m i t logging; IU, intermediate u t i l i z a t i o n logging; 0C, old clea r c u t ; RC, recent clearcut a i n the data pairs, the mean minimum temperature i s l i s t e d on top and the mean maximum temperature i s on the bottom b contains regression estimates f o r missing data 66 The d a t a p r o d u c e d d u r i n g t h e summers o f 1980, 1981, and 1982, p r i m a r i l y t h e l a t t e r t wo, s u p p o r t t h e h y p o t h e s i s t h a t moose s e e k s h e l t e r when a i r t e m p e r a t u r e s e x c e e d 2 0 ° . Moose showed a s i g n i f i c a n t p o s i t i v e r e l a t i o n s h i p w i t h h a b i t a t w i t h h e a v y c o v e r i n t h e warm summer o f 1981 t h a t s h i f t e d t o h a b i t a t s w i t h l e s s c o v e r d u r i n g a c o o l summer. T h e s e d a t a a r e i n c o n s i s t e n t w i t h t h e h y p o t h e s i s t h a t moose s e l e c t h a b i t a t s w i t h abundant f o r a g e . Even i n t h e c o o l summer o f 1982, t h e p a t t e r n o f h a b i t a t o c c u p a t i o n d i d n o t f o l l o w t h e d i s t r i b u t i o n o f f o r a g e ( T a b l e 7 ) . I n 1982 t h e h a b i t a t w i t h t h e g r e a t e s t amount of f o r a g e had a s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n w i t h moose l o c a t i o n s , b u t o t h e r h e a v y f o r a g e p r o d u c i n g a r e a s had s i g n i f i c a n t n e g a t i v e a s s o c i a t i o n s . 3. F a l l T h e f a l l s e a s o n e x t e n d e d f r o m 15 S e p t e m b e r u n t i l t h e b e g i n n i n g o f t h e f a l l m i g r a t i o n e a c h y e a r . S e p t e m b e r 15 i s c l o s e t o t h e b e g i n n i n g o f t h e r u t w h i c h m a r k s a c h a n g e f r o m t h e summer p e r i o d i n w h i c h week t o week movements a r e s m a l l t o l a r g e r week t o week f a l l m o v e m e n t s . The f a l l m i g r a t i o n i s c h a r a c t e r i z e d by t h r e e c h a n g e s : f i r s t , w e e k l y m o v e m e n t s become r e l a t i v e l y l a r g e , up t o 20 km, and a r e d i r e c t e d t o w a r d t h e a r e a s i n w h i c h moose w i l l w i n t e r ; s e c o n d , t h e m i g r a t o r y movements c a u s e an u n m i s t a k e a b l e s h i f t t o more open h a b i t a t ; 67 t h i r d , m i g r a t o r y m o v e m e n t s r e s u l t i n m o o s e o c c u p y i n g l o w e r e l e v a t i o n s . T h e f a l l m i g r a t i o n s w a s f i r s t r e c o r d e d o n 1 D e c e m b e r , 1 N o v e m b e r a n d 16 N o v e m b e r i n t h e t h r e e y e a r s 1 9 8 0 t o 1 9 8 2 r e s p e c t i v e l y . A l l t h r e e f a l l p e r i o d s p r o d u c e d s i g n i f i c a n t l y n o n r a n d o m 2 m o o s e l o c a t i o n s w i t h r e s p e c t t o h a b i t a t , p ( X s d f ) < 0.1. T h e p a s s a g e f r o m s u m m e r t o f a l l r e s u l t e d i n i n c r e a s e d m o o s e u s e o f u n l o g g e d f o r e s t a s c o m p a r e d w i t h l o g g e d h a b i t a t s ( T a b l e 3 a n d F i g u r e 3 ) . I n 1 9 8 0 a n d 1 9 8 1 u n l o g g e d f o r e s t h a d s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n s w i t h m o o s e l o c a t i o n s , w h i l e a l l l o g g e d h a b i t a t s h a d s i g n i f i c a n t n e g a t i v e a s s o c i a t i o n s ( F i g u r e s 4 , 5 , 6 , a n d T a b l e 4 ) . I n 1 9 8 2 t h e a s s o c i a t i o n s b e t w e e n m o o s e a n d c l e a r c u t s r e m a i n e d s i g n i f i c a n t l y n e g a t i v e , w h i l e d i a m e t e r l i m i t l o g g i n g i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g , a n d u n l o g g e d f o r e s t h a d n o n s i g n i f i c a n t a s s o c i a t i o n s . I n a l l t h r e e f a l l p e r i o d s t h e c o r r e l a t i o n c o e f f i c i e n t b e t w e e n C o l e i n d i c e s a n d f o r e s t c o v e r w e r e m u c h g r e a t e r t h a n t h o s e b e t w e e n C o l e i n d i c e s a n d a v a i l a b l e f o r a g e ( T a b l e 7 ) . F o r e s t c o v e r s a r e l i s t e d i n T a b l e 6 a n d a v a i l a b l e f o r a g e e s t i m a t e s a r e i n T a b l e 1 0 . C l e a r l y , t h e r e w a s a s t r o n g r e l a t i o n s h i p b e t w e e n h a b i t a t s w i t h h i g h f o r e s t c o v e r v a l u e s a n d m o o s e l o c a t i o n s i n a l l t h r e e y e a r s m o o s e l o c a t i o n s w e r e g a t h e r e d . N e i t h e r o f t h e a l t e r n a t e h y p o t h e s e s c o u l d a c c o u n t f o r t h e r e l a t i o n s h i p b e t w e e n m o o s e a n d f o r e s t c o v e r i n t h e f a l l . H a b i t a t a s s o c i a t i o n s w e r e n o t 6 8 Table 10. F a l l a v a i l a b l e forage (g/nr) Sedges Habitat Deciduous Twigs Grasses and Rushes T o t a l A v a i l a b l e Forest 4.59 ( 0 . 5 2 5 ) 3 0.10 (0.116) 0.26 (0.216) 4 . 9 5 (0.780) Diameter L i m i t 11.64 (1.925) 0.09 (0.116) 0 . 0 ( 0 .0) 11.72 (1.955) Intermediate U t i l i z a t i o n 14.72 (1.438) 0.88 (0.159) 0.14 (0.217) 15.74 (1.563) Old Clearcut 11.62 (1.069) 1.24 (0.198) 0.88 (0.249) 13.74 (1.262) Recent C l e a r c u t 3.95 (0.560) 2.21 (0.206) 2.34 (0.304) 7.50 (0.908) Upland Shrub 28.08 (6.339) 0.11 (0.117) 0 . 0 ( 0 . 0 ) 28.18 (6.348) R i p a r i a n Shrub 78.83 (6.059) 1.44 (0.180) 0.96 (0.353) 81.23 (6.103) Wetland 2.95 (0.630) 0.07 (0.116) 6.36 (0.352) 9.30 (0.930) bracketed f i g u r e s are standard d e v i a t i o n s 69 c a u s e d by t h e d i s t r i b u t i o n o f f o r a g e n o r was t h e r e a p r o b l e m w i t h t h e r m a l s t r e s s . I n a r e c e n t c l e a r c u t , t h e h a b i t a t i n w h i c h one e x p e c t s t h e m o s t e x t r e m e c o n d i t i o n s , t h e mean maximum t e m p e r a t u r e e q u a l l e d 10.9° i n t h e f a l l o f 1981 and 8.6° i n 1982 ( T a b l e 8). These t e m p e r a t u r e s a r e much l e s s t h a n K n o r r e ' s ( 1 9 5 9 ) 2 0 ° g u i d e l i n e f o r h e a t s t r e s s . Nor d o e s c o l d s t r e s s a p p e a r t o h a v e b e e n a p r o b l e m w i t h mean mi n i m u m t e m p e r a t u r e s o f 2.2° and 2.9° i n t h e two y e a r s r e s p e c t i v e l y . The mean minimum t e m p e r a t u r e s a r e much warmer t h a n t h e - 2 0 ° g u i d e l i n e f o r c o l d s t r e s s . 4. E a r l y W i n t e r The e a r l y w i n t e r s e a s o n was d e f i n e d as t h e p e r i o d f r o m f a l l m i g r a t i o n u n t i l 15 J a n u a r y e a c h y e a r . Moose l o c a t i o n s were g a t h e r e d d u r i n g two e a r l y w i n t e r p e r i o d s , t h e w i n t e r s of 1981-1982 and 1982-1983. F a l l m i g r a t i o n was f i r s t o b s e r v e d on 1 N o vember 1981 and 16 N o v e m b e r 1982. The d i s t r i b u t i o n o f m o o s e l o c a t i o n s w i t h r e s p e c t t o h a b i t a t t y p e s was 2 s i g n i f i c a n t l y nonrandom, p ( X 5 d f ) < 0.1, i n b o t h y e a r s . Moose l o c a t i o n s showed s i m i l a r t r e n d s f r o m f a l l t o e a r l y w i n t e r i n b o t h y e a r s b u t t h e i n t e n s i t y o f c h a n g e s d i f f e r e d . In 1982-1983 t h e r e was a l a r g e f a l l w i n t e r s h i f t o f moose f r o m u n l o g g e d f o r e s t and d i a m e t e r l i m i t l o g g i n g i n t o more open h a b i t a t s . I n 1 981-1982 moose movement t o c l e a r c u t s and 70 i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g f r o m u n l o g g e d f o r e s t was much l e s s t h a n i n 1982-1983 ( F i g u r e s 5 and 6, T a b l e s 3 and 4). T h r e e f o r e s t / 1 o g g e d h a b i t a t s w e r e c o n s i s t e n t i n t h e i r moose a s s o c i a t i o n s i n e a r l y w i n t e r 1981-1982 and 1 9 8 2 - 1 9 8 3 . U n l o g g e d f o r e s t had n e g a t i v e a s s o c i a t i o n s , i n t e r m e d i a t e u t i l i z a t i o n p o s i t i v e a s s o c i a t i o n s , and r e c e n t c l e a r c u t s n e u t r a l a s s o c i a t i o n s i n b o t h y e a r s . D i a m e t e r l i m i t l o g g i n g went f r o m a n e u t r a l a s s o c i a t i o n i n 1981-1982 t o a n e g a t i v e a s s o c i a t i o n i n 1982-1983. O l d c l e a r c u t s went f r o m n e u t r a l t o a p o s i t i v e a s s o c i a t i o n e a r l y w i n t e r 1981-1982 t o e a r l y w i n t e r 1982-1983. Of t h e two c o m p e t i n g a l t e r n a t e h y p o t h e s e s , t h e h y p o t h e s i s t h a t moose s e e k h a b i t a t s w i t h t h e g r e a t e s t p r o d u c t i o n o f f o r a g e came c l o s e r t o e x p l a i n i n g moose h a b i t a t a s s o c i a t i o n s i n e a r l y w i n t e r . H o w e v e r , t h e r e l a t i o n s h i p b e t w e e n moose and h a b i t a t s w i t h abundant f o r a g e was not s t r o n g . The c o r r e l a t i o n b e t w e e n a v a i l a b l e f o r a g e and moose h a b i t a t a s s o c i a t i o n s was h i g h e r t h a n t h a t between f o r e s t c o v e r and h a b i t a t a s s o c i a t i o n s i n 1982-1983. However, moose h a b i t a t a s s o c i a t i o n s were more c l o s e l y r e l a t e d t o f o r e s t c o v e r t h a n a v a i l a b l e f o r a g e i n 1981-1982 ( T a b l e 7 ) . A v a i l a b l e f o r a g e i s l i s t e d i n T a b l e 11 and f o r e s t c o v e r i s i n T a b l e 6. Snow d e p t h s h o u l d n o t h a v e a f f e c t e d m o o s e h a b i t a t a s s o c i a t i o n s i n e a r l y w i n t e r a s snow d e p t h s w e r e much l e s s t h a n t h e c r i t i c a l 90 cm d e p t h d u r i n g b o t h e a r l y w i n t e r p e r i o d s 71 Table 11. Early winter a v a i l a b l e forage (g/m ) T a l l Low Deciduous Deciduous T o t a l Habitat Twigs Twigs Available Forest 0.20 (0.415) 3 0.67 (0.375) 0.87 (0.271) Diameter Limit 3.86 (1.293) 0.13 (0.426) 3.99 (0.683) Intermediate U t i l i z a t i o n 6.98 (1.026) 1.11 (0.455) 8.08 (0.915) Old Clearcut 10.39 (1.644) 1.53 (0.492) 11.92 (1.702) Recent Clearcut 1.97 (0.828) 0.97 (0.461) 2.94 (0.464) a bracketed figures are standard deviations 72 ( F i g u r e 7, F i g u r e 8, T a b l e 12, and T a b l e 13). M o ose may h a v e s o u g h t s h e l t e r f r o m c o l d s t r e s s i n t h e l a t t e r p a r t o f t h e e a r l y w i n t e r of 1981-1982, p o s s i b l y e x p l a i n i n g t h e d i f f e r e n c e s i n t h e p r o p o r t i o n o f moose l o c a t i o n s i n t h e u n l o g g e d f o r e s t and d i a m e t e r l i m i t l o g g i n g between t h e two y e a r s . F i f t y - t h r e e p e r c e n t o f moose l o c a t i o n s w e r e i n u n l o g g e d f o r e s t and d i a m e t e r l i m i t l o g g i n g i n e a r l y w i n t e r 1981-1982 as compared w i t h 24% i n 1 9 8 2 - 1 9 8 3 ( T a b l e 3 ) . T h e r e was a c o l d p e r i o d , m i n i m u m t e m p e r a t u r e s b e l o w - 2 0 ° i n an o l d c l e a r c u t f o r 15 days, i n l a t e December 1981 and e a r l y J a n u a r y 1982. I n e a r l y w i n t e r 1982-1983 t h e r e were a t o t a l o f 3 days i n w h i c h minimum t e m p e r a t u r e s were below - 2 0 ° C i n t h e o l d c l e a r c u t . 5. M i d w i n t e r M i d w i n t e r e x t e n d e d f r o m 16 J a n u a r y t o 15 M a r c h each y e a r . M o o se l o c a t i o n s w e r e r e c o r d e d d u r i n g two m i d w i n t e r p e r i o d s 1982 a n d 1 9 8 3 . I n e a c h o f t h e m i d w i n t e r p e r i o d s t h e d i s t r i b u t i o n o f moose l o c a t i o n s w i t h r e s p e c t t o h a b i t a t t y p e s 2 was s i g n i f i c a n t l y nonrandom, p ( X 5 d f ) < 0.1. The p a t t e r n o f a s s o c i a t i o n b e t w e e n moose l o c a t i o n s and h a b i t a t a p p e a r q u i t e d i f f e r e n t between the two y e a r s . Moose had a s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n w i t h f o r e s t and a s i g n i f i c a n t n e g a t i v e a s s o c i a t i o n w i t h r e c e n t c l e a r c u t s i n b o t h y e a r s ( F i g u r e 5, F i g u r e 6 a n d T a b l e 4 ) . T h e r e was a 73 F i g u r e 7. Mean snow d e p t h s i n f o r e s t / l o g g e d h a b i t a t s d u r i n g t h e w i n t e r o f 1981-1982 180 Nov Dec Jan Feb Mar Apr May 7 5 F i g u r e 8. Mean snow d e p t h s i n f o r e s t / l o g g e d h a b i t a t s d u r i n g t h e w i n t e r of 1982-1983 100 -T 90-Nov Dec Jan Feb Mar Apr May 77 Table 12. Mean snow depths i n forest/logged habitats during the winter of 1981-1982 Range of Dates FR DL IU OC RC early winter 20 Nov 0 ( 0.0) a 0 ( 0.0) 0 ( 0.0) 0 ( 0.0) 0 (0.0) 29 Nov - 3 Dec 22 ( 2.2) 65 ( 3.7) 29 ( 3.6) - 32 (2.6) 30 Dec - 6 Jan 44 ( 6.2) 75 ( 7.1) 56 ( 7.4) 52 ( 3.6) 65 (6.5) midwinter 29 Jan - 31 Jan 143 (12.1) - 143 (21.1) 147 ( 9.5) -20 Feb - 21 Feb 136 ( 8.8) 175 (10.8) 149 (14.3) 148 ( 6.2) 152 (13.7) late winter 14 Mar - 23 Mar 136 (13.4) 152 (14.2) 145 (17.4) 134 (14.6) 155 (14.4) 6 Apr 104 ( 7.2) - 133 (14.9) - 130 (14.3) 20 Apr - 29 Apr 93 ( 9.2) 104 (11.6) 89 (17.0) 71 ( 6.0) 110 (10.7) 8 May 85 ( 7.6) - 33 (21.4) 10 (11.1) 49 (13.4) 29 May - 30 May 1 ( 2.0) 8 ( 8.8) 0 ( 0,0) 0 ( 0.0) 0 ( 0.0) FR, unlogged forest; DL, diameter l i m i t logging; IU, intermediate ut i l i z a t i o n logging; 0C, old clearcut; RC, recent clearcut figure i n brackets i s the standard deviation no data recorded during the stated period 78 Table 13. Mean snow depths i n forest/logged habitats during the winter of 1982-1983 Range of Dates FR DL IU OC RC early winter 4 Nov 0 ( 0.0) a 0 ( 0.0) 0 ( 0.0) 0 ( 0.0) 0 ( 0.0) 16 Nov - 23 Nov 10 ( 1.3) 17 ( 4.8) 17 ( 5.5) 18 ( 2.5) 21 ( 2.6) 3 Dec - 6 Dec 27 ( 5.2) 38 ( 8.8) 39 ( 9.7) 43 ( 4.9) 38 ( 5.2) 17 Dec - 22 Dec 41 ( 5.4) 62 ( 8.4) 51 (12.4) 61 ( 5.7) 58 ( 5.0) 7 Jan - 9 Jan 60 ( 8.0) 78 ( l l . D 79 (13.9) 79 ( 6.3) 70 ( 7.9) midwinter 20 Jan - 23 Jan 56 ( 8.4) 75 (12.9) 73 (13.8) 65 ( 8.6) 64 ( 8.9) 3 Feb - 5 Feb 60 ( 9.3) 80 (13.2) 74 (15.4) 67 ( 6.5) 69 ( 6.3) 21 Feb - 22 Feb 62 (11.5) 83 (11.0) 76 (17.5) 68 ( 8.0) 70 ( 7.9) 3 Mar - 8 Mar 58 (10.2) 75 (12.3) 67 (15.8) 56 (10.1) 57 ( 7.2) late winter 18 Mar 47 ( 9.2> 59 ( 8.7) 51 (18.2) 32 ( 8.1) 40 (11.2) 1 Apr 39 (11.3) 50 (10.4) 36 (18.6) 18 (10.0) 27 (14.0) 14 Apr - 15 Apr 25 (11.0) 31 (13.6) 17 (12.3) 0 ( 0.0) 2 ( 6.1) 28 Apr 0 ( 0.0) 1 ( 3.0) 0 ( 0.0) 0 ( 0.0) 0 ( 0.0) FR, unlogged forest; DL, diameter l i m i t logging; IU, intermediate u t i l i z a t i o n logging; 0C, old clearcut; RC, recent clearcut a figure i n brackets i s the standard deviation 79 s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n between moose and d i a m e t e r l i m i t l o g g i n g i n 1982 but a s i g n i f i c a n t n e g a t i v e a s s o c i a t i o n i n 1 9 8 3 . M o o s e w e r e n o n s i g n i f i c a n t 1 y a s s o c i a t e d w i t h i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g i n 1982 b u t p o s i t i v e l y a s s o c i a t e d i n 1983. O l d c l e a r c u t s went f r o m a s i g n i f i c a n t n e g a t i v e a s s o c i a t i o n w i t h m o o s e t o a n o n s i g n i f i c a n t a s s o c i a t i o n . C o l e i n d i c e s w e r e more h i g h l y c o r r e l a t e d w i t h f o r e s t c o v e r t h a n w i t h a v a i l a b l e f o r a g e i n b o t h m i d w i n t e r s ( T a b l e 7). F o r e s t c o v e r e s t i m a t e s a r e i n T a b l e 6. A v a i l a b l e f o r a g e f o r t h e m i d w i n t e r s e a s o n s a r e l i s t e d i n T a b l e 14. The moose b e h a v i o u r d a t a i n t h e m i d w i n t e r p e r i o d d i d not d i s t i n g u i s h b e t w e e n t h e a l t e r n a t e h y p o t h e s e s . Snow d e p t h g r e a t l y e x c e e d e d t h e 90 cm c r i t i c a l d e p t h i n a l l h a b i t a t s i n 1982 ( F i g u r e 7 and T a b l e 12) and moose h a b i t a t a s s o c i a t i o n s s u p p o r t t h e h y p o t h e s i s t h a t i n t h e s e c o n d i t i o n s moose occupy h a b i t a t s w i t h t h e g r e a t e s t f o r e s t c o v e r . M i d w i n t e r 1982 was a l s o c o l d . I n an o l d c l e a r c u t t h e r e w e r e 21 d a y s i n m i d w i n t e r 1982 when t h e t e m p e r a t u r e s f e l l below t h e - 2 0 ° g u i d e l i n e . I t i s p o s s i b l e t h a t moose w e r e u n d e r h e a v y c o v e r t o a v o i d c o l d s t r e s s . M o o se h a b i t a t a s s o c i a t i o n s d u r i n g m i d w i n t e r 1982 c a n n o t be used t o s o r t out w h e t h e r deep snow or c o l d d e t e r m i n e moose m i d w i n t e r h a b i t a t a s s o c i a t i o n s b e c a u s e b o t h o c c u r r e d . N e i t h e r f o r a g e p r o d u c t i o n c o n d i t i o n a l upon snow d e p t h nor c o l d s t r e s s c o u l d h a v e p r e d i c t e d t h e p a t t e r n o f h a b i t a t a s s o c i a t i o n i n m i d w i n t e r 1983. I n 1983 snow d e p t h d i d n o t 80 Table 14. Midwinter available forage (g/m ) 1981 - 1982 Habitat T a l l Deciduous Twigs Low Deciduous Twigs Abies Total Available Forest 0.03 (0.389) 3 0.01 (0.370) 2.06 ( 4.628) 2.10 (0.330) Diameter Limit 1.15 (0.651) 0.0 (0.0) 1.62 ( 4.119) 2.7 (0.315) Intermediate Uti l i z a t i o n 2.11 (0.587) 0.01 (0.410) 3.06 ( 7.208) 5.18 (0.543) Old Clearcut 3.52 (0.961) 0.0 (0.0) 7.30 (13.857) 10.81 (1.289) Recent Clearcut 0.15 (0.417) 0.0 (0.0) 1982 - 1983 1.01 (16.215) 1.16 (0.284) Forest 0.14 (0.415) 0.28 (0.374) 5.92 ( 9.822) 6.33 (0.588) Diameter Limit 3.53 (1.226) 0.02 (0.426) 6.81 (15.189) 10.35 (0.928) Intermediate Utilization 6.39 (1.005) 0.73 (0.439) 8.07 (17.171) 15.19 (1.035) Old Clearcut 9.10 (1.587) 0.93 (0.477) 16.62 (30.366) 36.65 (2.880) Recent Clearcut 1.10 (0.703) 0.08 (0.433) 2.93 (33.804) 4.11 (0.724) bracketed figures are standard deviations 81 e x c e e d 9 0 cm i n a n y h a b i t a t a t a n y t i m e ( F i g u r e 8 a n d T a b l e 1 3 ) a n d m i d w i n t e r 1 9 8 3 w a s w a r m . T h e m e a n m a x i m u m t e m p e r a t u r e w a s 1 . 0 ° i n a r e c e n t c l e a r c u t , a n d t h e m e a n m i n i m u m w a s - 4 . 4 ° ( T a b l e 8 ) . T h e m i n i m u m t e m p e r a t u r e w a s b e l o w - 2 0 ° o n o n e d a y . G i v e n t h e a l t e r n a t i v e s b e i n g d i s c u s s e d i n t h i s t h e s i s o n e w o u l d e x p e c t t o f i n d m o o s e i n h a b i t a t w i t h a b u n d a n t f o r a g e a n d n o t i n h a b i t a t c h a r a c t e r i z e d b y h i g h f o r e s t c o v e r v a l u e s . F o r e s t c o v e r , h o w e v e r , w a s m o r e h i g h l y c o r r e l a t e d t h a n f o r a g e a b u n d a n c e w i t h m o o s e h a b i t a t a s s o c i a t i o n s i n m i d w i n t e r 1 9 8 3 ( T a b l e 7 ) . S i n c e t h e r e w a s a n o n r a n d o m d i s t r i b u t i o n o f m o o s e w i t h r e s p e c t t o h a b i t a t i n m i d w i n t e r 1 9 8 3 , i n d i c a t i n g t h a t m o o s e w e r e s e l e c t i n g h a b i t a t , a f a c t o r o t h e r t h a n t h e e s c a p e o f d e e p s n o w , a t t r a c t i o n o f a b u n d a n t f o r a g e , o r a v o i d a n c e c o l d s t r e s s m u s t be i n v o l v e d . 6 . L a t e W i n t e r L a t e w i n t e r m o o s e l o c a t i o n s w e r e g a t h e r e d d u r i n g 1 9 8 2 a n d 1 9 8 3 . T h e l a t e w i n t e r p e r i o d s t a r t e d o n 16 M a r c h a n d e n d e d w i t h t h e o n s e t o f s p r i n g m i g r a t i o n . T h e f i r s t i n d i c a t i o n s o f s p r i n g m i g r a t i o n w e r e o n 15 M a y 1 9 8 2 a n d 22 A p r i l 1 9 8 3 . I n 1 9 8 2 t h e d i s t r i b u t i o n o f m o o s e l o c a t i o n s w i t h 2 r e s p e c t t o h a b i t a t t y p e s w a s s i g n i f i c a n t l y n o n r a n d o m P ( X 5 d f ) < 0 . 1 . I n 1 9 8 3 t h e d i s t r i b u t i o n o f m o o s e l o c a t i o n s w a s m a r g i n a l l y n o n s i g n i f i c a n t ( p = 0 . 1 1 1 ) i n a t e s t o f r a n d o m n e s s . 82 A s i m i l a r t r e n d m i d w i n t e r t o l a t e w i n t e r d e v e l o p e d i n b o t h y e a r s w i t h i n c r e a s e s i n t h e u s e o f c l e a r c u t s i n l a t e w i n t e r b u t w i t h s m a l l d i f f e r e n c e s i n t h e u s e o f t h e o t h e r t h r e e h a b i t a t s ( T a b l e 3, T a b l e 4, F i g u r e 5, and F i g u r e 6 ) . F o r e s t c o v e r was much more h i g h l y c o r r e l a t e d w i t h C o l e i n d i c e s t h a n was a v a i l a b l e f o r a g e ( T a b l e 7). L a t e w i n t e r a v a i l a b l e f o r a g e i s l i s t e d i n T a b l e 15. N e i t h e r t h e h y p o t h e s i s t h a t moose s h o u l d s e e k h a b i t a t s w i t h l a r g e amounts of f o r a g e t e m p e r e d by t h e c o n s i d e r a t i o n of a 90 cm c r i t i c a l snow d e p t h nor t h e h y p o t h e s i s t h a t moose seek s h e l t e r t o a v o i d c o l d s t r e s s a p p e a r t o e x p l a i n t h e p a t t e r n of moose h a b i t a t o c c u p a t i o n i n l a t e w i n t e r of e i t h e r y e a r . L a t e w i n t e r p e r i o d s were not c o l d , t h e mean minimum t e m p e r a t u r e i n a r e c e n t c l e a r c u t was - 3 . 0 ° i n 1982 and - 2 . 5 ° i n 1983 ( T a b l e 8 ) . T e m p e r a t u r e s d i d n o t d r o p b e l o w - 2 0 ° i n e i t h e r y e a r . Snow d e p t h s w e r e w e l l b e l o w t h e 90 cm c r i t i c a l d e p t h i n a l l h a b i t a t s i n 1 9 8 3 ( F i g u r e 8 a n d T a b l e 1 3 ) . I n 1 9 8 2 t h e s h a l l o w e s t snow was on an o l d c l e a r c u t , a l s o t h e h a b i t a t t y p e w i t h t h e l a r g e s t a m ount o f a v a i l a b l e b r o w s e ( T a b l e 1 5 ) , and t h e snow on t h a t s i t e was l e s s t h a n 90 cm f o r more t h a n h a l f t h e l a t e w i n t e r p e r i o d ( F i g u r e 7 and T a b l e 11). N e i t h e r snow d e p t h n o r c o l d s t r e s s c a n a c c o u n t f o r t h e a v o i d a n c e o f h e a v y b r o w s e b e a r i n g h a b i t a t and t h e o c c u p a t i o n o f h a b i t a t w i t h l i t t l e browse d u r i n g l a t e w i n t e r . 83 Table 15. Late winter available forage (g/m ) 1981 - 1982 Habitat T a l l Deciduous Twigs Low Deciduous Twigs Abies Total Available Forest 0.07 (0.412)a 0.02 (0.370) 4.24 ( 8.296) 4.32 (0.486) Diameter Limit 2.58 (1.024) 0.0 (0.0) 4.48 (10.750) .06 (0.636) Intermediate Utili z a t i o n 4.23 (0.868) 0.05 (0.412) 5.63 (12.081) .91 (0.820) Old Clearcut 5.56 (1.233) 0.00 (0.451) 15.34 (21.198) 20.90 (1.972) Recent Clearcut 0.53 (0.565) 0.00 (0.428) 1982 - 1983 1.79 (26.948) 2.30 (0.495) Forest 0.34 (0.434) 1.35 (0.387) 9.21 (12.903) 10.9 (0.808) Diameter Limit 4.73 (1.427) 0.58 (0.428) 0.63 (19.908) 15.24 (1.211) Intermediate Utili z a t i o n 9.17 (1.113) 2.44 (0.545) 11.80 (20.100) 23.41 (1.269) Old Clearcut 13.57 (1.728) 3.27 (0.551) 36.93 (32.060) 53.77 (3.428) Recent Clearcut 2.79 (0.948) 1.76 (0.521) 6.22 (37.457) 10.77 (1.373) a bracketed figures are standard deviations 84 7.0 Summary The a n n u a l p a t t e r n o f moose h a b i t a t o c c u p a t i o n i n n o r t h c e n t r a l B r i t i s h C o l u m b i a , f r o m A p r i l 1980 t o May 1983, was w i n t e r t o s p r i n g movement t o open h a b i t a t s ; i n summer, moose w e r e u s u a l l y a s s o c i a t e d w i t h h a b i t a t s b e a r i n g h e a v y f o r e s t c o v e r ; i n t h e f a l l t h e a s s o c i a t i o n w i t h f o r e s t c o v e r was even g r e a t e r t h a n i t was i n summer; w i t h t h e o n s e t of m i g r a t i o n i n e a r l y w i n t e r , moose a g a i n moved i n t o more o p e n a r e a s ; t h e y r e t r e a t e d t o h e a v i e r f o r e s t c o v e r i n m i d w i n t e r ; i n l a t e w i n t e r t h e r e was a l i t t l e movement t o open a r e a s ; and t h e movement t o open a r e a s became g e n e r a l w i t h t h e s p r i n g m i g r a t i o n ( F i g u r e 3 and T a b l e 3 ) . The i n t e n s i t y o f t h e s e a s o n t o s e a s o n c h a n g e s i n h a b i t a t o c c u p a t i o n v a r i e d f r o m y e a r t o y e a r . M o s t o f t h e h a b i t a t c h a n g e s i n v o l v e d moose m o v e m e n t s b e t w e e n u n l o g g e d f o r e s t and i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g . T h e s e two h a b i t a t t y p e s r a n k e d f i r s t and s e c o n d i n t e r m s o f l a n d a r e a . Moose a p p e a r t o s e l e c t h a b i t a t as t h e r e were s i g n i f i c a n t nonrandom d i s t r i b u t i o n s o f moose w i t h r e s p e c t t o h a b i t a t t y p e , 2 p ( X 5 d f ) < 0.1, e x c e p t i n two s p r i n g p e r i o d s , one summer, and one l a t e w i n t e r p e r i o d . I n t h e l a t e w i n t e r p e r i o d w h i c h f a i l e d t o r e c o r d a s i g n i f i c a n t n o n r a n d o m d i s t r i b u t i o n , t h e r e s u l t was m a r g i n a l l y n o n s i g n i f i c a n t . 8 5 7 . 1 R e l e v a n c e o f W o r k i n g H y p o t h e s e s N e i t h e r o f t h e a l t e r n a t e h y p o t h e s e s c o u l d e x p l a i n t h e p a t t e r n o f m o o s e / h a b i t a t a s s o c i a t i o n s r e c o r d e d i n t h i s t h e s i s . M o o s e a v o i d a n c e o f t h e r m a l s t r e s s b y s e e k i n g h a b i t a t s w i t h h e a v y f o r e s t c o v e r i n s u m m e r ( t e m p e r a t u r e s i n e x c e s s o f 2 0 ° ) a n d w i n t e r ( t e m p e r a t u r e s l e s s t h a n - 2 0 ° ) a c c o u n t e d f o r m o o s e h a b i t a t a s s o c i a t i o n s i n s u m m e r a n d i n m i d w i n t e r 1 9 8 2 . S n o w w a s v e r y d e e p i n m i d w i n t e r 1 9 8 2 , h o w e v e r , a n d m o o s e a s s o c i a t i o n s w i t h h e a v y f o r e s t c o v e r s i n t h a t s e a s o n c o u l d h a v e b e e n d u e t o t h e i r a v o i d i n g d e e p s n o w . A g a i n i n m i d w i n t e r 1 9 8 3 , m o o s e s o u g h t s h e l t e r . N o t e t h a t m i d w i n t e r 1 9 8 3 w a s w a r m . T h e d i r e c t i o n o f h a b i t a t a s s o c i a t i o n s d u r i n g s p r i n g a n d e a r l y w i n t e r p e r i o d s w a s i n a c c o r d a n c e w i t h t h e h y p o t h e s i s t h a t m o o s e w o u l d s e e k h a b i t a t w i t h a b u n d a n t f o r a g e . T w o o f f o u r s p r i n g p e r i o d s f a i l e d t o p r o d u c e a n o n r a n d o m d i s t r i b u t i o n o f m o o s e w i t h r e s p e c t t o h a b i t a t t y p e s , h o w e v e r , a n d o n l y t h r e e o f s i x s p r i n g a n d e a r l y w i n t e r p e r i o d s p r o d u c e d h i g h c o r r e l a t i o n s b e t w e e n a v a i l a b l e f o r a g e a n d C o l e a s s o c i a t i o n i n d i c e s ( T a b l e 7 ) . N e i t h e r a l t e r n a t e h y p o t h e s i s c o u l d h a v e p r e d i c t e d t h e m o o s e h a b i t a t a s s o c i a t i o n s w h i c h o c c u r r e d i n f a l l a n d l a t e w i n t e r . I n t h r e e f a l l s , m o o s e w e r e a l w a y s a s s o c i a t e d w i t h u n l o g g e d f o r e s t t h a t h a s a h e a v y f o r e s t c o v e r , 0 . 7 7 2 o f t h e s k y h e m i s p h e r e o b s c u r e d by v e g e t a t i o n , a n d l o w a v a i l a b l e f o r a g e , 4 . 9 5 g / m . W h i l e o l d c l e a r c u t s w i t h 8 6 l i g h t f o r e s t c o v e r , 0 . 0 6 0 o f t h e s k y h e m i s p h e r e , a n d l a r g e r a m o u n t s o f a v a i l a b l e f o r a g e , 1 3 . 7 4 g/m , w e r e a l w a y s a v o i d e d . T h e r e w a s n o a p p a r e n t t h e r m a l s t r e s s w h i c h c o u l d h a v e a c c o u n t e d f o r t h i s b e h a v i o u r . I n l a t e w i n t e r 1 9 8 2 a n d 1 9 8 3 , m o o s e l o c a t i o n s p r o d u c e d c o r r e l a t i o n c o e f f i c i e n t s b e t w e e n h a b i t a t a s s o c i a t i o n i n d i c e s a n d f o r e s t c o v e r t h a t w e r e l a r g e r t h a n t h o s e b e t w e e n a v a i l a b l e f o r a g e r a n k i n g s a n d h a b i t a t a s s o c i a t i o n s . C o l d s t r e s s c o u l d n o t a c c o u n t f o r t h e o b s e r v e d h a b i t a t a s s o c i a t i o n s ; b o t h l a t e w i n t e r s e a s o n s w e r e w a r m . 87 V DISCUSSION 1. F o r a g e P r o d u c t i o n C o n d i t i o n a l Upon Snow Depth D e s p i t e t h e f o r a g e p r o d u c t i o n h y p o t h e s i s ' i n a b i l i t y t o e x p l a i n t h e p a t t e r n of moose h a b i t a t a s s o c i a t i o n , one s h o u l d n o t d i s c a r d t h e i d e a t h a t f o r a g e p r o d u c t i o n i s an i m p o r t a n t c o n s i d e r a t i o n i n moose h a b i t a t . F o r a g e a t t r i b u t e s , o t h e r t h a n t o t a l p r o d u c t i o n , m i g h t h a v e e x p l a i n e d m o o s e h a b i t a t s e l e c t i o n . F o r e x a m p l e , Peek e t a l . ( 1 9 7 6 ) d o c u m e n t e d a s i m i l a r p a t t e r n o f moose h a b i t a t s e l e c t i o n t o t h a t i n t h i s s t u d y and t h e y m a i n t a i n e d t h a t o p e n a r e a s w e r e c r i t i c a l t o moose b e c a u s e o f f o r a g e r e q u i r e m e n t s . I n t h e i r ( P e e k e t a l . 1976) o p i n i o n , heavy moose use o f open a r e a s i n s p r i n g was due t o t h e h i g h p r o t e i n l e v e l s o f p l a n t s i n t h o s e a r e a s . T h e y s u g g e s t e d t h a t s h i f t s to h a b i t a t s w i t h g r e a t e r t r e e c o v e r i n summer w e r e due t o a d e c r e a s e i n p a l a t a b i l i t y o f p l a n t s i n open a r e a s . F o r e s t s a t t r a c t moose b e c a u s e p l a n t s m a t u r e s l o w e r and r e t a i n n u t r i e n t s l o n g e r . T h e r e i s i n d e p e n d e n t e v i d e n c e f o r t h i s a s s e r t i o n . T r e m b l a y ( 1 9 8 1 ) , w o r k i n g i n w h i t e - t a i l e d d e e r h a b i t a t , d o c u m e n t e d c r u d e p r o t e i n v a l u e s h i g h e r i n open h a b i t a t s i n s p r i n g b u t l o w e r t h a n i n c l o s e d h a b i t a t s i n summer. A l s o , B l a i r e t a l . ( 1 9 8 3 ) s h o w e d t h a t c r u d e p r o t e i n l e v e l s i n d e e r b r o w s e w e r e h i g h e r i n s h a d e d t h a n i n unshaded a r e a s . 88 F u r t h e r , P eek e t a l . ( 1 9 7 6 ) f o u n d t h a t l a t e f a l l e a r l y w i n t e r s e l e c t i o n of open a r e a s was c o r r e l a t e d w i t h h i g h e t h e r e x t r a c t a b l e p r o d u c t s and n i t r o g e n f r e e e x t r a c t s i n f o r a g e of o p e n a r e a s . W i n t e r u s e o f a r e a s w i t h d e n s e g r o w t h s o f t r e e s was dependent on t h e s e v e r i t y of the w i n t e r w i t h g r e a t e r use of open a r e a s d u r i n g m i l d w i n t e r s . Peek e t a l . (1976) a r g u e d t h a t moose r e q u i r e t h e p r o t e i n c o n t e n t i n t h e f o r a g e i n o p e n a r e a s i n s p r i n g b e c a u s e moose a r e g r o w i n g r a p i d l y a t t h a t t i m e and t h e h i g h l e v e l s of e t h e r and n i t r o g e n f r e e e x t r a c t a r e r e q u i r e d i n e a r l y w i n t e r t o r e c o v e r f a t r e s e r v e s f o l l o w i n g t h e r u t . Peek e t a l . ( 1 9 7 6 ) c o n c l u d e d t h a t t h e open a r e a s were i m p o r t a n t enough t o s t a t e ". . . t h e open c o v e r s u s e d i n l a t e f a l l and e a r l y w i n t e r , and a g a i n i n s p r i n g , must be t h e m a j o r h a b i t a t s s u s t a i n i n g t h i s p o p u l a t i o n . . . ". When a l l s e a s o n s w e r e c o n s i d e r e d , t h e moose p r o b a b l y showed no more a f f i n i t y f o r open h a b i t a t s t h a n t h o s e i n t h e p r e s e n t s t u d y . Y e t , Peek e t a l . ( 1 9 7 6 ) h a v e a r g u e d , not on t h e b a s i s of b i o m a s s of f o r a g e p l a n t s p r o d u c e d , b u t on t h e b a s i s o f t h e a b u n d a n c e o f p a r t i c u l a r n u t r i e n t s , t h a t open h a b i t a t s a r e c r i t i c a l t o t h e m a i n t e n a n c e of a l a r g e moose p o p u l a t i o n . I n t h e p r e s e n t s t u d y , i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g c o n s i s t e n t l y had a h i g h a s s o c i a t i o n w i t h moose l o c a t i o n s as compared w i t h the o t h e r f o r e s t / l o g g e d h a b i t a t s . The r e s p o n s e of moose t o i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g may be based i n 89 p a r t on t h e a b u n d a n t g r o w t h o f f o r a g e i n t h a t h a b i t a t . The o t h e r h a b i t a t o f t e n u s e d by moose, u n l o g g e d f o r e s t , had t h e l o w e s t f o r a g e a b u n d a n c e i n a l l s e a s o n s e x c e p t m i d - and l a t e w i n t e r when t h e b i o m a s s o f f o r a g e i n u n l o g g e d f o r e s t was m a r g i n a l l y g r e a t e r t h a n t h a t o f r e c e n t c l e a r c u t s . The g e n e r a l l y l o w l e v e l s o f a v a i l a b l e f o r a g e i n u n l o g g e d f o r e s t d o e s n o t mean t h a t moose w e r e u n a b l e t o o b t a i n s u f f i c i e n t f o r a g e t o s a t i s f y t h e i r n e e d s w i t h i n f o r e s t . F o r e s t s , p a r t i c u l a r l y o l d g r o w t h f o r e s t s , h a v e n u m e r o u s o p e n i n g s and wet a r e a s w h i c h c a n s u p p o r t l a r g e a m o u n t s o f f o r a g e . F o r e x a m p l e , on t h e m o r n i n g o f 27 A u g u s t 1982, a moose c r o s s e d one of t h e f o r e s t p l o t s e s t a b l i s h e d f o r t h e p r e s e n t s t u d y . W i t h i n a d i s t a n c e o f 30 m t h i s a n i m a l had p r o b a b l y s t r i p p e d e n o u g h d e c i d u o u s l e a v e s t o f i l l i t s rumen, y e t t h e o v e r a l l p r o d u c t i o n of t h a t s i t e was low. I f moose were g e n e r a l l y a b l e t o o b t a i n s u f f i c i e n t f o r a g e w i t h i n u n l o g g e d f o r e s t , t h e n t h e l a c k o f h i g h c o r r e l a t i o n s between moose a s s o c i a t i o n s and h a b i t a t w i t h a b u n d a n t f o r a g e ( T a b l e 7) w o u l d be e x p e c t e d . The d e n s i t y o f moose on t h e s t u d y a r e a was a p p r o x i m a t e l y 0.3 moose/ha, making i t u n l i k e l y t h a t f o r a g e was l i m i t i n g i n any h a b i t a t f r o m s p r i n g t o e a r l y w i n t e r . The e x p e c t a t i o n t h a t snow d e p t h i s t h e m o s t i m p o r t a n t f a c t o r i n f l u e n c i n g w i n t e r h a b i t a t s e l e c t i o n h a s c r e a t e d a c o n t r o v e r s y as t o how much snow i s n e c e s s a r y t o f o r c e moose 90 u n d e r c o v e r . The c h e s t h e i g h t s o f a d u l t moose o f a medium s i z e d s u b s p e c i e s , A 1_ c es a l c e s a_n ££J_ s_ o n_i j i , r a n g e f r o m 100 t o 110 cm ( T e l f e r and K e l s a l l 1 9 7 9). Many a u t h o r s h a v e a s s u m e d t h a t snow d e p t h c o m p r i s i n g a s i g n i f i c a n t p r o p o r t i o n o f t h a t h e i g h t , 80 t o 90 cm, i s r e q u i r e d t o i n h i b i t moose m o b i l i t y and f o r c e s e l e c t i o n of h a b i t a t s w i t h s h a l l o w e r snow. F o r example, T e l f e r ( 1 9 7 0 ) , K r e f t i n g ( 1 9 7 4 a ) , and Peek e t a l . ( 1 9 8 2 ) c i t e 90 cm as t h e d e p t h o f snow w h i c h r e q u i r e s moose t o s e e k s h e l t e r i n f o r e s t s . However, R o l l e y and K e i t h (1980) o b s e r v e d moose w i n t e r h a b i t a t s h i f t s f r o m open t o f o r e s t e d a r e a s w h i c h were " p r o b a b l y t r i g g e r e d by g r e a t e r snow a c c u m u l a t i o n i n open a r e a s " , b u t t h e i r snow d e p t h s n e v e r e x c e e d e d 75 cm and wer e o f t e n much l e s s . M o n t h e y ( 1 9 8 4 ) o b s e r v e d s i m i l a r h a b i t a t s h i f t s and b e l i e v e d t h a t 70 cm o f c r u s t e d snow i n t h e o p e n a r e a s was r e s p o n s i b l e f o r moose o c c u p y i n g f o r e s t s i n m i d -w i n t e r . P h i l l i p s e t a l . ( 1 9 7 3 ) b e l i e v e d t h a t 51 cm o f snow r e p r e s e n t e d a c r i t i c a l d e p t h b e c a u s e t h e y o b s e r v e d h a b i t a t s h i f t s when t h a t d e p t h of snow was p r e s e n t . F i n a l l y , Peek e t a l . (1976) r e p o r t e d 45 cm of c r u s t e d snow f o r c i n g moose out of o p e n h a b i t a t s and i n t o f o r e s t s . I t i s p o s s i b l e t h a t t h i s c o n t r o v e r s y a r i s e s b e c a u s e a f a c t o r o t h e r t h a n a b s o l u t e snow de p t h may be i n f l u e n c i n g moose b e h a v i o u r i n w i n t e r . Snow h i n d e r s movement due t o t h e d e p t h moose h o o v e s p e n e t r a t e and t h e amount o f d r a g due t o d e n s i t y and h a r d n e s s ( C o a d y 1 9 7 4 ) . The 90 cm c r i t i c a l d e p t h , o r any c r i t i c a l 91 d e p t h , a s s u m e s t h a t t h e moose I s p e n e t r a t i n g t h r o u g h a s u b s t a n t i a l p r o p o r t i o n o f t h e snow d e p t h . F o r e x a m p l e , i n l a t e J a n u a r y and e a r l y F e b r u a r y moose were f l o u n d e r i n g i n snow w e l l o v e r 100 cm d e e p , snow o f t e n r e a c h i n g 30 cm a b o v e t h e l e v e l o f t h e i r c h e s t s . H o w e v e r , t h e d e e p e s t snow r e c o r d e d ( F i g u r e 7 and T a b l e 12) was i n d i a m e t e r l i m i t l o g g i n g - - a h a b i t a t w i t h w h i c h moose w e r e p o s i t i v e l y a s s o c i a t e d i n m i d -w i n t e r 1982 ( F i g u r e 5 and T a b l e 4 ) . I f moose w e r e a v o i d i n g deep snow, th e y s h o u l d have a v o i d e d d i a m e t e r l i m i t l o g g i n g i n m i d w i n t e r 1982, but th e y d i d n o t . I n l a t e w i n t e r 1982 moose d i d n o t p e n e t r a t e d e e p l y i n t o t h e snow p a c k . On 14 M a r c h 1982 t h e mean snow d e p t h on a r e c e n t c l e a r c u t was 155 cm. Y e t , a moose c r o s s i n g t h a t a r e a , a few h o u r s b e f o r e t h e snow measurements, p e n e t r a t e d a mean of 50 cm. T h a t same day I n a mature f o r e s t 400 m away, t h e mean snow d e p t h was 131 cm and a moose m o v i n g t h r o u g h t h a t a r e a p e n e t r a t e d 57 cm p e r s t e p on a v e r a g e . S i x d a y s l a t e r on an o l d c l e a r c u t , t h e mean snow d e p t h was 134 cm and mean moose p e n e t r a t i o n was 41 cm. I f moose p e n e t r a t e d e e p e r i n t o snow i n h a b i t a t s w i t h h e a v y f o r e s t c o v e r t h a n i n o p e n a r e a s , t h e n moose h a b i t a t a s s o c i a t i o n s a r e not c r e d i b l y e x p l a i n e d by snow d e p t h e x c e e d i n g 90 cm i n t h e open a r e a s . T h e r e must have been o t h e r o r a d d i t i o n a l c a u s e s o f moose a v o i d a n c e o f open a r e a s and a s s o c i a t i o n w i t h f o r e s t c o v e r i n m i d w i n t e r and l a t e w i n t e r 1982. 92 C r u s t i n g o f snow i s s o m e t i m e s c i t e d as t h e r e a s o n moose move out of open h a b i t a t s b e c a u s e c r u s t e d snow a b r a d e s moose l e g s and c r e a t e s a d d i t i o n a l d r a g . Snow i n o p e n h a b i t a t s , i n t h e p r e s e n t s t u d y , was h e a v i l y c r u s t e d but not h a r d enough t o s u p p o r t moose d u r i n g m i d w i n t e r 1983. The c r u s t i n g o f snow i n t h e f o r e s t was w o r s e t h a n i n o p e n a r e a s , h o w e v e r , and c o u l d h a v e been c a l l e d l a y e r s o f i c e r a t h e r t h a n a snow p a c k . The s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n b e t w e e n moose and u n l o g g e d f o r e s t i n m i d w i n t e r 1983 ( F i g u r e 6 and T a b l e 4) d i d not r e s u l t f r o m moose e s c a p i n g c r u s t e d snow i n open h a b i t a t s . I t i s p r o b a b l e t h a t moose a b a n d o n m e n t o f o p e n a r e a s i n w i n t e r i s a t l e a s t p a r t l y c a u s e d by a f a c t o r o t h e r t h a n snow c o n d i t i o n s . W h e t h e r moose a v o i d a n c e o f open h a b i t a t i s a t t r i b u t e d t o snow d e p t h , snow c r u s t i n g , snow d e n s i t y o r p e n e t r a t i o n i n t o snow, t h e r e were o b s e r v a t i o n s made d u r i n g t h e p r e s e n t s t u d y w h i c h c o n t r a d i c t t h e p r e d i c t e d d i r e c t i o n o f h a b i t a t s e l e c t i o n . T he r a n g e o f snow d e p t h s and c o n d i t i o n s w h i c h h a v e been i m p l i c a t e d i n moose s e e k i n g c o v e r may be c a u s e d by snow o r i e n t e d o b s e r v e r s a s c r i b i n g t h a t b e h a v i o u r t o snow c o n d i t i o n s w i t h o u t f u l l j u s t i f i c a t i o n . 2. T h e r m a l S t r e s s The d i s c u s s i o n o f t h e r m a l s t r e s s w i l l p r o c e e d i n f o u r s t e p s : F i r s t , a s t a n d a r d i s n e c e s s a r y w i t h w h i c h t o compare 93 d i f f e r e n t e n v i r o n m e n t a l c o n d i t i o n s c o m p r i s i n g a i r t e m p e r a t u r e , r a d i a t i o n , and c o n v e c t i v e h e a t l o s s . The s t a n d a r d u s e d by b i o p h y s i c a l e c o l o g i s t s i s T g , t h e e q u i v a l e n t b l a c k body t e m p e r a t u r e ( C a m p b e l l 1 9 7 7 ) . T e d e s c r i b e s a c o m b i n a t i o n o f a i r t e m p e r a t u r e and r a d i a t i o n t h a t a t a g i v e n w i n d s p e e d p r o d u c e s t h e same t h e r m a l e n v i r o n m e n t a s an e n v i r o n m e n t c h a m b e r w i t h ( i ) a i r and w a l l s a t t h e same t e m p e r a t u r e as T g a n d ( i i ) s a m e w i n d s p e e d . A f t e r a d i s c u s s i o n o f t h e c a l c u l a t i o n of T e , w i n t e r c o l d s t r e s s , w i n t e r h e a t s t r e s s , and summer he a t s t r e s s w i l l be r e v i e w e d . 2.1 C a l c u l a t i o n of E q u i v a l e n t B l a c k b o d y T e m p e r a t u r e The c a l c u l a t i o n of t h e e q u i v a l e n t b l a c k b o d y t e m p e r a t u r e , T e , o v e r an a v e r a g e 24 h o u r c y c l e p r o c e e d s i n t h r e e s t e p s : (1) c a l c u l a t i o n of h o u r l y a i r t e m p e r a t u r e , (2) c a l c u l a t i o n of r a d i a t i o n a b s o r b e d , and (3) c o m b i n i n g a i r t e m p e r a t u r e and r a d i a t i o n a b s o r b e d w i t h r e s i s t a n c e s t o h e a t t r a n s f e r t o p r o d u c e T g . F o r the p u r p o s e s of c a l c u l a t i o n , a l l t e m p e r a t u r e s were e x p r e s s e d i n d e g r e e s K e l v i n . R e s u l t s w i l l be d i s c u s s e d i n d e g r e e s C e l s i u s . 94 2.1.1 A i r t e m p e r a t u r e A i r t e m p e r a t u r e t h r o u g h t h e a v e r a g e 24 h o u r c y c l e i s c a l c u l a t e d as f o l l o w s : < 2 3) T a = T a m i n + ( T a m a x - T a m i n ) { s i n [ ( i r / 2 ) ( ( t - t m i n ) / h s P a n ) ^ ' d u r i n g t h e w a r m i n g p a r t o f t h e day and (24) T ~ ^amax -(^amax ^ a m i n ^ s i n [ (TT/2) ( ( t - t m a x ) / c s p a n ) ] } , d u r i n g t h e c o o l i n g p a r t o f t h e day, w h e r e , T g i s a i r t e m p e r a t u r e a t t h e g i v e n h o u r , T _ m - and T o m „ v a r e t h e * ^ d IU J. II d 111 d A minimum and maximum a i r t e m p e r a t u r e s f o r t h e g i v e n t i m e p e r i o d ( T a b l e 8 o r 9 ) , t i s t h e t i m e i n h o u r s , t m i n i s t h e t i m e o f t h e m i n i m u m t e m p e r a t u r e 04:00 (4 h o u r s ) i n summer and 08:00 (8 h o u r s ) i n w i n t e r , tmax i s t h e t i m e o f maximum t e m p e r a t u r e , 14:00 (14 h o u r s ) i n b o t h summer and w i n t e r , hspan i s t h e l e n g t h of t h e p e r i o d of h e a t i n g d u r i n g t h e day, and c s p a n i s t h e l e n g t h of t h e p e r i o d of c o o l i n g d u r i n g t h e day. Moen ( 1 9 8 2 ) s u g g e s t e d t h e u s e o f s i n e f u n c t i o n s t o c a l c u l a t e h o u r l y a i r t e m p e r a t u r e s f r o m d a i l y maximum and m i n i m u m t e m p e r a t u r e s , b u t he d i d n o t p r o d u c e an e q u a t i o n . B e l o v s k y ( 1 9 8 1 ) p r o d u c e d an e q u a t i o n s i m i l a r t o e q u a t i o n s 23 and 24 w h i c h u s e d c o s i n e f u n c t i o n s . H o w e v e r , B e l o v s k y ' s e q u a t i o n s p r o v i d e d u n i n t e l l i g i b l e r e s u l t s . 2.1.2 A b s o r b e d r a d i a t i o n E q u a t i o n s 25 a b s o r b e d s h o r t w a v e t h r o u g h 31 i l l u s t r a t e t h e c a l c u l a t i o n o f r a d i a t i o n : 95 (25) SWdi = 0 . 3 8 4 - 0 . 4 1 6 { S W t / [ ( S c o n ) s i n ( S e l e v ) ] } ( S c o n ) s i n ( S e l e v ) , (26) SWbi = ( S W t - S W d i ) / s i n ( S e l e v ) , w h e r e , SWdi i s t h e i n t e r i m d i f f u s e s h o r t w a v e r a d i a n t f l u x d e n s i t y , SWt i s t h e mean t o t a l s h o r t w a v e r a d i a t i o n f o r a g i v e n t i m e p e r i o d w h i c h was c a l c u l a t e d f r o m Canada A t m o s p h e r i c E n v i r o n m e n t S e r v i c e ( 1981 t o 1983) d a t a r e c o r d e d a t t h e P r i n c e George A i r p o r t , Scon i s t h e s o l a r c o n s t a n t 1360 W m ( C a m p b e l l 1977), S e l e v i s t h e s o l a r e l e v a t i o n ( C a m p b e l l 1977 e q u a t i o n 5.8), and SWbi i s t h e i n t e r i m beam s h o r t w a v e r a d i a t i o n n o r m a l t o t h e i n c i d e n t r a d i a t i o n . I f t h e s o l a r e l e v a t i o n i s l e s s t h a n 1 5 ° , t h e n SWdi = SWt and SWbi = 0. E q u a t i o n s 25 and 26 a r e fr o m L i u and J o r d a n (1960). (27) SWd = SWdi x s k y , (28) SWb = SWbi x pen, where, SWd i s the amount of d i f f u s e s h o r t w a v e r e a c h i n g a moose, sky i s t h e mean p r o p o r t i o n of sky v i s i b l e on a l l th e f i s h - e y e p h o t o g r a p h s t a k e n on a p a r t i c u l a r h a b i t a t , SWb i s t h e am o u n t o f beam s h o r t w a v e p e n e t r a t i n g t h e f o r e s t c a n o p y , and pen i s t h e mean p r o p o r t i o n o f s k y v i s i b l e a t a p a r t i c u l a r s o l a r a n g l e on a l l f i s h - e y e p h o t o g r a p h s t a k e n on a p a r t i c u l a r h a b i t a t . (29) R r f l g = (SWb + S W d ) ( r f l g ) , (30) R r f l t = (SWb + S W d ) ( r f l t ) ( l - s k y ) , where, Rj-fjo i s t h e r a d i a t i o n r e f l e c t e d f r o m t h e ground, r f l g i s t h e p r o p o r t i o n of r a d i a t i o n s t r i k i n g t h e ground r e f l e c t e d , 0.2 i n summer ( B e l o v s k y 1981) and 0.7 i n w i n t e r ( a p p r o x i m a t e l y the a v e r a g e of f r e s h and w e a t h e r e d snow, Moen 1982), R r f i t i s t h e r a d i a t i o n r e f l e c t e d f r o m t r e e s , r f l t i s t h e r e f l e c t i v i t y of t r e e s 0.18 ( B e l o v s k y 1981) . 96 ( 3 1 ) R s w g = [ 0 . 5 ( S w d ) + a p a ( S w b ) + 0 . 5 ( R r f l g + R r f l t ) ] ( a s ) , w h e r e , ^ s w a i s t h e a m o u n t o f s h o r t w a v e r a d i a t i o n a b s o r b e d , apa i s t h e shadow a r e a of a moose (0.3, P a r k e r 1983), and as i s the a b s o r p t i v i t y of moose to s h o r t w a v e r a d i a t i o n s e t t o t h e same v a l u e as t h a t of a brown cow, 0.74 ( B e l o v s k y 1 9 8 1 ) . E q u a t i o n s 32 t h r o u g h 35 show t h e c a l c u l a t i o n of longwave r a d i a t i o n a b s o r b e d . (32) LWs = e s 0 T a 4 ( s k y ) , (33) LWt = £ t o T a 4 ( 1 - s k y ) , (34) LWg = £ g 0 T g 4 , w h e r e , LWs i s l o n g w a v e r a d i a t i o n f r o m t h e s k y , e i s t h e e m i s s i v i t y o f t h e s k y (0.72 + ( 0 . 0 0 5 ) ( T g - 273.16), C a m p b e l l 1 9 7 7 ) , C i s t h e S t e f a n - B o l t z m a n n c o n s t a n t (5.6x10" ), LWt i s longwa v e r a d i a t i o n f r o m t r e e s , £ f c i s t h e e m i s s i v i t y o f t r e e s ( 0 . 9 9 ) , LWg i s l o n g wave f r o m t h e g r o u n d , e i s t h e e m i s s i v i t y o f t h e g r o u n d (0.99 i n summer and 1.00 i n w i n t e r ) , and T i s t h e t e m p e r a t u r e of t h e g r o u n d (T = T g i n summer and T = 2 7 3 . 1 6 ° K i n w i n t e r ) . 8 8 W i n t e r v a l u e s of T and e a r e t h o s e of snow (Moen 1982). The amount of longwave r a d i a t i o n a b s o r b e d i s , (35) R l w a = (LWs+LWt+LWg)(0.5), where, R^ w a i s longwave r a d i a t i o n a b s o r b e d . T o t a l r a d i a t i o n a b s o r b e d i s , < 3 6> R a b s = R s w a + R l w a where, R a j j S i s t h e t o t a l r a d i a t i o n a b s o r b e d . 97 2.1.3 R e s i s t a n c e t o h e a t t r a n s f e r and Te The c a l c u l a t i o n of r e s i s t a n c e t o c o n v e c t i v e h e a t l o s s i s , (37) r h a = 3 0 7 ( D / U ) ' 5 , where, r n g i s t h e r e s i s t a n c e t o c o n v e c t i v e h e a t l o s s , D i s a c h a r a c t e r i s t i c d i m e n s i o n s e t a t 0.66 m f o r moose, and U i s t h e w i n d s p e e d s e t a t 0.5 m s . The c o e f f i c i e n t , 307, i n e q u a t i o n 37 p r o b a b l y p r o d u c e s an r n a l a r g e r t h a n t h a t w h i c h p r o d u c e d i n f i e l d c o n d i t i o n s . T u r b u l e n c e i s t y p i c a l l y more i n t e n s e o u t d o o r s t h a n i n w i n d t u n n e l s w h e r e r e s i s t a n c e c o e f f i c i e n t s w e r e c a l c u l a t e d . H o w e v e r , D=0.66 m a s s u m e s w i n d b l o w i n g p e r p e n d i c u l a r t o t h e l o n g a x i s o f a moose, v i e w e d as a c y l i n d e r . A moose f a c i n g i n t o t h e w i n d w o u l d h a v e a g r e a t e r r e s i s t a n c e t h a n one w i t h i t s s i d e t o t h e wind. The wind speed was c h o s e n t o p r o d u c e a r e s i s t a n c e t o f o r c e d c o n v e c t i v e h e a t l o s s t h a t i s s m a l l e r t h a n t h e r e s i s t a n c e t o f r e e c o n v e c t i v e h e a t l o s s w o u l d be. T h a t i s , e q u a t i o n 37 assumes an a c t i v e c o n v e c t i v e h e a t l o s s t h a t i s l a r g e r t h a n a p a s s i v e s y s t e m w o u l d p r o d u c e . A t o t a l i z i n g a n e m o m e t e r 1.8 m a b o v e t h e g r o u n d i n a r e c e n t c l e a r c u t c o n s i s t e n t l y r e c o r d e d mean m o n t h l y w i n d s p e e d s o f l e s s t h a n 0.5 m s " 1 . The r e s i s t a n c e t o r a d i a t i v e h e a t l o s s i s , 98 (38) r r = p c p / ( 4 e o T a 3 ) , where, r r i s t h e r e s i s t a n c e t o r a d i a t i v e h e a t l o s s , pc i s t h e v o l u m e t r i c h e a t c a p a c i t y of a i r a p p r o x i m a t e d by 1200 ( C a m p b e l l 1 9 7 7 ) , e i s t h e e m i s s i v i t y o f a moose s u r f a c e (0.9 9 ) , O i s t h e S t e p h a n - B o l t z m a n n c o n s t a n t (5.67 x 10 ), and T a i s t h e a i r t e m p e r a t u r e . The e q u i v a l e n t p a r a l l e l r e s i s t a n c e t o c o n v e c t i v e and r a d i a t i v e h e a t l o s s i s , (39) r e = [ ( r h a ) ( r r ) ] / ( r h a + r r ) , w h e r e , r g i s t h e e q u i v a l e n t p a r a l l e l r e s i s t a n c e t o c o n v e c t i v e a n d r a d i a t i v e h e a t l o s s , r h a * s t n e r e s i s t a n c e t o c o n v e c t i v e h e a t l o s s , a n d r i s t h e r e s i s t a n c e t o r a d i a t i v e h e a t l o s s . E q u a t i o n s 37 and 38 and 39 a r e f r o m C a m p b e l l (1977). F i n a l l y , t h e e q u i v a l e n t b l a c k b o d y t e m p e r a t u r e i s , (40) T e = T a + [ ( r e ) ( R a b s - e c T a 4 ) ] / p C p , where, T e i s t h e e q u i v a l e n t b l a c k b o d y t e m p e r a t u r e , T i s t h e a i r t e m p e r a t u r e , r g i s t h e e q u i v a l e n t p a r a l l e l r e s i s t a n c e t o c o n v e c t i v e and r a d i a t i v e h e a t l o s s , R a D S i s t h e t o t a l r a d i a t i o n a b s o r b e d , £ i s t h e e m m i s s i v i t y or moose, C i s t h e S t e p h a n - B o l t z m a n n c o n s t a n t , and pc i s the v o l u m e t r i c h e a t c a p a c i t y of a i r . E q u a t i o n 40 i s f r o m C a m p b e l l ( 1 9 7 7 ) . W h i l e e q u a t i o n 40 a p p e a r s t o be s u b s t i t u t i n g a i r t e m p e r a t u r e f o r s u r f a c e t e m p e r a t u r e t h i s i s n o t t h e c a s e . The p r o b l e m o f s u r f a c e t e m p e r a t u r e was s o l v e d i n C a m p b e l l ' s ( 1 9 7 7 ) e q u a t i o n s 7.7, 7.8, 7.9, a n d 7.10, w h i c h p r o d u c e d t h e c o n c e p t o f r e , a p p e a r i n g i n e q u a t i o n 40. T h e r e i s an e r r o r i n v o l v e d i n t h e 99 e s t i m a t i o n o f T e ^ » l e a d i n g t o e q u a t i o n 40. The b l a c k body l i n e , a c u r v e d r e l a t i o n s h i p between R a D S a n ^ ^ a ( s e e C a m p b e l l 1977, p. 9 0 - 9 2 ) , i s p r o j e c t e d as a s t r a i g h t l i n e a t T g . T h i s e s t i m a t e o f T g does not l e a d t o u n a c c e p t a b l e e r r o r u n l e s s t h e d i f f e r e n c e between an a n i m a l ' s body t e m p e r a t u r e and e s t i m a t e d T g i s v e r y l a r g e . W h i l e T g i s t h e f i n a l p r o d u c t of t h e h e a t f l u x e q u a t i o n s u s e d i n t h i s t h e s i s , C a m p b e l l ( 1 9 7 7 ) p r o d u c e s a f u r t h e r e q u a t i o n : (41) M-XE = [ p c p ( T b - T e ) ] / ( r H b + r e ) , w h e r e , M i s t h e m e t a b o l i c r a t e , XE i s t h e l a t e n t h e a t l o s s due t o e v a p o r a t i o n , T^ i s body t e m p e r a t u r e , and r ^ b i s t h e a v e r a g e c o a t p l u s t i s s u e r e s i s t a n c e t o he a t l o s s . E q u a t i o n 41 p e r m i t s t h e c a l c u l a t i o n of r ^ and M, f o r a g i v e n T , i f e i t h e r r ^ b o r M i s known. A s o l u t i o n t o e q u a t i o n 41 p e r m i t s an e s t i m a t e of t h e amount of f o o d r e q u i r e d f o r a g i v e n t i m e a t T e . A l s o , i f p h y s i o l o g i c a l e x t r e m e s of M and r ^ a r e used i n e q u a t i o n 41, t h e n t h e upper and l o w e r t h e r m a l l i m i t s , e x p r e s s e d as T e , can be d e f i n e d . 2.2 W i n t e r C o l d S t r e s s I n t h e summary t o t h e r e s u l t s s e c t i o n i t was n o t e d t h a t t h e r e was some d i f f i c u l t y e x p l a i n i n g t h e c o r r e l a t i o n between moose and f o r e s t c o v e r d u r i n g w i n t e r on t h e b a s i s o f c o l d 100 s t r e s s . M i d w i n t e r and l a t e w i n t e r o f b o t h 1982 and 1983 p r o d u c e d c o r r e l a t i o n s b e t w e e n moose a s s o c i a t i o n s and f o r e s t c o v e r w i t h a low p r o b a b i l i t y of h a v i n g been p r o d u c e d by random e v e n t s . Y e t , o n l y one of t h o s e f o u r p e r i o d s , m i d w i n t e r 1982, c o u l d be d e s c r i b e d as c o l d . A c o l d s e a s o n was d e f i n e d as one i n w h i c h m i n i m u m t e m p e r a t u r e s f r e q u e n t l y f e l l b e l o w - 2 0 ° . F u r t h e r , M a r k g r e n ( 1 9 6 6 ) r e p o r t e d t h a t moose c a l v e s a p p e a r u n a f f e c t e d by c o l d . T e m p e r a t u r e s as c o l d as - 2 8 ° p r o d u c e d no b e h a v i o u r a l r e s p o n s e and what was d e s c r i b e d a s " v e r y h a r d w e a t h e r " , - 2 4 ° w i t h a w i n d , and v e r y h i g h h u m i d i t y (85 t o 95 % ) , was n o t s u f f i c i e n t t o s t r e s s c a p t i v e moose ( M a r k g r e n 1966). The e v i d e n c e f r o m s t u d i e s o f t h e m e t a b o l i c r e s p o n s e of moose t o c o l d c o n d i t i o n s i s t h a t moose a r e v e r y r e s i s t a n t t o c o l d . R e n e c k e r e t a l . (1978) r e p o r t e d t h a t moose c a l v e s o n l y i n c r e a s e d t h e i r m e t a b o l i s m t o d e a l w i t h c o l d when a i r t e m p e r a t u r e s were l o w e r t h e n - 2 0 ° w i t h wind s p e e d s o f 8 m s - ^ (28.8 k m / h r ) . R e n e c k e r and H u d s o n ( 1 9 8 4 ) w e r e u n a b l e t o p r o d u c e a m e t a b o l i c r e s p o n s e i n a d u l t m o o s e when a i r t e m p e r a t u r e s were l e s s t h a n -30°C. The c o n d i t i o n s under w h i c h t h e s e a u t h o r s were a b l e t o p r o d u c e c o l d s t r e s s i n moose n e v e r o c c u r r e d d u r i n g t h e m i d w i n t e r and l a t e w i n t e r p e r i o d s of 1982 and 1983. I t i s u n l i k e l y t h a t c o l d was r e s p o n s i b l e f o r t h e p a t t e r n s of h a b i t a t a s s o c i a t i o n r e c o r d e d d u r i n g t h e w i n t e r i n t h e p r e s e n t s t u d y . 101 2.3 W i n t e r Heat S t r e s s I n c o n t r a s t w i t h t h e i r i n s e n s i t i v i t y t o c o l d , moose a r e e a s i l y h e a t s t r e s s e d i n w i n t e r . R e n e c k e r and H u d s o n ( 1 9 8 4 ) r e p o r t e d t h a t t h e upper c r i t i c a l a i r t e m p e r a t u r e f o r moose i n w i n t e r i s between -5.1°C and -0.4°C. At t e m p e r a t u r e s i n t h i s r a n g e , moose m e t a b o l i c r a t e s r o s e and t h e i r r e s p i r a t i o n r a t e s e l e v a t e d i n o r d e r t o l o s e h e a t . By c o m b i n i n g t h e a i r t e m p e r a -t u r e m e t a b o l i s m r e l a t i o n s h i p s ( R e n e c k e r and Hudson 1984) w i t h b l a c k g l o b e t e m p e r a t u r e (T ) m e t a b o l i s m r e l a t i o n s h i p s ( R e n e c k e r 1 9 8 5 ) , an e q u i v a l e n t b l a c k b o d y t e m p e r a t u r e ( T e » C a m p b e l l 1977) o f a b o u t 0 ° C p r o d u c e s t h e r m a l s t r e s s i n w i n t e r i n g moose. T d e s c r i b e s a t e m p e r a t u r e r e c o r d e d i n s i d e O an a i r t i g h t g l o b e , 10.2 cm i n d i a m e t e r , w h i c h had been p a i n t e d mat b l a c k and p o s i t i o n e d a t moose f l a n k h e i g h t . T e d e s c r i b e s a c o m b i n a t i o n of a i r t e m p e r a t u r e and r a d i a t i o n t h a t a t a g i v e n w i n d s p e e d p r o d u c e s t h e same t h e r m a l e n v i r o n m e n t as an e n v i r o n m e n t chamber w i t h a i r and w a l l s a t t h e same t e m p e r a t u r e a s T e . T g was c a l c u l a t e d f o r t h r e e h a b i t a t s , o l d c l e a r c u t , i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g and u n l o g g e d f o r e s t , f o r a v e r a g e 24 h o u r c y c l e s r e p r e s e n t i n g m i d w i n t e r o f 1982 and 1983. The a i r t e m p e r a t u r e s u s e d i n t h e s e c a l c u l a t i o n s came f r o m t h e s e a s o n a l d a t a g a t h e r e d on t h e s t u d y a r e a ( T a b l e 8) 102 a n d t h e s o l a r r a d i a t i o n d a t a w e r e g a t h e r e d a t t h e P r i n c e George a i r p o r t (Canada A t m o s p h e r i c E n v i r o n m e n t S e r v i c e 1981-1 9 8 3 ) . The r e s u l t s o f t h e s e c a l c u l a t i o n s a r e p r e s e n t e d i n F i g u r e 9. Of t h e s i x s i t u a t i o n s i n F i g u r e 9 o n l y one, u n l o g g e d f o r e s t d u r i n g m i d w i n t e r 1982, d i d n o t e x c e e d t h e u p p e r c r i t i c a l t e m p e r a t u r e a t any t i m e d u r i n g t h e a v e r a g e 24 h o u r c y c l e . On t h e o t h e r h a n d i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g e x c e e d e d t h e upper c r i t i c a l t e m p e r a t u r e f o r t h r e e and a h a l f h o u r s and s e v e n h o u r s d u r i n g a v e r a g e m i d w i n t e r d a y s i n 1982 and 1983 r e s p e c t i v e l y . O l d c l e a r c u t s e x c e e d e d upper c r i t i c a l t e m p e r a t u r e s f o r f i v e h o u r s and s e v e n h o u r s on a v e r a g e m i d -w i n t e r d a y s . O l d c l e a r c u t s and i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g s u b s t a n t i a l l y e x c e e d t h e upper c r i t i c a l t e m p e r a t u r e , o l d c l e a r c u t s more so t h a n i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g , d u r i n g m i d w i n t e r 1983. The l e n g t h o f t i m e t h e t h e r m a l c o n d i t i o n s i n a h a b i t a t e x c e e d u p p e r c r i t i c a l l e v e l s i s i m p o r t a n t t o moose. D u r i n g s u c h c o n d i t i o n s t h e r a n g e o f b e h a v i o u r o f an a n i m a l i s l i m i t e d , i n p a r t i c u l a r , f e e d i n g c e a s e s as t h e a n i m a l s engage i n b e h a v i o u r w h i c h w i l l r e s u l t i n th e l o s s of e x c e s s h e a t ( R e n e c k e r and Hudson 1984). R e n e c k e r and H u d s o n ( 1 9 8 4 ) n o t e d t h a t tame moose l o s t w e i g h t when w i n t e r t h e r m a l c o n d i t i o n s e x c e e d e d upper c r i t i c a l t e m p e r a t u r e s d e s p i t e n u t r i t i o u s f o o d b e i n g r e a d i l y a v a i l a b l e . 103 F i g u r e 9. E q u i v a l e n t b l a c k b o d y t e m p e r a t u r e (T ) d u r i n g an a v e r a g e d a i l y c y c l e i n m i d w i n t e r 1982 ( t o p ) and m i d w i n t e r 1983 ( b o t t o m ) 105 U n l i k e c o l d s t r e s s or f o r a g e abundance t e m p e r e d by snow d e p t h , h e a t s t r e s s p r o d u c e s e x p e c t a t i o n s o f w i n t e r h a b i t a t a s s o c i a t i o n c o n s i s t e n t w i t h t h o s e d o c u m e n t e d . When t h e e q u i v a l e n t b l a c k b o d y t e m p e r a t u r e (T ) e x c e e d s upper c r i t i c a l l i m i t s moose w o u l d g e n e r a l l y n o t be f o u n d i n o l d c l e a r c u t s b e c a u s e t h e r e i s v e r y l i t t l e s h a d e t o s h i e l d t hem f r o m s o l a r r a d i a t i o n and t h e c o n d i t i o n s i n t h e c l e a r c u t s c an be extr e m e . I n t e r m e d i a t e u t i l i z a t i o n l o g g i n g can be o c c u p i e d b e c a u s e i t i s p o s s i b l e t o f i n d d e ep s h a d e on t h e s e s i t e s . F o r e x a m p l e , on 22 M a r c h 1983 s i x o f e l e v e n r a d i o c o l l a r e d moose w e r e i n i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g . However, d e s p i t e abundant f r e s h t r a c k s i n i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g t h e r e were no moose s i g h t e d d u r i n g a t e l e m e t r y f l i g h t b e t w e e n 10:00 and 12:00 t h a t day. F i n a l l y , t h e t h e r m a l c o n d i t i o n s i n f o r e s t s a r e u s u a l l y a c c e p t a b l e t o moose. When t e m p e r a t u r e s e x c e e d e d upper c r i t i c a l l i m i t s , t h e amount was r e l a t i v e l y s m a l l . D u r i n g l a t e w i n t e r t h e p o s s i b i l i t y o f h e a t s t r e s s would be g r e a t e r t h a n i n m i d w i n t e r b e c a u s e a i r t e m p e r a t u r e s a r e h i g h e r and t h e r e i s g r e a t e r p o t e n t i a l f o r h i g h l e v e l s of s o l a r r a d i a t i o n . I n s p r i n g , p r o b l e m s w i t h h e a t s t r e s s s h o u l d be l e s s . Snow d i s a p p e a r s about t h e b e g i n n i n g of s p r i n g , p a r t i c u -l a r l y i n open a r e a s . R e f l e c t i o n of s o l a r r a d i a t i o n f r o m snow i s a l a r g e f a c t o r i n p r o d u c i n g h i g h r a d i a n t f l u x d e n s i t i e s i n w i n t e r . A l s o , moose l o s e t h e i r heavy w i n t e r c o a t s i n s p r i n g , 106 b e g i n n i n g i n A p r i l ( K n o r r e 1959). C a u t i o n s h o u l d be a d v i s e d i n a t t r i b u t i n g a l l w i n t e r moose b e h a v i o u r t o t h e a v o i d a n c e o f h e a t s t r e s s . F i r s t , t h e c a l c u l a t i o n s a s s u m e t h a t R e n e c k e r and Hudson's ( 1 9 8 4 ) b l a c k g l o b e t e m p e r a t u r e (T ) i s e q u a l t o C a m p b e l l ' s ( 1 9 7 7 ) e q u i v a l e n t b l a c k b o d y t e m p e r a t u r e ( T g ) . T h i s a s s u m p t i o n may not h o l d . H o w e v e r , R e n e c k e r and H u d s o n ( 1 9 8 4 ) s t a t e d t h a t t h e i r b l a c k g l o b e t e m p e r a t u r e m e a s u r e m e n t s a r e an a t t e m p t t o e s t i m a t e e q u i v a l e n t b l a c k b o d y t e m p e r a t u r e s . F u r t h e r , R e n e c k e r and Hudson's measurements of T can be compared t o e s t i m a t e s of T e g e n e r a t e d f o r i n d i v i d u a l h a b i t a t s i n t h e p r e s e n t s t u d y : F o r a g i v e n a i r t e m p e r a t u r e m e a s u r e m e n t s o f T a r e g r e a t e r t h a n e s t i m a t e s of T e f o r u n l o g g e d f o r e s t , a p p r o x i m a t e l y e q u a l t o e s t i m a t e s of T e f o r i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g , and a r e much l e s s t h a n e s t i m a t e s of T g f o r o l d c l e a r c u t s . T h i s i s p r e c i s e l y t h e s i t u a t i o n e x p e c t e d , i f R e n e c k e r and Hudson's (1984) measurements of T o c c u r r e d i n a s e m i - o p e n e n v i r o n m e n t . R e n e c k e r a n d H u d s o n ' s d e s c r i p t i o n o f t h e i r m e t h o d o f measurement and t h e r a d i a n t h e a t l o a d s r e c o r d e d s u g g e s t t h a t m e a s u r e m e n t s w e r e i n f a c t c o n d u c t e d i n a s e m i - o p e n e n v i r o n m e n t . The c r i t i c a l q u e s t i o n i s w h e t h e r moose were a b l e t o f o r a g e i n t h e c o n d i t i o n s p r e s e n t when moose l o c a t i o n s - w e r e r e c o r d e d . T g was c a l c u l a t e d f o r a s t a n d i n g a n i m a l . T h e r e would be an a d d i t i o n a l m e t a b o l i c h e a t l o a d on a f e e d i n g moose 107 ( R e n e c k e r 1985) w h i c h w o u l d r e d u c e t h e u p p e r c r i t i c a l T e . T h i s a d d i t i o n a l h e a t l o a d was not c o n s i d e r e d i n t h e e s t i m a t e s of t h e upper c r i t i c a l T g . The s e c o n d p o i n t t o c o n s i d e r i s t h a t t h e r e may be o t h e r f a c t o r s on w i n t e r r a n g e t h a t a r e i m p o r t a n t t o moose d e s p i t e th e l a c k of s i g n i f i c a n t c o r r e l a t i o n s w i t h moose l o c a t i o n s i n t h i s t h e s i s . T h e r e a r e d a t a w h i c h d e m o n s t r a t e t h a t f o r e s t c a n o p i e s can r e d u c e snow d e p t h s on moose w i n t e r r a n g e ( T e l f e r 1970, S i l v e r 1976, Eastman 1978, S i n g e r 1979, P i e r c e and Peek 1 9 8 4 ) w h i c h s h o u l d r e s u l t i n r e d u c e d e n e r g y c o s t s f o r l o c o m o t i o n . E n e r g y s a v i n g s d u r i n g w i n t e r can be c r i t i c a l t o the s u r v i v a l and r e p r o d u c t i o n of moose (Edwards 1956, R o l l e y and K e i t h 1980). A l s o , t h e r e l a t i v e l y l a r g e f o r a g e p r o d u c t i o n i n o l d c l e a r c u t s and i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g c o u l d w e l l be i m p o r t a n t t o t h e s u r v i v a l o f moose on t h e w i n t e r r a n g e under s t u d y . 2.4 Summer Heat S t r e s s M oose a p p e a r t o be e x t r a o r d i n a r i l y s e n s i t i v e t o summer h e a t . R e n e c k e r a n d H u d s o n ( 1 9 8 4 ) o b s e r v e d e l e v a t e d r e s p i r a t i o n r a t e s and m e t a b o l i s m when a i r t e m p e r a t u r e s e x c e e d e d 14° and p a n t i n g when a i r t e m p e r a t u r e s e x c e e d e d 20°. An u p p e r c r i t i c a l t e m p e r a t u r e f o r moose i n t h e r e g i o n o f 1 4 ° 108 t o 2 0 ° C i s a s much a s 1 5 ° l o w e r t h a n t h a t o f m u l e d e e r and e l k ( P a r k e r and R o b b i n s 1 9 8 4 ) . M o ose d e a l w i t h e x c e s s i v e h e a t p o o r l y . F i n c h (1972) n o t e d t h a t among A f r i c a n u n g u l a t e s l a r g e a n i m a l s , e.g. e l a n d ( T a u r o t r a g u s o r y x ) , l o s e h e a t by s w e a t i n g when t h e y have an e x c e s s i v e h e a t l o a d , w h i l e s m a l l e r a n i m a l s , e.g. h a r t e b e e s t ' (A.l£££.Jl£.l££ ££££££ll£ . L £ £ )» p a n t . P a r k e r ' s ( 1 9 8 3 ) o b s e r v a t i o n s on N o r t h A m e r i c a n , e l k (Ce_jrvjij3 ^J.a^jrjJhJJ^) and mule deer ( O d o c o i l e u s h e m i n o u s ) , i n d i c a t e d t h a t t h e t r e n d i s c o n s i s t e n t between t h e two c o n t i n e n t s . Moose, however, a r e an e x c e p t i o n . K n o r r e ( 1 9 5 9 ) and R e n e c k e r and H u d s o n ( 1 9 8 4 ) have b o t h o b s e r v e d moose p a n t i n g t o s l o u g h an e x c e s s i v e h e a t l o a d . A l a r g e a n i m a l s u c h a s a moose s h o u l d n o t be a b l e t o pant e f f i c i e n t l y b e c a u s e p a n t i n g would r e q u i r e a l a r g e amount of e n e r g y and would p r o d u c e a l a r g e amount o f m e t a b o l i c h e a t . T h e r e f o r e , h e a t l o a d s i n e x c e s s of c r i t i c a l t e m p e r a t u r e s a r e more s i g n i f i c a n t t o moose t h a n u n g u l a t e s w h i c h can d e a l w i t h h e a t e f f e c t i v e l y . E x c e e d i n g c r i t i c a l t e m p e r a t u r e s i n summer i s i m p o r t a n t f o r moose as i t r e s u l t s i n l e s s t i m e s p e n t f e e d i n g ( K n o r r e 1959, B e l o v s k y a n d J o r d a n 1978, R e n e c k e r and Hudson 1984) and w e i g h t can be l o s t ( R e n e c k e r and Hudson 1984) or w e i g h t g a i n s much r e d u c e d ( K n o r r e 1959). R e n e c k e r a n d H u d s o n ' s ( 1 9 8 4 ) v a l u e s f o r e l e v a t e d r e s p i r a t i o n and open mouthed p a n t i n g a r e e q u a l t o e q u i v a l e n t b l a c k b o d y t e m p e r a t u r e s ( T g ) of 20° and 30° r e s p e c t i v e l y . T e ' s w e r e c a l c u l a t e d f o r d a i l y c y c l e s r e p r e s e n t i n g mean t h e r m a l 109 c o n d i t i o n s i n f o u r p e r i o d s : summer 1981, summer 1982, J u l y and A u g u s t 1981, and J u l y and A u g u s t 1982. F i g u r e s 10 and 11 p r e s e n t t h e r e s u l t s of t h e s e c a l c u l a t i o n s . The c a l c u l a t i o n s w e r e f o r s t a n d i n g moose. The m e t a b o l i c h e a t l o a d o f a l y i n g moose w o u l d h a v e b e e n l e s s , w h i l e t h a t o f a f e e d i n g moose would have been g r e a t e r ( R e n e c k e r 1985). C l e a r l y moose were u n d e r h e a t s t r e s s i n b o t h 1981 and 1982. I n t h e c o o l e s t h a b i t a t i n t h e c o o l e r summer, u n l o g g e d f o r e s t i n 1982, t h e upper c r i t i c a l t e m p e r a t u r e was e x c e e d e d f o r about s i x h o u r s i n m i d d a y . The s i t u a t i o n was more s e v e r e i n i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g and much more s e v e r e i n o l d c l e a r c u t s . M o o se w o u l d h a v e b e e n p a n t i n g f o r s i x and a h a l f h o u r s p e r day i n o l d c l e a r c u t s i n t h e summer o f 1982 a n d f o r a b o u t e i g h t h o u r s i n 1981. F i g u r e 11, mean d a i l y c y c l e s o f T g f o r J u l y and A u g u s t , p r o b a b l y g i v e s a b e t t e r t h e r m a l p i c t u r e o f moose h a b i t a t s , d u r i n g t h e two summers. I n p a r t i c u l a r t h e s e d a t a a r e n o t i n f l u e n c e d by a v e r y h o t l a t e J u n e 1982 and a v e r y c o o l l a t e J u n e 1981. I n J u l y and A u g u s t 1981 moose w o u l d h a v e had t o e n g a g e i n t h e r m a l p a n t i n g f o r a l m o s t n i n e h o u r s i n o l d c l e a r c u t s and se v e n h o u r s i n i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g . I n J u l y and A u g u s t 1982 t h e r m a l p a n t i n g w o u l d h a v e b e e n r e q u i r e d f o r s i x and a h a l f h o u r s i n o l d c l e a r c u t s and a b o u t f o u r h o u r s i n i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g . In J u l y and 110 F i g u r e 10. E q u i v a l e n t b l a c k b o d y t e m p e r a t u r e (T ) d u r i n g an a v e r a g e d a i l y c y c l e summer 1981 ( t o p ) and summer 1982 ( b o t t o m ) 4 6 6 open mouth panting respiration rate elevated 1 1 r 16 16 17 u n l o g g » d ( o r « a t  l n ) « r m « d l o U u t i l i z a t i o n o l d c t o o r c u t 112 F i g u r e 11. E q u i v a l e n t b l a c k b o d y t e m p e r a t u r e ( T e ) d u r i n g an a v e r a g e d a i l y c y c l e J u l y and A u g u s t 1981 ( t o p ) and J u l y and August 1982 ( b o t t o m ) 114 August 1982 t h e r m a l c o n d i t i o n s i n f o r e s t b a r e l y e x c e e d e d upper c r i t i c a l t e m p e r a t u r e s . U n l i k e many of t h e h y p o t h e s e s s c r u t i n i z e d i n t h i s t h e s i s , t h e a v o i d a n c e o f summer h e a t s t r e s s i s c o n s i s t e n t w i t h t h e summer moose h a b i t a t a s s o c i a t i o n d a t a . The o b s e r v e d p a t t e r n s o f moose h a b i t a t a s s o c i a t i o n s ( F i g u r e s 4, 5, 6 and T a b l e 4) a r e e x p l i c a b l e on t h e b a s i s o f F i g u r e s 10 and 11. The f o r e s t was a r e l a t i v e l y c o m f o r t a b l e p l a c e f o r moose, d u r i n g t h e summers o f 1981 and 1982. When a v e r a g e c o n d i t i o n s e x c e e d e d moose t h e r m a l t o l e r a n c e , t h e n t h e y would have a c c e s s t o deep s h a d e and p o s s i b l y wet a r e a s t o c o o l t h e m s e l v e s . The same would be t r u e f o r moose i n i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g i n 1982, but i n 1981, p a r t i c u l a r l y J u l y and August, i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g would have been an i n h o s p i t a b l e p l a c e f o r moose r e q u i r i n g t h e r m a l p a n t i n g , d u r i n g a l a r g e p o r t i o n o f s u n l i g h t h o u r s , t o p r e v e n t h e a t o v e r l o a d . T h i s s i t u a t i o n c o u l d a c c o u n t f o r r e l a t i v e l y l i t t l e u s e o f i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g i n summer 1981 b u t r e l a t i v e l y h i g h u s e d u r i n g summer 1982 ( T a b l e 3, T a b l e 4, F i g u r e 5, and F i g u r e 6). O l d c l e a r c u t s w o u l d h a v e b e e n much t o o h o t f o r moose d u r i n g summer, e s p e c i a l l y t h e summer of 1981. N o t e , t h e c a l c u l a t i o n o f T g p r e s e n t e d i n t h i s s e c t i o n makes a c r i t i c a l a s s u m p t i o n : T i n R e n e c k e r and Hudson (1984) i s t h e same as T i n C a m p b e l l ( 1 9 7 7 ) . A j u s t i f i c a t i o n f o r 1 1 5 t h i s a s s u m p t i o n i s p r e s e n t e d i n t h e d i s c u s s i o n of w i n t e r h e a t s t r e s s ( S e c t i o n 2.3). The i n f o r m a t i o n p r e s e n t e d i n t h i s t h e s i s i n d i c a t e s t h a t s h e l t e r f r o m h e a t i s an i m p o r t a n t c o n s i d e r a t i o n f o r moose i n summer. W h i l e a moose h a b i t a t manager s h o u l d not i g n o r e s u c h c o n s i d e r a t i o n s as f o r a g e p r o d u c t i o n , summer r a n g e s h o u l d not be c o n s i d e r e d c o m p l e t e w i t h o u t p r o v i s i o n f o r t h e r m a l s h e l t e r . 3. U n t e s t e d H y p o t h e s e s T h e r e a r e t h r e e f a c t o r s w h i c h may have c o n t r i b u t e d t o t h e p a t t e r n o f moose h a b i t a t a s s o c i a t i o n s documented i n t h i s s t u d y t h a t w e r e n o t e x p l i c i t l y t e s t e d : p r e d a t o r s may a f f e c t t h e h a b i t a t moose c h o o s e , human d i s t u r b a n c e c o u l d a f f e c t t h e h a b i t a t s moose o c c u p i e d , and moose b e h a v i o u r p a t t e r n s c o u l d be l e a r n e d f r o m t h e i r dams and may not be an i m m e d i a t e r e s p o n s e t o p r e v a l e n t c o n d i t i o n s . K n o r r e ( 1 9 5 9 ) s u g g e s t e d t h a t i n w i n t e r moose u s e d e n s e f o r e s t as a v i s u a l b a r r i e r between t h e m s e l v e s and p r e d a t o r s . A p a r t f r o m t h a t r e f e r e n c e t h e a b i l i t y o f moose t o a v o i d p r e d a t i o n h a s b e e n i g n o r e d a s mo s t a u t h o r s e x p l o r e moose h a b i t a t s e l e c t i o n w i t h r e s p e c t t o t h e a v a i l a b i l i t y of f o r a g e and s h e l t e r . O t h e r a u t h o r s s u g g e s t t h a t moose s e l e c t h a b i t a t on t h e b a s i s o f f a c t o r s o t h e r t h a n p r e d a t o r s w h i c h a l t e r s 116 p r e d a t i o n r a t e s as a s e c o n d a r y e f f e c t . F o r example, P e t e r s o n and A l l e n ( 1 9 7 4 ) d i s c u s s e d how snow c o n d i t i o n s and t h e a v a i l a b i l i t y o f f o r a g e c a u s e d moose h a b i t a t s h i f t s w h i c h c h a n g e t h e v u l n e r a b i l i t y o f moose t o w o l v e s . H o w e v e r , many a u t h o r s h a v e d o c u m e n t e d moose s h i f t i n g t o d e n s e r h a b i t a t as w i n t e r p r o g r e s s e s . Whatever t h e o t h e r e x p l a n a t i o n s w h i c h may a c c o u n t f o r t h i s b e h a v i o u r , i t i s a l s o p o s s i b l e t h a t as w i n t e r p r o g r e s s e s and t h e p h y s i c a l c o n d i t i o n of moose d e t e r i o r a t e s , e n c o u n t e r s w i t h p r e d a t o r s must be a v o i d e d b e c a u s e e n e r g y e x p e n d i t u r e s a t t h a t t i m e c o u l d be f a t a l even i f t h e p r e d a t o r s a r e u n s u c c e s s f u l . A l s o , moose s e e k i n g h a b i t a t w i t h l o w v i s i b i l i t y t o a v o i d p r e d a t i o n c o u l d e x p l a i n t h e h i g h p r o p o r t i o n o f moose l o c a t i o n s i n h a b i t a t s w i t h h i g h f o r e s t c o v e r v a l u e s i n a l l s e a s o n s . M oose a v o i d a n c e o f human d i s t u r b a n c e i s a p o t e n t i a l f a c t o r i n moose h a b i t a t s e l e c t i o n w h i c h has a l s o been l i t t l e d i s c u s s e d . However, Hancock (1976) i n c o m p a r i n g a r e a s w h i c h r a n k e d e q u a l l y as moose w i n t e r r a n g e , i . e . t h e r e was e q u a l b r o w s e p r o d u c t i o n i n a l l a r e a s , f o u n d t h a t moose p r e f e r r e d w i n t e r i n g a r e a s w i t h l i t t l e human d i s t u r b a n c e . Tomm ( 1 9 7 8 ) fo u n d t h a t moose made l e s s use of c u t b l o c k s e x p o s e d t o human h a r r a s s m e n t t h a n u n d i s t u r b e d c u t b l o c k s . R o l l e y and K e i t h ( 1 9 8 0 ) and M y t t o n and K e i t h ( 1 9 8 1 ) f o u n d moose f a r t h e r f r o m human d i s t u r b a n c e t h a n e x p e c t e d a t r a n d o m e x c e p t d u r i n g 117 m i d w i n t e r and l a t e w i n t e r . The g r e a t e s t human d i s t u r b a n c e t o moose, i n t h e P r i n c e George a r e a , o c c u r s d u r i n g h u n t i n g s e a s o n . A l m o s t a l l h u n t e r a c c e s s t o moose i s t h r o u g h l o g g i n g r o a d s w h i c h means t h a t t h e g r e a t e s t d i s t u r b a n c e w i l l o c c u r w h e r e l o g g i n g h a s o c c u r r e d . Moose a v o i d a n c e of l o g g e d a r e a s , p a r t i c u l a r l y i n t h e f a l l , may r e s u l t f r o m moose a v o i d i n g human d i s t u r b a n c e . H u n t i n g was c l o s e d d u r i n g t h e s t u d y i n a c c e s s i b l e a r e a s b u t t h a t d i d n o t p r e v e n t t r a f f i c f r o m moving t h r o u g h t h o s e a r e a s . M y t t o n and K e i t h (1981) n o t e d moose a v o i d a n c e of humans even when moose were not hunted. O f t e n a r e s e a r c h e r w i l l assume t h a t t h e p a t t e r n of moose h a b i t a t o c c u p a t i o n b e i n g o b s e r v e d i s an i m m e d i a t e r e s p o n s e t o t h e c o n d i t i o n s p r e v a l e n t a t t h e t i m e c o m p a r i s o n s a r e b e i n g made. T h i s need not be t h e c a s e . F o r example, Gasaway e t a l . (1980) d e t e c t e d no d i f f e r e n c e between t h e home r a n g e s of moose up t o t h r e e y e a r s o f age and t h e i r dams. G i v e n s u c h a s t r o n g r e l a t i o n s h i p b e t w e e n moose and t h e i r dams, t h e a p p a r e n t s e l e c t i o n o f one h a b i t a t and a v o i d a n c e o f a n o t h e r c o u l d be a r e s u l t o f o f f s p r i n g l e a r n i n g a b e h a v i o u r p a t t e r n f r o m t h e i r dams w h i c h d i d not i n c l u d e an a r e a where a p a r t i c u l a r h a b i t a t was p r e v a l e n t . 118 4. F o r a g e Biomass E s t i m a t i o n T e c h n i q u e s 4.1 Shrub Biomass E s t i m a t i o n No t e c h n i q u e h a s b e e n d e v e l o p e d w h i c h c a n e c o n o m i c a l l y i n v e n t o r y s h r u b b i o m a s s p r o d u c t i o n a c c u r a t e l y and p r e c i s e l y ( P i t t and Schwab 1985). The r e g r e s s i o n e s t i m a t e of i n d i v i d u a l s h r u b p r o d u c t i o n c o m b i n e d w i t h a c o r r e c t e d p o i n t d i s t a n c e (CPD) e s t i m a t e o f s h r u b d e n s i t y ( B a t c h e l e r 1971) was c h o s e n b e c a u s e i t d e a l t w i t h some p r o b l e m s known t o a f f e c t s h r u b b i o m a s s e s t i m a t i o n . F o r example, an e x c e s s i v e amount of t i m e i s r e q u i r e d t o p r o d u c e p r e c i s e e s t i m a t e s when c l i p p i n g q u a d r a t e s ( S h a f e r 1963, B a r n e s 1976, H a r l o w 1977, R u t h e r f o r d 1979, T e l f e r 1981). A l s o , a stem d e n s i t y e s t i m a t e i s r e q u i r e d when a b r o w s e i n v e n t o r y i s b a s e d on r e g r e s s i o n e s t i m a t e s o f t h e p r o d u c t i o n of i n d i v i d u a l s tems. Some d i s t a n c e measurement e s t i m a t e s o f d e n s i t y p r o d u c e b i a s e d e s t i m a t e s o f d e n s i t y b e c a u s e t h e y a s s u m e t h a t s h r u b s a r e d i s t r i b u t e d r a n d o m l y ( L a y c o c k and B a t c h e l e r 1975, O l d e m e y e r and R e g e l i n 1 9 8 0 ) . S h r u b s a r e more l i k e l y t o h a v e a c l u m p e d d i s t r i b u t i o n t h a n a r a n d o m d i s t r i b u t i o n . The CPD e s t i m a t e o f s t e m d e n s i t y c o r r e c t s f o r d e p a r t u r e s f r o m randomness ( B a t c h e l e r 1971). T h e r e a r e , h o w e v e r , p r o b l e m s w i t h t h e u s e o f r e g r e s s i o n e s t i m a t e s o f i n d i v i d u a l s t e m p r o d u c t i o n c o m b i n e d w i t h a CPD e s t i m a t e of d e n s i t y . F o r t h e e s t i m a t i o n of t h e p r o d u c t i o n of 119 i n d i v i d u a l s h r u b s t h e e s t i m a t e s and v a r i a n c e were c a l c u l a t e d a s i f t h e y w e r e p r o d u c e d by a d o u b l e s a m p l e ( C o c h r a n 1 9 6 3 ) . I n f a c t t h e y w e r e n o t p r o d u c e d w i t h a d o u b l e s a m p l e . A d o u b l e s a m p l e w o u l d r e q u i r e t h a t p r e d i c t o r v a r i a b l e s be m e a s u r e d on e a c h s h r u b i n t h e s a m p l e and t h a t p r o d u c t i o n be m e a s u r e d on a s u b p o p u l a t i o n o f t h e s h r u b s m e a s u r e d . The r e g r e s s i o n of stem p r o d u c t i o n on p r e d i c t o r v a r i a b l e s would be c a l c u l a t e d f r o m t h e s u b p o p u l a t i o n . I n t h e m e t h o d u s e d t o i n v e n t o r y s h r u b p r o d u c t i o n , i n t h i s t h e s i s , p r e d i c t o r v a r i a b l e s w e r e m e a s u r e d on e a c h s h r u b i n t h e s a m p l e , b u t t h e r e g r e s s i o n was n o t c a l c u l a t e d f r o m a s u b p o p u l a t i o n o f t h e sample. The r e g r e s s i o n e q u a t i o n was c a l c u l a t e d f r o m a sample i n d e p e n d e n t of p l o t e s t i m a t e s of s h r u b b i o m a s s . The use o f a r e g r e s s i o n e q u a t i o n made up a t a d i f f e r e n t t i m e and i n many c a s e s w i t h s h r u b s f r o m a d i f f e r e n t p l a c e t h a n t h o s e o f i n t e r e s t must assume t h a t s u c h a r e g r e s s i o n makes an u n b i a s e d a s s e s s m e n t of t h e s h r u b s b e i n g i n v e n t o r i e d . T h i s a s s u m p t i o n i s n e v e r t r u e b e c a u s e s h r u b b i o m a s s and d i m e n s i o n s v a r y a c c o r d i n g t o a number of f a c t o r s w h i c h a r e l i n k e d t o t i m e and s i t e . F o r e x a m p l e , s h r u b b i o m a s s v a r i e s a c c o r d i n g t o p h e n o l o g y (Dean e t a l . 1981), s p e c i f i c s i t e c o n d i t i o n s ( T e l f e r 1 9 6 9 a, T e l f e r 1 9 6 9 b ) , a n n u a l g r o w i n g c o n d i t i o n s ( S t o n e and C r a w f o r d 1 9 8 1 ) , s e a s o n s a m p l e s a r e c o l l e c t e d ( T e l f e r 1969b, Bobek e t a l . 1979, P o t v i n 1981), and o v e r s t o r y canopy c l o s u r e 120 ( P e e k 1 9 7 0 ) . T h e r e w e r e r e a s o n s f o r t h e m e t h o d u s e d , i n t h i s t h e s i s , t o e s t i m a t e s h r u b b i o m a s s p r o d u c t i o n . F i r s t , a d o u b l e sample would have been f a r t o o t i m e c o n s u m i n g t o employ. Second, t h e m e t h o d u s e d a t t e m p t s t o o v e r c o m e s e a s o n a l d i f f e r e n c e s i n p r o d u c t i o n . T h a t i s , i f a p l o t were a c c u r a t e l y i n v e n t o r i e d i n m i d - J u n e and a g a i n a t t h e end o f A u g u s t , t h e n s u b s t a n t i a l d i f f e r e n c e s i n s h r u b p r o d u c t i o n w o u l d be r e c o r d e d . To overcome t h i s p r o b l e m a l l p r e d i c t o r v a r i a b l e s a r e r e l a t e d t o m i d s u m m e r s h r u b p r o d u c t i o n . A s s u m i n g t h a t t h e r e i s l e s s v a r i a t i o n i n p r e d i c t o r v a r i a b l e s t h a n i n t h e c u r r e n t a n n u a l g r o w t h borne by a s h r u b , t h e p r o b l e m w i t h r e c o r d i n g s p u r i o u s d i f f e r e n c e s among a r e a s i s r e d u c e d . N e v e r t h e l e s s , t h e a s s u m p t i o n t h a t t h e r e g r e s s i o n e s t i m a t e i s u n b i a s e d i s i m p o r t a n t . I f t h a t a s s u m p t i o n i s i n e r r o r t o a l a r g e d e g r e e , t h e n t h e v a l u e o f t h e s h r u b b i o m a s s e s t i m a t i o n d a t a a r e q u e s t i o n a b l e . F u r t h e r , t h e n a t u r a l l o g t r a n s f o r m a t i o n of t h e p r e d i c t e d v a r i a b l e , t o c o r r e c t f o r h e t e r o s c e d a s t i c i t y a n d c u r v i l i n e a r i t y , i n t r o d u c e s b i a s ( B a s k e r v i l l e 1965, Beauchamp and O l s s o n 1973, M o u n t f o r d and Bunce 1973). T h i s p r o b l e m i s , a p p a r e n t l y , o v e r c o m e by t h e a d d i t i o n o f a c o r r e c t i o n f a c t o r f o r b i a s ( e m s e / ^ 2 ) t o t h e b a c k t r a n s f o r m e d i n i t i a l e s t i m a t e ( M o u n t f o r d and Bunce 1973). 121 F i n a l l y , t h e l a c k o f an a p p r o p r i a t e b a c k t r a n s f o r m e d v a r i a n c e e s t i m a t e makes c o m b i n i n g t h e r e g r e s s i o n v a r i a n c e w i t h a d e n s i t y v a r i a n c e m a t h e m a t i c a l l y u n a c c e p t a b l e . I n t h e back t r a n s f o r m e d s c a l e , m e a s u r e s o f d i s p e r s i o n t h a t o r i g i n a t e i n t h e l o g a r i t h m i c s c a l e a r e not s y m m e t r i c . The method used i n t h i s s t u d y t o p r o d u c e an u n t r a n s f o r m e d v a r i a n c e o f a p p r o x i m a t e l y t h e r i g h t m a g n i t u d e does n o t i n f a c t p r o d u c e a m a t h e m a t i c a l l y c o r r e c t v a r i a n c e . A l t h o u g h CPD e s t i m a t e s o f s h r u b d e n s i t i e s a r e u n b i a s e d ( L a y c o c k and B a t c h e l e r 1975, O l d e m e y e r and R e g e l i n 1 9 8 0 ) , t h e r e a r e p r o b l e m s w i t h t h e e s t i m a t e s o f d e n s i t y v a r i a n c e b a s e d upon p r o b a b l e l i m i t s o f e r r o r ( P L E ) . A P L E i s an e m p i r i c a l e s t i m a t e o f e r r o r and not a m a t h e m a t i c a l l y r i g o r o u s c a l c u l a t i o n . T h i s s i t u a t i o n o f t e n r a i s e s q u e s t i o n s as t o t h e v a l i d i t y o f a P L E c a l c u l a t i o n . P L E c a l c u l a t i o n s h a v e b e e n shown t o o v e r e s t i m a t e t h e t r u e e r r o r o f a CPD e s t i m a t e o f d e n s i t y (Boyd 1979). 4.2 Herb L a y e r Biomass E s t i m a t i o n As compared w i t h t h e p r o b l e m s of s h r u b b i o m a s s e s t i m a t i o n t h e p r o b l e m s w i t h e s t i m a t i n g b i o m a s s i n t h e h e r b l a y e r a r e r e l a t i v e l y s t r a i g h t f o r w a r d . H e r b l a y e r b i o m a s s e s t i m a t i o n d o e s n o t h a v e t h e p r o b l e m , d i s c u s s e d u n d e r s h r u b b i o m a s s e s t i m a t i o n , o f r e g r e s s i o n e q u a t i o n s p r o d u c e d a t a d i f f e r e n t 122 t i m e a n d p l a c e f r o m t h e s a m p l e b e c a u s e t h e h e r b l a y e r p r o d u c t i o n e s t i m a t e s were p r o d u c e d w i t h a d o u b l e sample. Nor i s t h e r e a p r o b l e m w i t h a m a t h e m a t i c a l l y o b s c u r e c a l c u l a t i o n o f d e n s i t y v a r i a n c e b e c a u s e i t was not n e c e s s a r y t o e s t i m a t e t h e d e n s i t y of h e r b s . However, t h e r e g r e s s i o n e q u a t i o n s w h i c h p r e d i c t t h e b i o m a s s a v a i l a b l e i n a h a b i t a t were p r o d u c e d w i t h a n a t u r a l l o g t r a n s f o r m a t i o n of t h e p r e d i c t e d v a r i a b l e . T h i s t r a n s f o r m a t i o n i n c u r s t h e same d i f f i c u l t i e s w i t h b i a s i n t h e e s t i m a t e and l a c k of a back t r a n s f o r m e d v a r i a n c e e q u i v a l e n t t o t h e v a r i a n c e i n t h e n a t u r a l l o g s c a l e as were d i s c u s s e d under s h r u b b i o m a s s e s t i m a t i o n . 5. A s s o c i a t i o n I n d i c e s I f a s e t o f a n i m a 1 / h a b i t a t r e l a t i o n s h i p d a t a i s t o be i n t e r p r e t e d f o r management i m p o r t a n c e , t h e n c a r e must be t a k e n i n t h e u s e o f s e l e c t i o n d a t a p r o d u c e d w i t h an a s s o c i a t i o n i n d e x . I t r e q u i r e s a v e r y l a r g e p r o p o r t i o n o f l o c a t i o n s t o p r o d u c e a p o s i t i v e a s s o c i a t i o n , i f a h a b i t a t t y p e makes up a l a r g e p r o p o r t i o n o f t h e a v a i l a b l e a r e a . C o n v e r s e l y , i t i s e asy t o show n e u t r a l or n e g a t i v e a s s o c i a t i o n s even when t h e r e i s a r e l a t i v e l y l a r g e p r o p o r t i o n o f l o c a t i o n s i n s u c h h a b i t a t s . F o r e x a m p l e , u n l o g g e d f o r e s t made up 54.9% o f t h e a v a i l a b l e summer h a b i t a t . I n t h e summer o f 1982, u n l o g g e d f o r e s t was c r e d i t e d w i t h 76.7 moose l o c a t i o n s , or 54.8% of t h e 123 140 l o c a t i o n s r e c o r d e d . T h i s r e s u l t s i n a n o n s i g n i f i c a n t a s s o c i a t i o n . The number of l o c a t i o n s i n u n l o g g e d f o r e s t was more t h a n t h r e e t i m e s t h e number i n t h e h a b i t a t s e c o n d r a n k e d i n moose l o c a t i o n s . T he s e c o n d r a n k e d h a b i t a t r e c o r d e d a s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n . I f a h a b i t a t t y p e makes up a v e r y l a r g e p r o p o r t i o n o f a r e g i o n , t h e n a s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n b e t w e e n t h a t h a b i t a t and moose i s v e r y d i f f i c u l t t o p r o d u c e . On t h e o t h e r e x t r e m e , i t i s d i f f i c u l t t o show a n e g a t i v e r e l a t i o n s h i p b e t w e e n moose and a h a b i t a t m a k i n g up a s m a l l p r o p o r t i o n o f t h e a v a i l a b l e a r e a . F u r t h e r , few l o c a t i o n s a r e n e c e s s a r y t o show a p o s i t i v e r e l a t i o n s h i p . O l d c l e a r c u t s were 6.2% o f t h e a v a i l a b l e w i n t e r a r e a . I n m i d w i n t e r 1983, 7.7 l o c a t i o n s o f 1 6 9 , 4.5%, w e r e s c o r e d f o r o l d c l e a r c u t s , p r o d u c i n g a n o n s i g n i f i c a n t a s s o c i a t i o n . R e c e n t c l e a r c u t s were 7.3% o f t h e a v a i l a b l e s p r i n g a r e a . I n t h e s p r i n g o f 1983, 9 l o c a t i o n s of 67, 13.4%, were i n r e c e n t c l e a r c u t s , p r o d u c i n g a s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n . A s s o c i a t i o n o r s e l e c t i o n i n d i c e s d e s c r i b e d e p a r t u r e s f r o m randomness. They do not measure management v a l u e , nor do t h e y d e f i n e i t . Management s i g n i f i c a n c e m i g h t w e l l be based on the a b s o l u t e p e r c e n t a g e o f a n i m a l l o c a t i o n s i n a p a r t i c u l a r h a b i t a t . The s e c o n d p r o b l e m i s t h a t i n t h i s s t u d y , s e a s o n s w e r e d e l i m i t e d by o b s e r v e d h a b i t a t s h i f t s by m o o s e . T h e 124 o b j e c t i v i t y o f t h e a s s o c i a t i o n i n d e x was r e d u c e d by t h e c i r c u l a r i t y o f d o c u m e n t i n g c h anges i n h a b i t a t a l r e a d y known t o e x i s t . I t was n e c e s s a r y t o d e l i m i t s e a s o n s on t h e b a s i s o f moose b e h a v i o u r b e c a u s e i n s u f f i c i e n t d a t a w e r e g e n e r a t e d t o c a l c u l a t e h a b i t a t a s s o c i a t i o n s on a m o n t h l y b a s i s . F u r t h e r , i t w ould have been i n a p p r o p r i a t e t o v i e w h a b i t a t a s s o c i a t i o n s m o n t h by m o n t h , a s many c a l e n d a r m o n t h s c o n t a i n e d t w o d i f f e r e n t k i n d s o f moose h a b i t a t s e l e c t i o n b e h a v i o u r . A f u r t h e r p r o b l e m i s t h e q u e s t i o n o f a v a i l a b i l i t y . I n t h e l a t e w i n t e r and s p r i n g o f 1982, d i a m e t e r l i m i t l o g g i n g s c o r e d p o s i t i v e a s s o c i a t i o n s w i t h moose l o c a t i o n s . I t i s p o s s i b l e , h o w e v e r , t h a t t h e s e p o s i t i v e a s s o c i a t i o n s w e r e c a u s e d by moose t r a p p e d on t h e s o u t h s i d e of t h e F r a s e r R i v e r by i c e c o n d i t i o n s . M o s t o f t h e i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g , o l d and r e c e n t c l e a r c u t s were not a v a i l a b l e t o t h e s e moose. A more s e r i o u s p r o b l e m w i t h a v a i l a b i l i t y i s t h a t t h e o b s e r v e r d e c i d e s what i s a v a i l a b l e . The o b s e r v e r t r i e s t o s e l e c t t h e a r e a a v a i l a b l e s u c h t h a t t h e r e p r e s e n t a t i v e a s s o c i a t i o n i n d i c e s a r e d e v e l o p e d . In t h e c a s e of t h i s s t u d y , t h e r e was no p r o b l e m w i t h t h e s p r i n g t h r o u g h f a l l r a n g e s , as t h e a r e a d e f i n e d as a v a i l a b l e e n c o m p a s s e d a v e r y b r o a d a r e a c o n t a i n i n g a l l moose l o c a t i o n s . The a r e a i n w h i c h moose w i n t e r , however, was much s m a l l e r t h a n th e b r o a d e r n o n - w i n t e r 125 a r e a . The w i n t e r r a n g e ne a r t h e F r a s e r R i v e r was d o m i n a t e d by l o g g e d a r e a s , w h e r e a s away f r o m t h e r i v e r , u n l o g g e d f o r e s t d o m i n a t e d . I f one s e l e c t e d a v e r y n a r r o w a v a i l a b l e w i n t e r i n g a r e a , l o g g e d a r e a s would n e v e r show p o s i t i v e a s s o c i a t i o n s w i t h moose. I f a v e r y l a r g e a v a i l a b l e w i n t e r i n g a r e a were cho s e n , u n l o g g e d f o r e s t would n e v e r show p o s i t i v e a s s o c i a t i o n s . The a v a i l a b l e w i n t e r a r e a c h o s e n was a b r o a d s w a t h o v e r t h e m i d w i n t e r and l a t e w i n t e r l o c a t i o n s . W i n t e r r a n g e d e f i n e d on t h e b a s i s o f m i d w i n t e r a n d l a t e w i n t e r moose l o c a t i o n s c r e a t e d p r o b l e m s w i t h t h e c a l c u l a t i o n o f h a b i t a t a s s o c i a t i o n s i n e a r l y w i n t e r . M o o s e s t a r t e d t h e e a r l y w i n t e r p e r i o d as w i d e s p r e a d a s t h e y w e r e i n s p r i n g t h r o u g h f a l l . A t t h e end o f t h a t s e a s o n , moose w e r e a s n a r r o w l y d i s t r i b u t e d a s t h e y w e r e d u r i n g t h e r e s t o f t h e w i n t e r . The b r o a d s p r i n g , summer, and f a l l r e g i o n was c h o s e n as t h e a r e a a v a i l a b l e d u r i n g e a r l y w i n t e r . I f t h e m i d w i n t e r a n d l a t e w i n t e r r e g i o n h a d b e e n c h o s e n a s t h e a r e a a v a i l a b l e t o moose, t h e a s s o c i a t i o n i n d i c e s w o u l d h a v e b e e n much d i f f e r e n t t h a n shown. I n s u m m a r y , t h e r e a r e w e a k n e s s e s i n t h e u s e o f a s s o c i a t i o n i n d i c i e s t o d o c u m e n t s e l e c t i o n o r a v o i d a n c e o f b o t h common and r a r e h a b i t a t t y p e s . I t i s d i f f i c u l t t o d e m o n s t r a t e s e l e c t i o n o f common h a b i t a t , b u t e a s y t o show s e l e c t i o n f o r r a r e h a b i t a t s . I f s e l e c t i o n d a t a a r e b e i n g i n t e r p r e t e d t o i n d i c a t e management i m p o r t a n c e , c a r e s h o u l d be 126 t a k e n i n t h e i n t e r p r e t a t i o n o f n e g a t i v e and n o n s i g n i f i c a n t s e l e c t i o n o f a h a b i t a t t h a t o c c u p i e s a l a r g e p r o p o r t i o n of t h e a v a i l a b l e a r e a . I t may be t h a t t h e l a c k o f p o s i t i v e s e l e c t i o n i n d i c e s may be due t o t h e i n a b i l i t y of t h e s e l e c t i o n i n d e x t o p r o d u c e a p o s i t i v e v a l u e f o r t h a t h a b i t a t . I f , h o w e v e r , one w i s h e s t o d i s c o u n t t h e p o s s i b i l i t y t h a t moose a r e o c c u p y i n g h a b i t a t a t random, t h e n some s e l e c t i o n or a s s o c i a t i o n i n d e x i s n e c e s s a r y . 6. N o n f o r e s t H a b i t a t s T h e r e w e r e f i v e n o n f o r e s t h a b i t a t t y p e s r e c o g n i z e d i n t h i s t h e s i s : u p l a n d s h r u b , r i p a r i a n s h r u b , w e t l a n d s , l a k e s , and t h e F r a s e r R i v e r . The f o l l o w i n g i s an a s s e s s m e n t o f t h e r e l a t i o n s h i p b e t w e e n e a c h h a b i t a t and moose b a s e d upon c a l c u l a t i o n s o f t h e C o l e i n d e x o f a s s o c i a t i o n ( F i g u r e s 12, 13, and 1 4 ) , t h e number o f l o c a t i o n s i n a h a b i t a t ( T a b l e 16), and o b s e r v a t i o n s made d u r i n g v i s i t s t o p l o t s r e p r e s e n t a t i v e o f t h e s e h a b i t a t t y p e s . 6.1 U p l a n d Shrub U p l a n d s h r u b h a t i t a t s a r e f o r t h e most p a r t dense g r o w t h s o f a l d e r ( A l n u s v i r i d i s ) a n d b r a c k e n f e r n ( P t e r i d i u m 127 a_£_ui_linum), b u t w i t h a few p a l a t a b l e s h r u b s . F o r t h e m o s t p a r t , t h e s e a r e a s a r e l i t t l e u s e d by moose p r o b a b l y b e c a u s e t h e g r o w t h i s so t h i c k t h a t i t w o u l d r e q u i r e a l o t o f e f f o r t t o p e n e t r a t e . H o w e v e r , w h e r e a b r e a k i n t h e g r o w t h o c c u r s , moose w i l l u s e t h e s e a r e a s as t r a v e l c o r r i d o r s and h e a v i l y used t r a i l s d e v e l o p . F o r e s t e d edges a r e a l s o h e a v i l y used by moose p r o b a b l y b o t h as t r a v e l c o r r i d o r s and t o e x p l o i t a heavy g r o w t h of p a l a t a b l e f o r b s and browse w h i c h o f t e n o c c u r s a t t h e i n t e r f a c e b e t w e e n a l d e r t h i c k e t s and f o r e s t . The p o s i t i v e a s s o c i a t i o n d o c u m e n t e d b e t w e e n moose l o c a t i o n s and u p l a n d s h r u b h a b i t a t , i n some s e a s o n s ( F i g u r e s 12, 13, and 14) was p r o b a b l y t h e r e s u l t o f moose u s i n g t h e f o r e s t e d e d g e s o f t h e a l d e r t h i c k e t s . T h e r e was a h i g h and o f t e n s i g n i f i c a n t l y p o s i t i v e a s s o c i a t i o n between moose l o c a t i o n s and u p l a n d s h r u b i n f a l l ( F i g u r e s 12, 13 and 14). T h i s may i n d i c a t e t h a t a l d e r t h i c k e t s w e r e i m p o r t a n t d u r i n g r u t t i n g a c t i v i t y , o r p e r h a p s a l d e r o r some o t h e r f o r a g e i n t h e a l d e r p a t c h e s i s i m p o r t a n t f o r f a l l n u t r i t i o n . F o r e x a m p l e , t h e r u m e n o f a b u l l moose e x a m i n e d i n O c t o b e r 1982 a p p e a r e d t o c o n t a i n a l d e r l e a v e s a l m o s t e x c l u s i v e l y . However, a more b r o a d l y b ased e x a m i n a t i o n o f moose f o o d h a b i t s i n t h e P r i n c e George a r e a ( Eastman 1978) c o n t a i n e d no i n d i c a t i o n of moose p r e f e r e n c e f o r a l d e r i n the f a l l . 128 F i g u r e 12. C o l e (Cy) i n d e x of a s s o c i a t i o n between moose l o c a t i o n s and n o n f o r e s t h a b i t a t t y p e s f r o m s p r i n g 1980 t o f a l l 1980. O p e n a n d c l o s e d c i r c l e s r e p r e s e n t n o n s i g n i f i c a n t a s s o c i a t i o n s and s i g n i f i c a n t a s s o c i a t i o n s r e s p e c t i v e l y . 0 . 3 X - 0 . 3 <D O O O - 0 . 6 - 0 . 9 •1.2 O sprlng — • / — o o summer fall t o early winter midwinter upland shrub  r i p a r i a n shrub wet lands lakes Fraser River 130 F i g u r e 13 C o l e (Cy) i n d e x of a s s o c i a t i o n between moose l o c a t i o n s and n o n f o r e s t h a b i t a t t y p e s f r o m s p r i n g 1981 t o l a t e w i n t e r 1982. O p e n a n d c l o s e d c i r c l e s r e p r e s e n t n o n s i g n i f i c a n t a s s o c i a t i o n s and s i g n i f i c a n t a s s o c i a t i o n s r e s p e c t i v e l y . c o spring summer fall early winter midwinter late winter upland shrub  r i p a r i a n shrub wet lands lakes Fraser River 132 F i g u r e 14 C o l e ( C y ) i n d e x o f a s s o c i a t i o n b e t w e e n moose l o c a t i o n s and n o n f o r e s t h a b i t a t t y p e s f r o m s p r i n g 1982 t o s p r i n g 1983. O p e n a n d c l o s e d c i r c l e s a s s o c i a t i o n s and s i g n i f i c a n t r e p r e s e n t a s s o c i a t i o n s n o n s i g n i f i c a n t r e s p e c t i v e l y . upland shrub  r i p a r i a n shrub wet lands lakes Fraser R iver 134 T a b l e 16. Number of moose l o c a t i o n s i n n o n f o r e s t h a b i t a t s by y e a r and season Season Upland Shrub R i p a r i a n Shrub W e t l a n d s L a k e s F r a s e r R i v e r T o t a l A] H a b i t a t 1980 1 . 0 1 . 8 6 . 3 0 . 5 0 . 0 5 7 . 0 S p r i n g 1981 3 . 0 2 . 1 4 . 3 0 . 0 0 . 0 4 2 . 0 1982 5 . 2 2 . 8 4 . 7 0 . 0 0 . 3 7 5 . 0 1983 7.7 1 . 0 2 . 8 0 . 0 0 . 5 6 7 . 0 1980 1 . 5 1 . 0 3 . 3 0 . 0 0 . 0 3 1 . 0 Summer 1981 5 . 3 4 . 3 5 . 7 0 . 5 0 . 0 6 4 . 0 1982 1 0 . 5 4 . 2 1 0 . 3 0 . 0 0 . 0 1 4 0 . 0 1980 1 . 3 1 . 5 3 . 3 0 . 5 0 . 0 3 1 . 0 F a l l 1981 4 . 0 1 . 0 3 . 0 0 . 0 0 . 0 3 1 . 0 1982 1 1 . 0 4 . 2 6 . 2 0 . 0 0 . 0 8 2 . 0 E a r l y 1981-82 3 . 0 0 . 8 2 . 8 0 . 0 0 . 0 3 9 . 0 W i n t e r 1982-83 1 . 5 2 . 3 2 . 8 0 . 0 0 . 0 6 6 . 0 M i d w i n t e r 1982 0 . 5 3 . 5 2 . 3 0 . 0 4 . 7 1 0 2 . 0 1983 1 . 3 6 . 0 4 . 7 0 . 0 0 . 0 1 6 9 . 0 L a t e 1982 4 . 7 5 . 0 3 . 7 0 . 0 4 . 8 1 6 2 . 0 W i n t e r 1983 4 . 0 0 . 5 2 . 0 0 . 0 0 . 0 7 5 . 0 1 3 5 6 . 2 R i p a r i a n S h r u b I n g e n e r a l , t h e r e w e r e t w o k i n d s o f r i p a r i a n s h r u b h a b i t a t s e x a m i n e d d u r i n g t h i s s t u d y : t h e f i r s t d o m i n a t e d by a d e n s e g r o w t h o f w i l l o w ( S a l i x s p p . ) w i t h m i n o r c o m p o n e n t s o f r e d - o s i e r d o g w o o d ( C o r n u s s e r i c e a ) a n d s p i r e a ( S p i r a e a s p p . ) . T h e s e c o n d h a d a n o v e r s t o r y o f a l d e r ( A l n u s i H ^ j L C J i ) w i t h a s u b s t a n t i a l g r o w t h o f b l a c k t w i n b e r r y ( L o n i c e r a i n v o l u c r a t a ) a n d o s t r i c h f e r n ( ^ i i t e u c e i a js £ £ u ^ h d^££ t_ e_ r ^ j g ) ± n t h e u n d e r s t o r y . T h e t h i c k g r o w t h o n t h e r i p a r i a n s h r u b h a b i t a t s a p p e a r e d t o d e t e r m o o s e u s e o f t h e s e a r e a s . T h e a s s o c i a t i o n o f m o o s e l o c a t i o n s w i t h r i p a r i a n s h r u b h a b i t a t w a s r a r e l y o t h e r t h a n n e u t r a l ( F i g u r e s 1 2 , 1 3 , a n d 1 4 ) . H o w e v e r , t h e r e w a s a s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n b e t w e e n m o o s e l o c a t i o n s a n d r i p a r i a n s h r u b d u r i n g t h e s u m m e r o f 1 9 8 1 a n d t h e f a l l o f 1 9 8 2 . D e s p i t e t h e l a c k o f c o n s i s t e n t p o s i t i v e a s s o c i a t i o n s b e t w e e n m o o s e a n d r i p a r i a n s h r u b , t h i s h a b i t a t p r o b a b l y c o n t r i b u t e s a g r e a t d e a l o f f o r a g e t o m o o s e r e s i d e n t i n a d j a c e n t f o r e s t s . On f o r e s t e d g e s , u s e b y m o o s e w a s v e r y h e a v y a n d w h e r e p a s s a g e i n t o t h e r i p a r i a n a r e a s w a s f a i r l y e a s y t h e r e w a s a b u n d a n t l e a f s t r i p p i n g o f b l a c k t w i n b e r r y a n d o s t r i c h f e r n . T h e r e w a s a l s o h e a v y u s e o f t h e e d g e s o f r i p a r i a n h a b i t a t s i n w i n t e r w h e n m o o s e h a d a c c e s s t o t h e m o v e r t h e i c e o n c r e e k s o r t h e F r a s e r R i v e r . 1 3 6 6 . 3 W e t l a n d s W e t l a n d s w e r e c h a r a c t e r i z e d a s a r e a s w i t h s o i l s s a t u r a t e d w i t h m o i s t u r e . A r e a s w i t h o p e n w a t e r a n d b l a c k s p r u c e ( P i c e a m a r i a n a ) f o r e s t w e r e n o t i n c l u d e d u n d e r t h i s c l a s s i f i c a t i o n . W e t l a n d s i n c l u d e d a r e a s c o v e r - e d b y h e a v y m a t s o f m o s s w i t h l i t t l e s e d g e a n d f o r b g r o w t h t o a r e a s d o m i n a t e d b y b o g b i r c h ( B ^ t i i J L a g l a n d u l o s a ) . O c c a s i o n a l l y s m a l l g r o u p s o f t r e e s , l o d g e p o l e p i n e (P_i n u s^ c o n t o r t a ) o r b l a c k s p r u c e , g r e w o n s l i g h t e l e v a t i o n s i n t h e s o i l . M o o s e m a d e a l o t o f u s e o f w e t l a n d s d u r i n g t h e s t u d y . V i s i t s t o w e t l a n d s n o r m a l l y s h o w e d n u m e r o u s f r e s h t r a c k s i n t h e w e t l a n d i t s e l f a n d a l s o i n t h e f o r e s t i m m e d i a t e l y s u r r o u n d i n g t h e w e t l a n d . M o o s e u s e o f w e t l a n d s a p p e a r e d t o b e d i r e c t e d t o w a r d g r a z i n g b u c k b e a n ( M e n j r a n t h e s t ^ ^ f _ £ 3 . _ i j a _ t a ) w h i c h t h e y u p r o o t e d f r o m t h e s o i l . A n y t a l l g r o w i n g w i l l o w w h i c h o c c u r r e d i n a w e t l a n d w a s h e a v i l y b r o w s e d . W e t l a n d s s u r r o u n d e d b y c l e a r c u t s d i d n o t s h o w m u c h u s e b y m o o s e . W e t l a n d s w a s t h e o n l y n o n f o r e s t h a b i t a t t y p e w h i c h c o n s i s t e n t l y s h o w e d s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n s w i t h m o o s e l o c a t i o n s ( F i g u r e s 1 2 , 13 a n d 1 4 ) p a r t i c u l a r l y i n s p r i n g , s u m m e r , a n d f a l l . . 6 . 4 L a k e s A l t h o u g h m o o s e w e r e s t a n d i n g b y s m a l l l a k e s , o c c a s i o n a l l y s e e n s w i m m i n g i n o r l a k e s w e r e c r e d i t e d w i t h o n l y 1 . 5 137 l o c a t i o n s d u r i n g t h e c o u r s e o f t h e s t u d y ( T a b l e 1 6 ) . T h i s i s u n u s u a l c o n s i d e r i n g t h e n u m b e r o f r e p o r t s o f m o o s e f e e d i n g o n s o d i u m r i c h e m e r g e n t a n d s u b m e r g e n t a q u a t i c v e g e t a t i o n d u r i n g t h e s p r i n g a n d s u m m e r ( K n o r r e 1 9 5 9 , R i t c e y a n d V e r b e e k 1 9 6 9 , V a n B a l l e n b e r g e a n d P e e k 1 9 7 1 , P e e k e t a l . 1 9 7 6 , B e l o v s k y 1 9 7 8 , F r a s e r e t a l . 1 9 8 4 ) . H o w e v e r , m o s t o f t h e s m a l l l a k e s o n t h e s t u d y a r e a w e r e s u r r o u n d e d b y a f l o a t i n g m a t o f m o s s . E m e r g e n t a n d s u b m e r g e n t a q u a t i c s w e r e r a r e e v e n w h e n t h e y d i d o c c u r . T h e l a c k o f t h e h a b i t a t f e a t u r e t h o u g h t t o b e m o s t d e s i r a b l e t o m o o s e i n s m a l l l a k e s may e x p l a i n t h e c o n s i s t e n t n e g a t i v e a s s o c i a t i o n o f m o o s e l o c a t i o n s w i t h s m a l l l a k e s ( F i g u r e s 1 2 , 1 3 , a n d 1 4 ) . When a s m a l l l a k e w a s s u r r o u n d e d by f o r e s t , t h e r e w a s a l m o s t i n v a r i a b l y a h e a v i l y u s e d m o o s e t r a i l r i n g i n g t h e l a k e . 6 . 5 F r a s e r R i v e r W i t h t h e e x c e p t i o n o f m i d w i n t e r a n d l a t e w i n t e r 1 9 8 2 t h e r e w e r e f e w m o o s e l o c a t i o n s a s s o c i a t e d w i t h t h e F r a s e r R i v e r ( T a b l e 1 6 ) . H o w e v e r , d u r i n g m i d - a n d l a t e w i n t e r 1 9 8 2 t h e r e w e r e s u f f i c i e n t m o o s e l o c a t i o n s o n o r n e a r t h e F r a s e r t o p r o d u c e a s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n b e t w e e n m o o s e l o c a t i o n s a n d t h e r i v e r . T h e u s e o f t h e r i v e r i n m i d - a n d l a t e w i n t e r 1 9 8 2 w a s g r e a t e r t h a n i n d i c a t e d i n T a b l e 16 a n d F i g u r e 1 3 . M o s t c o l l a r e d m o o s e o b s e r v e d o n t h e r i v e r w e r e 138 n e a r t h e r i v e r ' s e d g e . A m o o s e l o c a t e d o n t h e r i v e r ' s e d g e w o u l d h a v e t o s h a r e t h e l o c a t i o n b e t w e e n t h e r i v e r a n d a t l e a s t o n e o t h e r h a b i t a t t y p e . T h e r e a s o n t h e F r a s e r R i v e r w a s u s e d r e l a t i v e l y h e a v i l y i n t h e w i n t e r o f 1 9 8 2 , p a r t i c u l a r l y t h e e a r l y p a r t o f m i d w i n t e r 1 9 8 2 , w a s t h a t w h i l e m o o s e w e r e f l o u n d e r i n g i n t h e d e e p s o f t s n o w o n l a n d t h e y w e r e a b l e t o m o v e f r e e l y o n t h e r i v e r i c e . M o o s e h o o v e s r a r e l y p e n e t r a t e d m o r e t h a n 4 5 cm i n t o s n o w o n t h e r i v e r , w h i l e t h e y w e r e s t r u g g l i n g w i t h s n o w 30 cm d e e p e r t h a n t h e i r c h e s t . h e i g h t s o n l a n d . 7 . M o o s e H a b i t a t M a n a g e m e n t 7 . 1 M a n a g e m e n t I m p l i c a t i o n s o f R e s u l t s I n 1 9 7 8 B e l o v s k y a n d J o r d a n p r o d u c e d t h e f i r s t p a p e r p u b l i s h e d o n N o r t h A m e r i c a n m o o s e b e h a v i o u r w h i c h i l l u s t r a t e d t h e i m p o r t a n c e t o m o o s e o f a v o i d i n g h e a t s t r e s s i n s u m m e r . W i t h a s u c c e e d i n g p a p e r ( B e l o v s k y 1 9 8 1 ) , t h e h y p o t h e s i s o f m o o s e a v o i d a n c e o f s u m m e r h e a t s t r e s s b e c a m e e s t a b l i s h e d . R e n e c k e r a n d H u d s o n ( 1 9 8 4 ) f o u n d m o o s e s e n s i t i v e e v e n t o r e l a t i v e l y l o w t e m p e r a t u r e s i n s u m m e r . M o o s e p a n t e d i n a i r t e m p e r a t u r e s e x c e e d i n g 2 0 ° . T h e h e a t s t r e s s a v o i d a n c e h y p o t h e s i s i s n o t g e n e r a l l y a c c e p t e d , h o w e v e r , a s many a u t h o r s h a v e o b s e r v e d m o o s e i n o p e n a r e a s t h a t p r o d u c e a b u n d a n t f o r a g e o r b e l i e v e t h a t a c c e s s t o q u a l i t y f o r a g e e x p l a i n s e n t i r e l y 139 s u m m e r m o o s e b e h a v i o u r ( s e e l i t e r a t u r e r e v i e w ) . T h e d a t a p r o d u c e d i n t h i s t h e s i s i n d i c a t e t h a t m o o s e b e h a v e a s i f a v o i d a n c e o f h e a t s t r e s s i s i m p o r t a n t i n s u m m e r : T h e r e w a s a s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n b e t w e e n m o o s e a n d u n l o g g e d f o r e s t i n t h e s u m m e r o f 1 9 8 1 , w h e n a i r t e m p e r a t u r e s g e n e r a l l y e x c e e d e d 2 0 ° . M o r e o p e n h a b i t a t s , c h a r a c t e r i z e d by i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g , w e r e s e l e c t e d i n t h e s u m m e r 1 9 8 2 , w h e n t e m p e r a t u r e s d i d n o t g e n e r a l l y e x c e e d 2 0 ° ( F i g u r e s 5 a n d 6 ) . T h e h a b i t a t s e l e c t e d b y m o o s e i n t h e w a r m s u m m e r , u n l o g g e d f o r e s t , h a d t h e l o w e s t f o r a g e p r o d u c t i o n ( 2 0 . 6 0 g/m T a b l e 5 ) o f a n y f o r e s t / l o g g e d h a b i t a t . I n t h e c o o l s u m m e r , 1 9 8 2 , a l a r g e p r o p o r t i o n o f m o o s e l o c a t i o n s w e r e a g a i n r e c o r d e d i n u n l o g g e d f o r e s t , b u t n o t i n s u f f i c i e n t n u m b e r s t o p r o d u c e a s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n . R e g a r d l e s s o f w h e t h e r s u m m e r t e m p e r a t u r e s w e r e w a r m o r c o o l , h a b i t a t w i t h a b u n d a n t f o r a g e , o l d c l e a r c u t s ( 5 1 . 5 7 g / m 2 ) , w e r e a v o i d e d ( F i g u r e 5 , F i g u r e 6 , T a b l e 3 , a n d T a b l e 4 ) . T h e r e i s a p o p u l a r p e r c e p t i o n t h a t i n w i n t e r m o o s e s e l e c t h a b i t a t w h i c h p r o v i d e s t h e m o s t f o r a g e e x c e p t w h e n s n o w d e p t h s e x c e e d a c r i t i c a l d e p t h ( s e e l i t e r a t u r e r e v i e w ) . T h e c r i t i c a l d e p t h i s o f t e n s e t a t 9 0 c m . T h e h y p o t h e s i s t h a t m o o s e a r e a s v u l n e r a b l e t o h e a t s t r e s s i n w i n t e r a s i n s u m m e r h a s o n l y r e c e n t l y b e e n s u g g e s t e d b y R e n e c k e r a n d H u d s o n ( 1 9 8 4 ) . 140 Snow d e p t h s were w e l l i n e x c e s s of 90 cm i n a l l h a b i t a t s i n m i d - and l a t e w i n t e r 1982, b u t were much l e s s t h a n 90 cm i n t h e w i n t e r of 1983. Moose s h o u l d have s e l e c t e d h a b i t a t on t h e b a s i s of c o v e r i n 1982 but on t h e b a s i s of f o r a g e abundance i n 1983, a c c o r d i n g t o t h e f o r a g e p r o d u c t i o n c o n d i t i o n a l upon snow de p t h h y p o t h e s i s . Moose s h o u l d have s e l e c t e d h a b i t a t on t h e b a s i s o f c o v e r i n b o t h w i n t e r s a c c o r d i n g t o t h e a v o i d a n c e o f h e a t s t r e s s h y p o t h e s i s b e c a u s e t h e r m a l c o n d i t i o n s e x c e e d e d moose upper c r i t i c a l t e m p e r a t u r e s i n b o t h y e a r s . The d a t a i n t h i s t h e s i s a r e more c o n s i s t e n t w i t h t h e a v o i d a n c e o f h e a t s t r e s s h y p o t h e s i s t h a n w i t h t h e f o r a g e abundance c o n d i t i o n a l u p on snow d e p t h h y p o t h e s i s . I n m i d w i n t e r 1982 f o r e s t c o v e r was more h i g h l y c o r r e l a t e d w i t h a s s o c i a t i o n s i n d i c e s t h a n was f o r a g e a b u n d a n c e . The s i t u a t i o n i n m i d w i n t e r 1982 was e x p e c t e d ( T a b l e 7). In m i d w i n t e r 1983 t h e c o r r e l a t i o n between f o r e s t c o v e r and a s s o c i a t i o n i n d i c e s was h i g h e r t h a n t h e c o r r e l a t i o n between f o r a g e abundance and a s s o c i a t i o n i n d i c e s . The s i t u a t i o n i n 1983 was n o t e x p l i c a b l e u n l e s s moose were s e e k i n g s h e l t e r a s w e l l a s f o r a g e . A l s o , t h e c o r r e l a t i o n between f o r e s t c o v e r and a s s o c i a t i o n i n d i c e s was much g r e a t e r t h a n t h a t between f o r a g e abundance and a s s o c i a t i o n i n d i c e s i n b o t h l a t e w i n t e r p e r i o d s . T h e s e c o r r e l a t i o n s i n d i c a t e t h a t moose s o u g h t s h e l t e r r a t h e r t h a n f o r a g e i n l a t e w i n t e r . F u r t h e r , i n t o t a l t h e r e were p r o p o r t i o n a l l y more moose l o c a t e d 141 i n u n l o g g e d f o r e s t d u r i n g m i d - and l a t e w i n t e r s o f s h a l l o w snow, 1983, t h a n d u r i n g m i d and l a t e w i n t e r s o f deep snow, 1982 ( T a b l e 3, T a b l e 12, and T a b l e 13). The d a t a p r o d u c e d i n t h i s t h e s i s i n d i c a t e t h a t s h e l t e r f r o m h e a t s t r e s s i s an i m p o r t a n t c o n s i d e r a t i o n f o r moose h a b i t a t d u r i n g b o t h s u m m e r a n d w i n t e r . T h e r e a r e t w o i m p o r t a n t i m p l i c a t i o n s of t h i s f i n d i n g : (1) Q u a l i t y summer h a b i t a t i s based on more t h a n abundant f o r a g e . Summer h a b i t a t must a l s o p r o v i d e s h e l t e r f r o m h e a t s t r e s s . (2) Moose w i n t e r h a b i t a t i s b e t t e r managed t h a n summer h a b i t a t b e c a u s e i t has b e e n a s s u m e d t h a t m o o s e r e q u i r e m a t u r e t r e e s on t h e i r w i n t e r i n g a r e a s t o r e d u c e snow d e p t h s and p r o t e c t t hem f r o m c o l d s t r e s s . Many of t h e o b s e r v a t i o n s w h i c h s u p p o r t e d t h e s e c o n c e p t s may have r e s u l t e d f r o m m i s i n t e r p r e t i n g moose h a b i t a t s e l e c t i o n c a u s e d by h e a t s t r e s s . I t i s i m p o r t a n t t o u n d e r s t a n d t h e e n v i r o n m e n t a l i n f l u e n c e s w h i c h c a u s e a p a r t i c u l a r b e h a v i o u r b e c a u s e moose p r o b a b l y r e q u i r e m a t u r e t r e e s f o r s h e l t e r f r o m h e a t s t r e s s r e g a r d l e s s of t h e d e p t h of snow a c c u m u l a t i o n . T hese s t a t e m e n t s s h o u l d not be i n t e r p r e t e d as i n d i c a t i n g t h a t c o n s i d e r a t i o n s o t h e r t h a n s h e l t e r f r o m h e a t s t r e s s a r e not i m p o r t a n t . The p r o v i s i o n of a d e q u a t e f o r a g e i n summer and w i n t e r , and s h e l t e r f r o m deep snow and c o l d s t r e s s i n w i n t e r s h o u l d a l s o be c o n s i d e r e d i n t h e management o f moose h a b i t a t . 142 F u r t h e r , w i t h o u t t h e c h o i c e o f h a b i t a t w i t h f o r e s t c o v e r m o o s e w o u l d p o s s i b l y o c c u p y h a b i t a t i n w h i c h t h e r m a l s t r e s s w a s a f r e q u e n t o c c u r r e n c e . B a s e d o n t h e o b s e r v a t i o n s o f K n o r r e ( 1 9 5 9 ) , M y t t o n a n d K e i t h ( 1 9 8 1 ) , a n d R e n e c k e r a n d H u d s o n ( 1 9 8 4 ) t h e s e a n i m a l s w o u l d p r o b a b l y h a v e l o w e r g r o w t h a n d r e p r o d u c t i o n r a t e s t h a n m o o s e a b l e t o a v o i d t h e r m a l s t r e s s . 7 . 2 M a n a g e m e n t I m p o r t a n c e o f S p e c i f i c H a b i t a t s 7 . 2 . 1 U n l o g g e d F o r e s t M a n y a u t h o r s h a v e w r i t t e n a b o u t t h e n e e d t o r e m o v e f o r e s t t o s t i m u l a t e t h e p r o d u c t i o n o f p a l a t a b l e s h r u b s ( s e e l i t e r a t u r e r e v i e w ) . I n f a c t t h e o b s e r v a t i o n t h a t a b u n d a n t f o r a g e i s l i m i t e d i n f o r e s t w a s s u p p o r t e d b y t h e d a t a c o l l e c t e d f o r t h i s t h e s i s ( T a b l e s 5 , 1 0 , 1 1 , 1 4 , a n d 1 5 ) . T h e e x p e c t a t i o n t h a t t h i s p h e n o m e n o n s h o u l d r e s u l t i n u n l o g g e d f o r e s t b e i n g a v o i d e d i n f a v o u r o f h a b i t a t s b e a r i n g g r e a t e r f o r a g e w a s n o t s u p p o r t e d by t h e m o o s e l o c a t i o n d a t a . U n l o g g e d f o r e s t r e c o r d e d a s m a n y s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n s ( s e v e n ) w i t h m o o s e l o c a t i o n s a s a n y o t h e r f o r e s t / l o g g e d h a b i t a t t y p e ( F i g u r e s 4 , 5 , 6 , a n d T a b l e 4 ) . M o o s e w e r e m o s t c l o s e l y a s s o c i a t e d w i t h u n l o g g e d f o r e s t i n s u m m e r , f a l l , m i d w i n t e r , a n d l a t e w i n t e r . N e g a t i v e a s s o c i a t i o n s o c c u r r e d i n s p r i n g a n d e a r l y w i n t e r . 1 4 3 T h e m a n a g e m e n t i m p o r t a n c e o f f o r e s t i s i l l u m i n a t e d m o r e c l e a r l y b y t h e p r o p o r t i o n o f t o t a l l o c a t i o n s i n u n l o g g e d f o r e s t t h a n by c a l c u l a t i n g w h e t h e r m o o s e o c c u p i e d f o r e s t a t a g r e a t e r o r l e s s e r r a t e t h a n e x p e c t e d a t r a n d o m . U n l o g g e d f o r e s t made up a l a r g e p r o p o r t i o n ( 5 4 . 9 % ) o f a v a i l a b l e h a b i t a t f r o m s p r i n g t h r o u g h e a r l y w i n t e r . A r i t h m e t i c a l l y n e g a t i v e a s s o c i a t i o n s a r i s e e a s i l y , w h i l e p o s i t i v e a s s o c i a t i o n s o c c u r r a r e l y f o r h a b i t a t s m a k i n g up a l a r g e p r o p o r t i o n o f a v a i l a b l e a r e a . F o r e x a m p l e , t h e r e w e r e n e g a t i v e a s s o c i a t i o n s r e c o r d e d b e t w e e n u n l o g g e d f o r e s t a n d m o o s e i n t h e s p r i n g s o f 1 9 8 0 , 1 9 8 2 , a n d 1 9 8 3 w h e n 3 7 % , 4 3 % , a n d 4 0 % , r e s p e c t i v e l y , o f m o o s e l o c a t i o n s w e r e i n u n l o g g e d f o r e s t ( T a b l e 3 ) . S i m i l a r l y , t h e f o l l o w i n g s a m p l i n g p e r i o d s p r o d u c e d n e u t r a l a s s o c i a t i o n s b e t w e e n m o o s e a n d u n l o g g e d f o r e s t i n s p i t e o f h i g h p r o p o r t i o n s o f m o o s e l o c a t i o n s , i n d i c a t e d i n p a r e n t h e s e s , i n u n l o g g e d f o r e s t : s p r i n g 1 9 8 1 ( 4 8 % ) , e a r l y w i n t e r 1 9 8 1 - 1 9 8 2 ( 4 5 % ) , s u m m e r 1 9 8 2 ( 5 5 % ) , a n d f a l l 1 9 8 2 ( 5 8 % ) . T h e r e a r e r e s o u r c e s o r c o n d i t i o n s w i t h i n u n l o g g e d f o r e s t s t h a t a r e a t t r a c t i v e t o m o o s e . I n s u m m e r a n d w i n t e r a v o i d a n c e o f h e a t s t r e s s i s p r o b a b l y t h e f a c t o r t h a t c a u s e d m o o s e t o o c c u p y u n l o g g e d f o r e s t . I n f a l l m o o s e w e r e p o s s i b l y a v o i d i n g h u m a n d i s t u r b a n c e w h i c h r e s u l t s i n g r e a t e r o c c u p a t i o n o f u n l o g g e d f o r e s t b e c a u s e a c c e s s i s t h r o u g h r o a d s t h a t a r e o n l y p r e s e n t w h e r e l o g g i n g h a s o c c u r r e d . 144 7.2.2 D i a m e t e r l i m i t l o g g i n g G i v e n a s i m i l a r f o r e s t c o v e r to u n l o g g e d f o r e s t ( T a b l e 6) and g r e a t e r f o r a g e a b u n d a n c e t h a n f o r e s t i n a l l s a m p l i n g p e r i o d s ( T a b l e s 5, 10, 11, 14, and 15) one m i g h t have e x p e c t e d t h i s h a b i t a t t y p e t o be r e l a t i v e l y a t t r a c t i v e t o moose. The a s s o c i a t i o n of moose w i t h d i a m e t e r l i m i t l o g g i n g was, however, i n c o n s i s t e n t . T a b l e 3, l i s t i n g t h e p r o p o r t i o n o f moose l o c a t i o n s i n t h e f o r e s t / 1 o g g e d h a b i t a t s b e s t i l l u s t r a t e s t h i s a s s e r t i o n . I n t h e s p r i n g o f 1981 t h e r e w e r e no m o o s e l o c a t i o n s i n d i a m e t e r l i m i t l o g g i n g , b u t i n t h e s p r i n g o f 1982, 14% of a l l l o c a t i o n s were i n d i a m e t e r l i m i t l o g g i n g . In t h e summers o f 1980 and 1981 a g a i n t h e r e a r e no l o c a t i o n s i n t h i s h a b i t a t , b u t i n 1 9 8 2 , 8% o f l o c a t i o n s o c c u r r e d i n d i a m e t e r l i m i t l o g g i n g . A n n u a l s h i f t s i n t h e number of moose i n d i a m e t e r l i m i t l o g g i n g were so g r e a t t h a t b o t h m i d - and l a t e w i n t e r 1982 r e c o r d e d s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n s between moose and t h i s h a b i t a t , but m i d - and l a t e w i n t e r 1983 b o t h r e c o r d e d s i g n i f i c a n t n e g a t i v e a s s o c i a t i o n s ( F i g u r e s 5 and 6 ) . D i a m e t e r l i m i t l o g g i n g was c o n f i n e d t o t h e s o u t h w e s t p o r t i o n o f t h e s t u d y a r e a a t l o w e l e v a t i o n s . I t i s p o s s i b l e t h a t moose o c c u p a t i o n of d i a m e t e r l i m i t l o g g i n g was c o n f o u n d e d by t h e s e f a c t o r s . Movement away from t h e s o u t h w e s t c o r n e r and 1 4 5 t o h i g h e r e l e v a t i o n s w o u l d m a k e d i a m e t e r l i m i t l o g g i n g l e s s a v a i l a b l e d u r i n g s p r i n g t h r o u g h e a r l y w i n t e r a n d d u r i n g m i d a n d l a t e w i n t e r s w h e n s n o w w a s s h a l l o w . D i a m e t e r l i m i t l o g g i n g r e c o r d e d r e l a t i v e l y f e w ( t h r e e ) s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n s w i t h m o o s e c o m p a r e d t o e i g h t n e g a t i v e a s s o c i a t i o n s a n d f i v e n e u t r a l a s s o c i a t i o n s . N o n e t h e l e s s , t w o o f t h e s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n s o c c u r r e d d u r i n g t h e s e v e r e m i d a n d l a t e w i n t e r o f 1 9 8 2 w h e n s n o w d e p t h s e x c e e d e d t h e 9 0 c m c r i t i c a l d e p t h b y a l a r g e m a r g i n . R e l o g g i n g o f d i a m e t e r l i m i t l o g g i n g o n w i n t e r r a n g e s h o u l d be s e n s i t i v e t o t h e p o s s i b i l i t y t h a t t h i s h a b i t a t i s a n i m p o r t a n t s h e l t e r f o r m o o s e d u r i n g i n c l e m e n t w i n t e r c o n d i t i o n s . 7 . 2 . 3 I n t e r m e d i a t e u t i l i z a t i o n l o g g i n g T h e i n t e r m e d i a t e u t i l i z a t i o n l o g g e d a r e a s o n t h e s t u d y s i t e r a n k e d h i g h i n t h e i r s e l e c t i o n b y m o o s e t h r o u g h a l m o s t a l l s a m p l i n g p e r i o d s . I n t e r m e d i a t e u t i l i z a t i o n l o g g i n g r e c o r d e d s e v e n s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n s w i t h m o o s e l o c a t i o n s a s c o m p a r e d w i t h t h r e e s i g n i f i c a n t n e g a t i v e a s s o c i a t i o n s , d u r i n g t h e 16 s a m p l i n g p e r i o d s s t u d i e d ( F i g u r e s 4 , 5 a n d 6 ) . T h e s e a r e a s w e r e e s p e c i a l l y a t t r a c t i v e t o m o o s e f r o m e a r l y w i n t e r t o s p r i n g w h e n s i x o f t h e s e v e n p o s i t i v e a s s o c i a t i o n s w e r e r e c o r d e d . T w o o f t h e t h r e e n e g a t i v e 146 a s s o c i a t i o n s o c c u r r e d d u r i n g t h e f a l l , t h e o n l y s e a s o n t o s c o r e c o n s i s t e n t l y low a s s o c i a t i o n i n d i c e s between moose and i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g . The p r o p o r t i o n o f moose l o c a t i o n s i n i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g v a r i e d most w i d e l y i n summer ( T a b l e s 3 and 4 ) , a s e a s o n w h i c h p r o d u c e d a p o s i t i v e , n e g a t i v e , and a n e u t r a l a s s o c i a t i o n b e t w e e n moose and i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g . The l a r g e number o f moose w h i c h occupy t h e w i n t e r r a n g e on t h e n o r t h s i d e o f t h e F r a s e r R i v e r may be t h e r e i n r e s p o n s e t o t h e h e t e r o g e n e o u s c o m b i n a t i o n o f f o r a g e and c o v e r f o u n d i n i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g . I n t e r m e d i a t e u t i l i z a t i o n l o g g i n g r a n k e d s e c o n d t o o l d c l e a r c u t s i n t h e amount of f o r a g e a v a i l a b l e i n a l l w i n t e r p e r i o d s ( T a b l e s 11, 14, and 15) and l i k e l y a v e r a g e s s u f f i c i e n t f o r e s t c o v e r (0.355 o f t h e s k y h e m i s p h e r e , T a b l e 6) t o p r o t e c t moose from t h e r m a l s t r e s s , as i n f e r r e d f r o m a s s o c i a t i o n i n d i c e s . T h e i m p o r t a n c e o f i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g t o w i n t e r i n g moose on t h e s t u d y a r e a can not be o v e r s t a t e d . U n f o r t u n a t e l y , i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g i s r e s p o n s i b l e f o r s u p p r e s s i n g r e g r o w t h of c o m m e r i c a l l y d e s i r a b l e t r e e s . As s u c h , t h e s e a r e a s a r e t h e t a r g e t o f a B.C. F o r e s t S e r v i c e " s i l v i c u l t u r a l s l u m c l e a r i n g p r o g r a m " ( S t e v e n s o n 1 9 8 4 ) . I n t h e i n t e r e s t s o f w i n t e r i n g a l a r g e number o f moose i n t h e s o u t h w e s t c o r n e r o f T F L 30, i n t e r m e d i a t e u t i l i z a t i o n l o g g e d a r e a s i n t h e D a r b y C r e e k , 147 B a r n e y C r e e k , L i m e s t o n e C r e e k , a n d O l s s o n C r e e k a r e a s s h o u l d n o t be c l e a r e d . On t h e p o s i t i v e s i d e , a f o r e s t m a n a g e r h a s e x p r e s s e d c o n c e r n a b o u t r e d u c t i o n s i n t h e a m o u n t o f b r o w s e o n i n t e r m e d i a t e u t i l i z a t i o n l o g g e d s i t e s w h i c h w i n t e r l a r g e d e n s i t i e s o f m o o s e ( W e i n a r d 1 9 8 3 ) . I t i s p o s s i b l e t h a t , a s p a l a t a b l e s h r u b s g r o w o u t o f t h e r e a c h o f m o o s e a n d t h o s e w i t h i n r e a c h h a v e t h e i r p r o d u c t i o n r e d u c e d by h e a v y b r o w s i n g , t h e r e w i l l b e a s h o r t a g e o f f o r a g e f o r w i n t e r i n g m o o s e . T h e p r o d u c t i o n o f t h e o p e n a r e a s w i t h i n i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g c a n b e r e j u v e n a t e d w i t h s m a l l s c a l e m e c h a n i c a l d i s t u r b a n c e . 7 . 2 . 4 O l d c l e a r c u t s F o r e s t r e s o u r c e m a n a g e r s i n t h e P r i n c e G e o r g e a r e a g e n e r a l l y a s s u m e t h a t s e v e n t o f i f t e e n y e a r o l d c l e a r c u t s a r e p r e f e r r e d m o o s e h a b i t a t ( W e i n a r d 1 9 8 0 ) . T h i s a s s u m p t i o n i s b u i l t l a r g e l y u p o n o b s e r v a t i o n s o f l a r g e a m o u n t s o f p a l a t a b l e b r o w s e o n c l e a r c u t s o f t h i s a g e . T h e s e a r e a s d o p r o d u c e a b u n d a n t f o r a g e . A m o n g f o r e s t / l o g g e d h a b i t a t s , o l d c l e a r c u t s r a n k e d f i r s t i n a v a i l a b l e f o r a g e d u r i n g t h e t h r e e w i n t e r s e a s o n s ( e a r l y w i n t e r , m i d w i n t e r , a n d l a t e w i n t e r ) a n d w e r e r a n k e d s e c o n d t o i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g f r o m s p r i n g t o f a l l ( T a b l e s 5 , 1 0 , 1 1 , 1 4 , a n d 1 5 ) . H o w e v e r , o l d 1A8 c l e a r c u t s d i d n o t p r o v e a t t r a c t i v e t o moose. S p r i n g 1980 p r o d u c e d a s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n between moose and o l d c l e a r c u t s , but s p r i n g 1983 p r o d u c e d a s i g n i f i c a n t n e g a t i v e a s s o c i a t i o n . E a r l y w i n t e r 1982-1983 p r o d u c e d t h e o n l y o t h e r s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n . Two summers, a l l f a l l s , one m i d w i n t e r , and one l a t e w i n t e r a l l p r o d u c e d s i g n i f i c a n t n e g a t i v e a s s o c i a t i o n s . The r e m a i n i n g s i x s a m p l i n g p e r i o d s a l l p r o d u c e d n e u t r a l a s s o c i a t i o n s between moose and o l d c l e a r c u t s . O l d c l e a r c u t s , w i t h 3.7% o f t h e s p r i n g t h r o u g h e a r l y w i n t e r a v a i l a b l e a r e a and 6.2% o f t h e m i d - and l a t e w i n t e r a r e a , r a n k e d f i f t h of t h e f i v e f o r e s t / l o g g e d h a b i t a t s i n t e r m s o f t h e p r o p o r t i o n o f t h e s t u d y a r e a o c c u p i e d . As a h a b i t a t m a k i n g up a s m a l l p r o p o r t i o n o f t h e a v a i l a b l e a r e a , v e r y few moose l o c a t i o n s were r e q u i r e d t o p r e v e n t n e g a t i v e a s s o c i a t i o n s t o p r o d u c e p o s i t i v e a s s o c i a t i o n s w i t h o l d c l e a r c u t s . F o r e x a m p l e , 0.5 l o c a t i o n s i n o l d c l e a r c u t s p r o d u c e d a n e u t r a l a s s o c i a t i o n between t h a t h a b i t a t t y p e and moose i n t h e summer of 1980 ( T a b l e 3, T a b l e A, and F i g u r e A). G i v e n t h e ease w i t h w h i c h n e g a t i v e a s s o c i a t i o n s c o u l d be a v o i d e d and p o s i t i v e r e c o r d e d , e i g h t n e g a t i v e a s s o c i a t i o n s v e r s u s two p o s i t i v e a s s o c i a t i o n s must be r e g a r d e d as a s t r o n g i n d i c a t i o n t h a t moose wer e a v o i d i n g o l d c l e a r c u t s . I n s p i t e o f a b u n d a n t f o r a g e , o l d c l e a r c u t s w e r e a v o i d e d by moose. The p r o b a b l e r e a s o n f o r moose a v o i d i n g o l d c l e a r c u t s i n summer and w i n t e r i s t h e l a c k o f s h e l t e r f r o m h e a t s t r e s s . The f a l l n e g a t i v e 149 a s s o c i a t i o n s a r e p o s s i b l y r e l a t e d t o m o o s e a v o i d i n g h u m a n d i s t u r b a n c e i n t h a t h a b i t a t . T h e l a c k o f c l e a r c u t s p l a n t e d a n d g r o w n i n t o y o u n g s t a n d s o f c o n i f e r m a k e s a d e f i n i t i v e s t a t e m e n t o n t h e e f f e c t s o f s u c h d e v e l o p m e n t s i m p o s s i b l e . H o w e v e r , P e e k e t a l . ( 1 9 7 6 ) s p e c u l a t e d t h a t l a r g e a r e a s o f u n i f o r m a g e d c o n i f e r w i l l b e d e t r i m e n t a l t o m o o s e . I t i s p o s s i b l e t h a t b y t h e t i m e p l a n t a t i o n s h a v e g r o w n s u f f i c i e n t t h e r m a l s h e l t e r f o r m o o s e t h e t r e e s w i l l h a v e s u p p r e s s e d a l l g r o w t h o f p a l a t a b l e s h r u b s a n d h e r b s . T h e e f f e c t o f p l a n t a t i o n s o n m o o s e w i l l p r o b a b l y b e t h e g r e a t e s t i n f l u e n c e o n m o o s e h a b i t a t o n t h e s t u d y a r e a i n f u t u r e b e c a u s e c u r r e n t p o l i c y i s t o p l a n t a l l c l e a r c u t s n o t n a t u r a l l y r e g e n e r a t e d . 7 . 2 . 5 R e c e n t c l e a r c u t s W i t h r e l a t i v e l y l i t t l e f o r a g e p r o d u c t i o n ( T a b l e s 5 , 1 0 , 1 1 , 1 4 , a n d 1 5 ) a n d a l m o s t n o s h e l t e r ( T a b l e 6 ) r e c e n t c l e a r c u t s s h o u l d n o t b e a t t r a c t i v e t o m o o s e . I n f a c t m o o s e w e r e l i t t l e a t t r a c t e d t o r e c e n t c l e a r c u t s a s s i g n i f i c a n t n e g a t i v e a s s o c i a t i o n s b e t w e e n m o o s e a n d r e c e n t c l e a r c u t s w e r e r e c o r d e d d u r i n g n i n e o f t h e 16 s a m p l i n g p e r i o d s d a t a w e r e g a t h e r e d , n e u t r a l a s s o c i a t i o n s o c c u r r e d o n f i v e o c c a s i o n s , a n d s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n s w e r e l i m i t e d t o t w o s a m p l i n g p e r i o d s ( F i g u r e s 4 , 5 , a n d 6 a n d T a b l e 4 ) . T h e r e w e r e s e a s o n s 1 5 0 m o o s e a v o i d e d r e c e n t c l e a r c u t s . S i g n i f i c a n t n e g a t i v e a s s o c i a t i o n s a l w a y s o c c u r r e d i n s u m m e r , f a l l , a n d m i d w i n t e r . T h e h i g h e s t a s s o c i a t i o n i n d i c e s r e c o r d e d f o r r e c e n t c l e a r c u t s w e r e i n s p r i n g . T h e t w o p o s i t i v e a s s o c i a t i o n s o c c u r r e d d u r i n g t h e s p r i n g s o f 1 9 8 1 a n d 1 9 8 3 . R e c e n t c l e a r c u t s a r e p r o b a b l y i m p o r t a n t t o m o o s e i n p r o d u c i n g t h e f i r s t n e w g r o w t h i n s p r i n g . T h e c o m p l e t e a b s e n c e o f a f o r e s t c a n o p y m e a n s t h a t t h e s n o w i n r e c e n t c l e a r c u t s w i l l b e t h e f i r s t t o d i s a p p e a r a n d t h e s u n c a n w a r m t h e s o i l p r o d u c i n g n e w g r o w t h . T h e c o n t i n u a l p r o d u c t i o n o f s m a l l r e c e n t c l e a r c u t s w o u l d p r o b a b l y b e n e f i t t h e p r o d u c t i v i t y o f m o o s e i n t h e s t u d y a r e a . 7 . 2 . 6 N o n f o r e s t h a b i t a t s T h e r e w e r e f i v e n o n f o r e s t h a b i t a t s r e c o g n i z e d i n t h i s t h e s i s : w e t l a n d s , u p l a n d s h r u b , r i p a r i a n s h r u b , l a k e s , a n d t h e F r a s e r R i v e r . T h e n o n f o r e s t h a b i t a t m o s t o f t e n p o s i t i v e l y a s s o c i a t e d w i t h m o o s e w a s w e t l a n d s w i t h e i g h t s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n s i n t h e 16 s a m p l i n g p e r i o d s s t u d i e d ( F i g u r e s 1 2 , 1 3 , a n d 1 4 ) . A l l e i g h t p o s i t i v e a s s o c i a t i o n s o c c u r r e d d u r i n g s p r i n g , s u m m e r , a n d f a l l . T h e n e e d t o c o n s e r v e w e t l a n d s a n d a c c e s s t o t h e m i s c l e a r l y d e m o n s t r a t e d . R i p a r i a n s h r u b h a b i t a t p r o d u c e d o n l y t w o s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n s w i t h m o o s e , i n s u m m e r 1 9 8 1 a n d f a l l 1 9 8 2 ( F i g u r e s 1 2 , 1 3 , a n d 1 4 ) . A l l o t h e r s a m p l i n g p e r i o d s p r o d u c e d 151 n e u t r a l a s s o c i a t i o n s . T h e r i p a r i a n s h r u b a r e a o f t e n s u p p o r t s s u c h a d e n s e g r o w t h o f s h r u b s t h a t e n t r y i s d e t e r r e d . O b s e r v a t i o n s o f f o r e s t e d e d g e s , h o w e v e r , f o u n d t h e m h e a v i l y u s e d b y m o o s e . T h e i m p o r t a n c e o f t h e r i p a r i a n s h r u b - f o r e s t e c o t o n e s h o u l d b e r e c o g n i z e d a n d t h e s e a r e a s c o n s e r v e d a s m o o s e h a b i t a t . O b s e r v a t i o n s o f u p l a n d s h r u b ( a l d e r t h i c k e t s ) i n d i c a t e v i r t u a l l y n o m o o s e u s e . T h e f o r e s t o n t h e e d g e o f u p l a n d s h r u b i s u s u a l l y h e a v i l y u s e d w h i c h p r o b a b l y a c c o u n t s f o r t h e f o u r s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n s b e t w e e n m o o s e a n d u p l a n d s h r u b r e c o r d e d d u r i n g t h e s t u d y ( F i g u r e s 1 2 , 1 3 , a n d 1 4 ) . M o o s e m a y b e r e s p o n d i n g t o a h e a v y g r o w t h o f p a l a t a b l e s h r u b s a n d h e r b s w h i c h o f t e n o c c u r s o n t h e u p l a n d s h r u b - f o r e s t e d g e . C l e a r i n g o f a l d e r t h i c k e t s d o e s n o t r e p r e s e n t a t h r e a t t o m o o s e h a b i t a t , b u t c l e a r i n g o f a l d e r t h i c k e t s a n d c l e a r c u t t i n g o f a d j a c e n t f o r e s t w o u l d p r o d u c e a l a r g e a r e a t h a t i s i n h o s p i t a b l e t o m o o s e t h r o u g h m o s t o f t h e y e a r . L a k e s r e c e i v e d a l m o s t n o u s e f r o m r a d i o c o l l a r e d m o o s e d u r i n g t h e s t u d y ( T a b l e 1 6 ) . T h e s m a l l l a k e s i n t h e s t u d y a r e a g e n e r a l l y l a c k t h e s u b m e r g e n t a n d e m e r g e n t a q u a t i c v e g e t a t i o n s o u g h t b y m o o s e . O b s e r v e d n e g a t i v e a n d n e u t r a l a s s o c i a t i o n s b e t w e e n m o o s e a n d l a k e s c a n n o t b e e x t e n d e d t o o t h e r a r e a s . I n t h e B o w r o n L a k e P a r k j u s t s o u t h o f t h e s t u d y a r e a , m o o s e a r e k n o w n t o f o r a g e f o r a q u a t i c p l a n t s i n l a k e s ( R i t c e y a n d V e r b e e k 1 9 6 9 ) . T h e p o t e n t i a l h e a v y u s e o f l a k e s 152 s h o u l d b e r e c o g n i z e d i n t h e c o n s e r v a t i o n o f f o r e s t a s a c c e s s t o l a k e s h o r e s . I n g e n e r a l m o o s e m a d e l i t t l e o r n o u s e o f t h e F r a s e r R i v e r o r e d g e s o f t h e r i v e r ( T a b l e 1 6 , F i g u r e s 1 2 , 1 3 , a n d 1 A ) . H o w e v e r , d u r i n g t h e s e v e r e m i d a n d l a t e w i n t e r o f 1 9 8 2 s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n s b e t w e e n m o o s e a n d t h e r i v e r o c c u r r e d a s m o o s e t o o k t o t h e r i v e r t o a v o i d d e e p s n o w a n d , p o s s i b l y w o l v e s w h i c h w e r e a b u n d a n t i n t h e a r e a d u r i n g 1 9 8 2 . T h e u s e o f t h e r i v e r a s e s c a p e t e r r a i n d u r i n g h a r s h w i n t e r c o n d i t i o n s m e a n s t h a t i t i s i m p o r t a n t t o e n s u r e t h a t m o o s e c a n m o v e q u i c k l y f r o m f o r e s t t o t h e r i v e r . F o r e s t a l o n g t h e r i v e r s h o u l d be c o n s e r v e d a n d , w h e r e i t c a n n o t , m o v e m e n t c o r r i d o r s f r o m f o r e s t t o t h e r i v e r s h o u l d be e s t a b l i s h e d . 7 . 3 F o r e s t C u t t i n g R e c o m m e n d a t i o n s 7 . 3 . 1 L i t e r a t u r e r e v i e w a n d p r a c t i c e s o n a n d n e a r t h e s t u d y  a r e a S e v e r a l a u t h o r s i n t e r e s t e d i n t h e c o n s e r v a t i o n o f m o o s e h a b i t a t h a v e p r e s c r i b e d r e l a t i v e l y l a r g e c l e a r c u t s . P e e k a t a l . ( 1 9 7 6 ) s u g g e s t e d 8 0 h a . c l e a r c u t s . P a r k e r a n d M o r t o n ( 1 9 7 8 ) p r e s c r i b e d c l e a r c u t s o f AO t o 5 0 h a i n N e w f o u n d l a n d . I n O n t a r i o t h e m a x i m u m c l e a r c u t g u i d e l i n e i s 1 3 5 h a ( C h a m b e r l i n 1 9 8 1 , E u l e r 1 9 8 1 ) . 1 5 3 I n B r i t i s h C o l u m b i a t h e r e a r e no p r o v i n c e w i d e g u i d e l i n e s t o l i m i t c l e a r c u t s i z e i n m o o s e h a b i t a t , b u t i n t h e P r i n c e G e o r g e r e g i o n a n a t t e m p t i s m a d e t o r e s t r i c t c l e a r c u t s t o a m a x i m u m 8 0 h a o n w i n t e r r a n g e . On a n e s p e c i a l l y i m p o r t a n t w i n t e r r a n g e , c l e a r c u t s w i l l be l i m i t e d t o 4 0 h a , i f p o s s i b l e ( S t e v e n s o n 1 9 8 4 ) . On s u m m e r r a n g e , t h e c l e a r c u t s i z e a p p e a r s t o b e l i m i t e d o n l y t o t h e e x t e n t t h a t l o g g i n g r a t e s m u s t b e c o m m e n s u r a t e w i t h f o r e s t r o t a t i o n a g e . L a r g e c l e a r c u t s o n m o o s e s u m m e r r a n g e h a v e b e e n j u s t i f i e d o n t h e g r o u n d s t h a t l o g g i n g p r o d u c e s g o o d m o o s e s u m m e r r a n g e , b y s t i m u l a t i n g b r o w s e p r o d u c t i o n , w i t h no h a r m f u l e f f e c t s . R e s t r i c t i o n s o n t h e s i z e o f c l e a r c u t s w e r e l a r g e l y a b a n d o n e d i n t h e s t u d y a r e a , d u r i n g a s p r u c e b a r k b e e t l e i n f e s t a t i o n w h i c h b e g a n i n 1 9 8 0 . C l e a r c u t s o n t h e s t u d y a r e a w e r e c o m m o n l y g r e a t e r t h a n 3 0 0 h a a n d o n e c u t i n t h e s o u t h e a s t e r n c o r n e r o f t h e s t u d y a r e a e x c e e d e d 2 , 2 0 0 h a . I n t h e i m m e d i a t e s t u d y a r e a , t h e b e e t l e i n f e s t a t i o n w i t h a s s o c i a t e d l a r g e c l e a r c u t s h a s e n d e d . S o u t h o f t h e s t u d y a r e a h o w e v e r , b e e t l e s a l v a g e l o g g i n g m a y p r o d u c e a c l e a r c u t o f a b o u t 3 1 , 0 0 0 h a ( S t e v e n s o n 1 9 8 4 ) . F i s h a n d W i l d l i f e s t a f f h o p e t h a t t h i s a r e a w i l l b e p r o d u c t i v e m o o s e h a b i t a t i n 15 y e a r s ( S t e v e n s o n 1 9 8 4 ) . T h e a l l o w a n c e f o r l a r g e c l e a r c u t s o n m o o s e r a n g e r e p r e s e n t s a c o n c e s s i o n t o l o g g i n g e c o n o m i c s . T h e e v i d e n c e 1 5 4 f r o m m a n y s o u r c e s i s t h a t m o o s e d o w e l l i n a r e a s w h i c h a r e h e t e r o g e n e o u s w i t h r e s p e c t t o t h e a v a i l a b i l i t y o f f o r a g e a n d c o v e r . I n A l a s k a , m o o s e s u c c e s s h a s b e e n a t t r i b u t e d t o e x t r e m e l y h e t e r o g e n e o u s h a b i t a t f o l l o w i n g b u r n s ( L e R e s c h e e t a l . 1 9 7 4 ) . I n S c a n d i n a v i a , " c l e a r c u t s " o f a p p r o x i m a t e l y 2 h a may be p a r t i a l l y r e s p o n s i b l e f o r r e c e n t v e r y h i g h d e n s i t i e s o f m o o s e i n t h a t r e g i o n , b u t w i t h m i n i m u m c l e a r c u t s i z e s now g r e a t e r t h a n 5 h a t h e q u a l i t y o f m o o s e h a b i t a t may b e r e d u c e d ( M a r k g r e n 1 9 7 4 ) . H a m i l t o n a n d D r y s d a l e ( 1 9 7 5 ) n o t e d t h a t c u t o v e r s l e s s t h a n 2 0 0 m w i d e a r e n a r r o w e n o u g h t h a t d i s t a n c e f r o m c o v e r d o e s n o t a f f e c t m o o s e , b u t t h a t c u t o v e r s m o r e t h a n 3 0 0 m w i d e c a u s e a r a p i d d e c l i n e i n m o o s e u s e a f t e r 4 0 m a n d n o u s e m o r e t h a n 1 0 0 m f r o m t h e e d g e . I n n o r t h e a s t e r n M i n n e s o t a a s t y l e o f l o g g i n g s i m i l a r t o i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g i n t h e p r e s e n t s t u d y i s b e l i e v e d t o b e r e s p o n s i b l e f o r a l a r g e d e n s i t y o f m o o s e ( P e e k e t a l . 1 9 7 6 ) . I n S a s k a t c h e w a n , H u n t ( 1 9 7 6 ) n o t e d t h a t t h e a v a i l a b i l i t y o f a b u n d a n t r e s i d u a l c o v e r w i t h i n a c u t o v e r r e s u l t s i n m o o s e b e i n g a b l e t o u s e a c u t o v e r w i t h o u t r e s t r i c t i o n . E a s t m a n ( 1 9 7 8 ) n o t e d t h a t p a r t i a l c u t o v e r s w e r e t h e p r e f e r r e d h a b i t a t i n t h e P r i n c e G e o r g e r e g i o n . E a s t m a n ( 1 9 7 8 ) a l s o d i s c u s s e d t h e r e l e v a n c e o f h e t e r o g e n e o u s h a b i t a t t o m o o s e , c o n c l u d i n g t h a t h e t e r o g e n e o u s h a b i t a t i s g e n e r a l l y p r e f e r r e d b y m o o s e . I n O n t a r i o , m o o s e p r e f e r e n c e f o r s e l e c t i v e l y l o g g e d a r e a s w i t h 1% t o 4 0 % o f t h e t r e e s r e m o v e d w a s n o t e d b y W e l s h e t a l . 1 5 5 ( 1 9 8 0 ) . E v e n i n a r e a s w h e r e l a r g e c l e a r c u t s h a v e b e e n r e c o m m e n d e d , t h e r e a r e r e s t r i c t i o n s i m p o s e d o r s u g g e s t e d . P e e k e t a l . ( 1 9 7 6 ) s u g g e s t e d t h a t a s p e n s t a n d s b e c o n s e r v e d . P a r k e r a n d M o r t o n ( 1 9 7 8 ) s u g g e s t e d t h a t 4 0 t o 50 h a c l e a r c u t s b e c o n d u c t e d i n m a n n e r w h i c h w i l l r e s u l t i n a h e t e r o g e n e o u s m i x o f c l e a r c u t s a n d m a t u r e f o r e s t . I n O n t a r i o , t h e r e i s a m a x i m u m c l e a r c u t w i d t h o f 4 0 0 m, a m a x i m u m d i s t a n c e b e t w e e n s h e l t e r b l o c k s o f 2 0 0 m, p r o v i s i o n f o r m o v e m e n t c o r r i d o r s f o r w i l d l i f e , a n d t h e m a x i m u m c l e a r c u t s i z e i s o n l y a l l o w e d i f l o c a l w i l d l i f e o f f i c i a l s b e l i e v e t h a t s u f f i c i e n t e d g e i s b e i n g c r e a t e d by t h e l a r g e c u t ( C h a m b e r l i n 1 9 8 1 , E u l e r 1 9 8 1 ) . T h e v a l u e o f h e t e r o g e n e o u s h a b i t a t , w h i c h p l a c e s s h e l t e r f r o m d e e p s n o w i n t h e v i c i n i t y o f a b u n d a n t f o r a g e , h a s b e e n r e c o g n i z e d a s a n i m p o r t a n t f e a t u r e o f m o o s e w i n t e r r a n g e ( P e e k e t a l . 1 9 7 6 , E a s t m a n 1 9 7 8 , W e l s h e t a l . 1 9 8 0 ) . I t s h o u l d b e r e c o g n i z e d t h a t s h e l t e r f r o m h e a t s t r e s s i s a n i m p o r t a n t b e n e f i t o f f o r e s t c o v e r f o r m o o s e o n w i n t e r r a n g e ( R e n e c k e r a n d H u d s o n 1 9 8 4 , t h i s t h e s i s ) . M o o s e c a n be h e a t s t r e s s e d a t c o m p a r a t i v e l y l o w a i r t e m p e r a t u r e s , e v e n d u r i n g s u m m e r ( K n o r r e 1 9 5 9 , R e n e c k e r a n d H u d s o n 1 9 8 4 ) , m a k i n g t h e p r o v i s i o n o f d e e p s h a d e i n t h e v i c i n i t y o f a b u n d a n t f o r a g e a n i m p o r t a n t c o n s i d e r a t i o n o n m o o s e s u m m e r r a n g e . W i t h r e s p e c t t o t h e p r o d u c t i o n o f e x t r e m e l y l a r g e c l e a r c u t s o n m o o s e s u m m e r r a n g e i n t h e P r i n c e G e o r g e r e g i o n , t h e h o p e t h a t t h e s e a r e a s w i l l be 156 p r o d u c t i v e m o o s e h a b i t a t i n 15 y e a r s , w i t h o u t m a t u r e t r e e s t o p r o v i d e t h e r m a l s h e l t e r , i s p r o b a b l y u n r e a s o n a b l e . H i g h m o o s e d e n s i t i e s u p t o 4 0 0 m f r o m f o r e s t c o v e r ( P a r k e r a n d M o r t o n 1 9 7 8 , E a s t m a n 1 9 7 8 ) a r e p r o b a b l y a n e a r l y w i n t e r p h e n o m e n o n . T h i s s t u d y s u p p o r t s t h e w o r k o f P e e k e t a l . ( 1 9 7 6 ) , W e l s h e t a l . ( 1 9 8 0 ) , a n d T h o m p s o n e t a l . ( 1 9 8 1 ) , w h o s h o w e d t h a t m o o s e m a k e m o r e u s e o f o p e n a r e a s i n e a r l y w i n t e r t h a n a t o t h e r t i m e s o f t h e y e a r . B a s i n g l o g g i n g p r e s c r i p t i o n s o n m o o s e b e h a v i o u r d u r i n g e a r l y w i n t e r c o u l d p r e j u d i c e m a n a g e m e n t a g a i n s t t h e h a b i t a t r e q u i r e m e n t s o f m o o s e i n o t h e r s e a s o n s , p a r t i c u l a r l y s u m m e r , m i d w i n t e r , a n d l a t e w i n t e r . 7 . 3 . 2 C u t t i n g r e c o m m e n d a t i o n s f o r f o r e s t A l t e r n a t i n g 4 h a l o g g e d p a t c h e s w i t h 4 h a p a t c h e s o f s t a n d i n g t i m b e r , s i m i l a r t o t h e i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g i n t h e L i m e s t o n e C r e e k a r e a , w o u l d b e c l o s e t o i d e a l m o o s e h a b i t a t f o r b o t h s u m m e r a n d w i n t e r r a n g e . M a r k g r e n ( 1 9 7 4 ) b e l i e v e d t h a t a l o g g i n g p a t t e r n s i m i l a r t o t h i s w a s r e s p o n s i b l e f o r h i g h m o o s e d e n s i t i e s i n S c a n d i n a v i a . F u r t h e r , i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g i n t h e L i m e s t o n e C r e e k a r e a w a s o b s e r v e d t o s u p p o r t a l a r g e n u m b e r o f m o o s e , a n d i n t e r m e d i a t e u t i l i z a t i o n l o g g i n g i n g e n e r a l w a s p r e f e r r e d by 157 m o o s e d u r i n g t h i s s t u d y ( F i g u r e s 4 , 5 a n d 6 a n d T a b l e 4 ) . L a r g e r l o g g e d p a t c h e s a n d l e s s d i s p e r s i o n o f t h e l o g g i n g i n r e m a i n i n g t i m b e r i s c o m p r o m i s i n g m o o s e p r o d u c t i o n t o l o g g i n g e c o n o m i c s . M o o s e w e r e n o t o b s e r v e d t o u s e c l e a r c u t s o f 4 0 h a a n d l a r g e r , e x c e p t i n e a r l y w i n t e r a n d s p r i n g . C l e a r c u t s o f 4 0 h a a n d l a r g e r a r e t o o l a r g e f o r m o o s e u n l e s s s t r i c t l i m i t s a r e p l a c e d o n t h e l o g g i n g . T h e d i s t a n c e f r o m a n y p o i n t w i t h i n t h e c l e a r c u t t o c o v e r s h o u l d b e l i m i t e d — 2 0 0 m m a y b e a p p r o p r i a t e - - a n d t h a t c o v e r be s t r i c t l y d e f i n e d i n a m a n n e r s i m i l a r t o t h a t i n O n t a r i o : a m i n i m u m o f 3 h a w i t h a m i n i m u m 11 m / h a b a s a l a r e a o f t r e e s a t l e a s t 6 m t a l l ( E u l e r 1 9 8 1 ) . F o r e s t m a n a g e r s w i l l p r o b a b l y f i n d t h e s e r e c o m m e n d a t i o n s f a r t o o r e s t r i c t i v e f o r t h e e c o n o m i c m a n a g e m e n t o f t i m b e r r e s o u r c e s . H o w e v e r , i f a l a n d a r e a i s t o p r o d u c e b o t h t i m b e r a n d m o o s e , t h e n a c o m p r o m i s e w i l l h a v e t o b e s t r u c k b e t w e e n t h e c o m p e t i n g i n t e r e s t s . A f i r s t s t e p i n t h e p r o d u c t i o n o f a m a n a g e m e n t p l a n i s f o r t i m b e r a n d w i l d l i f e m a n a g e r s t o p r e s e n t r e a l i s t i c g o a l s f o r t h e p r o d u c t i o n o f b o t h r e s o u r c e s . F o r e x a m p l e , t h e s t u d y a r e a c o u l d s u p p o r t a t a r g e t d e n s i t y o f 0 . 5 m o o s e / k m , a n i n c r e a s e o f a b o u t 67% f r o m t h e p r e s e n t d e n s i t y . I f s m a l l c l e a r c u t s w e r e a l l o w e d i n a r e a s i m p o r t a n t t o m o o s e , t h e n b o t h 0 . 5 m o o s e / k m a n d l a r g e c l e a r c u t s c o u l d be s u p p o r t e d by t h e s t u d y a r e a . 1 5 8 7 . 3 . 3 Nj3nf_o_ire_£!jt h a b i t a t s , c u r r e n t p r a c t i c e s , a n d r e c o m m e n d a t i o n s T h e i m p o r t a n c e o f w e t l a n d s t o m o o s e h a s b e e n r e c o g n i z e d a n d l o g g i n g i n t h o s e a r e a s i s s u b j e c t t o s t r i c t g u i d e l i n e s w h e r e p o s s i b l e ( S t e v e n s o n 1 9 8 4 ) . F o r e x a m p l e , a b a n o n t h e b u i l d i n g o f l a n d i n g s o n w e t l a n d s a n d a n 8 0 m l e a v e s t r i p a r o u n d w e t l a n d s a r e i m p l e m e n t e d w h e r e p o s s i b l e . An i m p o r t a n t a s p e c t o f t h e s e g u i d e l i n e s i s t h a t t h e i m p o r t a n c e o f t h e i n t e g r i t y o f t h e f o r e s t - w e t l a n d e d g e i s r e c o g n i z e d . T h e i n t e g r i t y o f t h e e d g e b e t w e e n f o r e s t a n d o t h e r h a b i t a t s s h o u l d a l s o b e r e c o g n i z e d . C o n s i d e r a t i o n s h o u l d b e g i v e n t o t h e d i s t a n c e m o o s e w o u l d h a v e t o c r o s s f r o m l a r g e a r e a s o f c o v e r t o w e t l a n d s . A w e t l a n d s u r r o u n d e d b y a n 8 0 m l e a v e s t r i p w o u l d p r o b a b l y r e c e i v e l e s s u s e i s o l a t e d i n a l a r g e c l e a r c u t t h a n i f t h e r e w e r e a c o n t i n u o u s c o n n e c t i o n b e t w e e n t h e w e t l a n d a n d a l a r g e a r e a o f c o v e r , o r i f t h e o p e n a r e a b e t w e e n t h e l e a v e s t r i p a n d c o v e r w e r e l i m i t e d . A l i m i t o f a m a x i m u m o f 2 0 0 m b e t w e e n w e t l a n d s a n d l a r g e a r e a s o f c o v e r m a y b e a p p r o p r i a t e . R i p a r i a n s h r u b a r e a s a r e u s u a l l y u n a f f e c t e d b y l o g g i n g b e c a u s e l o g g e r s h a v e n o i n t e r e s t i n r i p a r i a n s h r u b a r e a s a n d t h e r e f o r e , w h e n a s k e d t o l e a v e t h e m a l o n e , u s u a l l y c o m p l y ( S t e v e n s o n 1 9 8 4 ) . H o w e v e r , t h e r e a p p e a r s t o be no r e c o g n i t i o n o f t h e i m p o r t a n c e o f t h e f o r e s t - r i p a r i a n s h r u b e d g e a s i s 159 d e m o n s t r a t e d f o r t h e f o r e s t - w e t l a n d e d g e . R i p a r i a n s h r u b w h i c h p r o v i d e s f o r a g e , c o m b i n e d w i t h f o r e s t t o p r o v i d e s h e l t e r , y i e l d v a l u a b l e m o o s e h a b i t a t . W i t h o u t t h e f o r e s t t o p r o v i d e s h e l t e r a n d r o u t e s i n t o t h e d e n s e g r o w t h o f t h e r i p a r i a n s h r u b , t h e v a l u e o f t h e s e a r e a s w i l l be r e d u c e d . An 8 0 m l e a v e s t r i p o n r i p a r i a n s h r u b a r e a s m a y l e a v e t h e f o r e s t -s h r u b e d g e i n t a c t a n d t h e e s t a b l i s h m e n t o f c o r r i d o r s o r m a x i m u m a l l o w a b l e d i s t a n c e s f r o m r i p a r i a n a r e a s t o c o v e r may p r e s e r v e t h e v a l u e o f t h i s h a b i t a t . L o g g i n g r e s t r i c t i o n s a r e a p p l i e d t o l a k e s , i n t h e i n t e r e s t s o f p r o t e c t i n g f i s h h a b i t a t : a m a x i m u m 40% o f l a k e e d g e c a n b e l o g g e d a t o n e t i m e w i t h l o g g i n g f o r b i d d e n o n a d j a c e n t a r e a s u n t i l a 2 t o 4 m r e g r o w t h o f c o n i f e r s h a s b e e n a c h i e v e d . A n 8 0 m l e a v e s t r i p a n d m o v e m e n t c o r r i d o r s c o n s i s t i n g o f t r e e s o f a m i n i m u m h e i g h t , e . g . 6 m, o r m i n i m u m d i s t a n c e t h a t m o o s e w o u l d h a v e t o c r o s s t o r e a c h t h e l a k e w o u l d e n h a n c e . t h e p r o t e c t i o n o f l a k e e d g e s f o r m o o s e . T h e F r a s e r R i v e r may b e i m p o r t a n t o n l y a s e s c a p e t e r r a i n f o r m o o s e d u r i n g w i n t e r s o f d e e p s n o w , b u t t h e a b i l i t y o f m o o s e t o m o v e f r e e l y o n t h e r i v e r i n t h o s e c o n d i t i o n s m a y b e i m p o r t a n t t o t h e s u r v i v a l o f a l a r g e m o o s e p o p u l a t i o n i n t h e s t u d y a r e a . F i s h a n d W i l d l i f e s t a f f a t t e m p t t o p r o v i d e f o r f o r e s t a l o n g r i v e r s t o s e r v e a s e s c a p e c o v e r f o r m o o s e . B o t h s i d e s o f a r i v e r m a y n o t b e l o g g e d a t o n c e a n d t h e m a x i m u m 1 6 0 l e n g t h o f r i v e r b a n k w h i c h may be e x p o s e d by a c o n t i n u o u s c u t may be l i m i t e d t o 3 0 0 m. A d d i t i o n a l a t t e n t i o n s h o u l d be g i v e n t o t h e d i s t a n c e m o o s e w o u l d h a v e t o m o v e t h r o u g h o p e n a r e a s t o r e a c h r i v e r i c e . A m i d w i n t e r r u n o f 2 0 0 m m a y b e t o o f a r f o r a m o o s e t r y i n g t o e s c a p e p r e d a t o r s . 161 V I SUMMARY AND CONCLUSIONS T h i s t h e s i s i s p r i m a r i l y d i r e c t e d t o d o c u m e n t i n g t h e e f f e c t s o f p a s t l o g g i n g p r a c t i c e s o n m o o s e h a b i t a t s e l e c t i o n . T h e h a b i t a t o c c u p a t i o n p a t t e r n s o f r a d i o c o l l a r e d m o o s e w e r e r e c o r d e d d u r i n g t h e p e r i o d A p r i l 1 9 8 0 t o M a y 1 9 8 3 i n a n a r e a c e n t e r e d 50 km n o r t h e a s t o f P r i n c e G e o r g e B . C . T h e h y p o t h e s e s m o s t c o m m o n l y u s e d t o e x p l a i n m o o s e h a b i t a t s e l e c t i o n p a t t e r n s w e r e d e t e r m i n e d f r o m a l i t e r a t u r e r e v i e w . T h e h a b i t a t a s s o c i a t i o n s o f t h e r a d i o c o l l a r e d m o o s e w e r e t e s t e d a g a i n s t t h e s e h y p o t h e s e s . F o r t h e w i n t e r p e r i o d : ( 1 ) M o o s e a r e r a n d o m l y d i s t r i b u t e d w i t h r e s p e c t t o h a b i t a t t y p e s . ( 2 ) M o o s e a r e n o t r a n d o m l y d i s t r i b u t e d b u t s e l e c t h a b i t a t s w i t h g r e a t e r a m o u n t s o f f o r a g e e x c e p t w h e n s n o w d e p t h e x c e e d s 9 0 c m . I f s n o w d e p t h e x c e e d s 9 0 c m , t h e n m o o s e w i l l s e l e c t h a b i t a t s w i t h g r e a t e r f o r e s t c o v e r . ( 3 ) M o o s e a r e n o t r a n d o m l y d i s t r i b u t e d b u t s e l e c t t h e h a b i t a t w i t h t h e g r e a t e r f o r e s t c o v e r a s p r o t e c t i o n f r o m c o l d s t r e s s w h e n a i r t e m p e r a t u r e s a r e l e s s t h a n - 2 0 ° C . 1 6 2 F o r t h e summer ( n o n w i n t e r ) p e r i o d : ( 1 ) M o o s e a r e r a n d o m l y d i s t r i b u t e d w i t h r e s p e c t t o h a b i t a t t y p e . ( 2 ) M o o s e a r e n o t r a n d o m l y d i s t r i b u t e d b u t s e l e c t h a b i t a t s w i t h g r e a t e r a m o u n t s o f f o r a g e . ( 3 ) M o o s e a r e n o t r a n d o m l y d i s t r i b u t e d b u t s e l e c t h a b i t a t s w i t h g r e a t e r f o r e s t c o v e r w h e n a i r t e m p e r a t u r e s e x c e e d 2 0 ° C . T h e s t u d y a r e a w a s d i v i d e d i n t o 10 h a b i t a t t y p e s , 5 f o r e s t / l o g g e d h a b i t a t s a n d 5 n o n f o r e s t h a b i t a t s . T h e f o r e s t / l o g g e d h a b i t a t s i n c l u d e d : ( 1 ) r e c e n t c l e a r c u t s , c l e a r c u t s l o g g e d l a t e r t h a n t h e w i n t e r o f 1 9 7 3 ; ( 2 ) o l d c l e a r c u t s , c l e a r c u t s l o g g e d t h e w i n t e r o f 1 9 7 3 o r e a r l i e r ; ( 3 ) i n t e r -m e d i a t e u t i l i z a t i o n l o g g i n g ; ( 4 ) d i a m e t e r l i m i t l o g g i n g ; a n d ( 5 ) f o r e s t . T h e n o n f o r e s t u n l o g g e d h a b i t a t t y p e s i n c l u d e : ( 6 ) u p l a n d s h r u b , a l d e r ; ( 7 ) r i p a r i a n s h r u b ; ( 8 ) w e t l a n d s ; ( 9 ) t h e F r a s e r R i v e r ; a n d ( 1 0 ) l a k e s . T o t e s t w h e t h e r m o o s e w e r e d i s t r i b u t i n g t h e m s e l v e s r a n d o m l y i n e a c h s e a s o n t h e f r e q u e n c y d i s t r i b u t i o n o f m o o s e l o c a t i o n s a c r o s s s i x c a t e g o r i e s , f i v e f o r e s t / l o g g e d h a b i t a t s p l u s n o n f o r e s t h a b i t a t s a s o n e c a t e g o r y , w a s t e s t e d a g a i n s t t h e f r e q u e n c y d i s t r i b u t i o n o f t h o s e h a b i t a t t y p e s , u s i n g t h e X t e s t ( S n e d e c o r a n d C o c h r a n 1 9 8 0 ) . T h e h a b i t a t t y p e s 1 6 3 r e s p o n s i b l e f o r n o n r a n d o m n e s s w e r e i l l u s t r a t e d w i t h t h e C o l e ( 1 9 4 9 ) i n d e x o f a s s o c i a t i o n . T o d r a w o u t t h e r e l a t i o n s h i p b e t w e e n C o l e i n d i c e s a n d f o r a g e p r o d u c t i o n o r f o r e s t c o v e r , c o r r e l a t i o n s b e t w e e n h a b i t a t a s s o c i a t i o n s a n d a v a i l a b l e f o r a g e a n d b e t w e e n h a b i t a t a s s o c i a t i o n s a n d f o r e s t c o v e r w e r e c a l c u l a t e d . S e a s o n s w e r e d e f i n e d a s f o l l o w s : ( 1 ) s p r i n g , f r o m t h e o n s e t o f s p r i n g m i g r a t i o n u n t i l 1 5 J u n e ; ( 2 ) s u m m e r , f r o m 16 J u n e t o 15 S e p t e m b e r ; ( 3 ) f a l l , f r o m 16 S e p t e m b e r u n t i l t h e o n s e t o f f a l l m i g r a t i o n ; ( 4 ) e a r l y w i n t e r , f r o m f a l l m i g r a t i o n u n t i l 1 5 J a n u a r y ; ( 5 ) m i d w i n t e r , f r o m 16 J a n u a r y u n t i l 15 M a r c h ; a n d ( 6 ) l a t e w i n t e r , f r o m 16 M a r c h u n t i l t h e o n s e t o f s p r i n g m i g r a t i o n . M o o s e f o l l o w e d a g e n e r a l a n n u a l p a t t e r n t h a t v a r i e d a c c o r d i n g t o t h e c o n d i t i o n s p r e v a l e n t d u r i n g i n d i v i d u a l s e a s o n s . I n s p r i n g , m o o s e w e r e f o u n d i n o p e n h a b i t a t s . T h e s p r i n g t o s u m m e r c h a n g e o f s e a s o n s a w m o o s e m o v e i n t o h a b i t a t c h a r a c t e r i z e d b y h e a v y f o r e s t c o v e r . I n f a l l , m o o s e w e r e a g a i n a s s o c i a t e d w i t h h e a v y f o r e s t c o v e r s , p e r h a p s e v e n m o r e s o t h a n d u r i n g s u m m e r . E a r l y w i n t e r p r o d u c e d a d r a m a t i c s h i f t o f m o o s e i n t o o p e n h a b i t a t s . I n m i d w i n t e r a n d l a t e w i n t e r , m o o s e r e t r e a t e d t o h a b i t a t s w i t h h e a v y f o r e s t c o v e r . M o o s e w e r e u s u a l l y n o n r a n d o m l y d i s t r i b u t e d w i t h r e s p e c t t o h a b i t a t t y p e . H o w e v e r , t h e a l t e r n a t e h y p o t h e s e s g e n e r a l l y f a i l e d t o a c c o u n t f o r t h e d e p a r t u r e s f r o m r a n d o m n e s s . I n 164 s p r i n g , h a b i t a t a s s o c i a t i o n s w e r e i n t h e d i r e c t i o n o f s e l e c t i o n f o r f o r a g e p r o d u c t i o n . H o w e v e r , t h e r e l a t i o n s h i p was weak as one o f f o u r s p r i n g p e r i o d s p r o d u c e d a n e g a t i v e c o r r e l a t i o n between a s s o c i a t i o n s i n d i c i e s and a v a i l a b l e f o r a g e and a n o t h e r s p r i n g had a l o w p o s i t i v e c o r r e l a t i o n b e t w e e n a s s o c i a t i o n i n d i c i e s and a v a i l a b l e f o r a g e . In summer, moose h a b i t a t a s s o c i a t i o n s c o u l d be e x p l a i n e d by moose a v o i d i n g heat s t r e s s . F a l l h a b i t a t a s s o c i a t i o n s w e r e n e v e r e x p l i c a b l e by t h e t e s t e d a l t e r n a t e h y p o t h e s e s . I n e a r l y w i n t e r , h a b i t a t a s s o c i a t i o n s w e r e o n c e a g a i n i n t h e d i r e c t i o n e x p e c t e d by moose s e l e c t i n g f o r abundant f o r a g e . However, i n one of two e a r l y w i n t e r p e r i o d s s t u d i e d , t h e c o r r e l a t i o n between f o r e s t c o v e r and h a b i t a t a s s o c i a t i o n s was g r e a t e r t h a n t h a t between a v a i l a b l e f o r a g e and h a b i t a t a s s o c i a t i o n s . In one m i d w i n t e r s e a s o n , moose h a b i t a t a s s o c i a t i o n s were c o n s i s t e n t w i t h t h e h y p o t h e s i s t h a t moose s e e k h a b i t a t w i t h h e a v y f o r e s t c o v e r when snow d e p t h s e x c e e d 90 cm, b u t a v o i d a n c e o f c o l d s t r e s s would have p r o d u c e d t h e same p a t t e r n of h a b i t a t a s s o c i a t i o n . No t e s t e d a l t e r n a t e h y p o t h e s i s c o u l d e x p l a i n moose h a b i t a t a s s o c i a t i o n s i n t h e o t h e r m i d w i n t e r s e a s o n , n o r i n two l a t e w i n t e r s e a s o n s . E q u i v a l e n t b l a c k body t e m p e r a t u r e s (T ) were c a l c u l a t e d f o r f o r e s t / 1 o g g e d h a b i t a t s and c o m p a r e d w i t h known t h e r m a l l i m i t s f o r moose i n summer and w i n t e r . T g v a l u e s f o r summer 165 c o n f i r m e d h e a t s t r e s s a s a n i m p o r t a n t f a c t o r a n d t h a t o b s e r v e d h a b i t a t a s s o c i a t i o n s w e r e i n a c c o r d a n c e w i t h m o o s e a v o i d i n g h e a t s t r e s s . T g v a l u e s f o r w i n t e r i n d i c a t e d t h a t h e a t s t r e s s r a t h e r t h a n c o l d s t r e s s w a s a f f e c t i n g m o o s e i n w i n t e r . T h e o b s e r v e d w i n t e r h a b i t a t a s s o c i a t i o n s , w h i c h c o u l d n o t b e e x p l a i n e d b y c o l d , s n o w , o r f o r a g e , w e r e e x p l i c a b l e o n t h e b a s i s o f m o o s e a v o i d i n g h e a t s t r e s s . O t h e r t h a n h e a t s t r e s s i n w i n t e r , t h e r e a r e a d d i t i o n a l h y p o t h e s e s w h i c h c a n a l s o e x p l a i n m o o s e h a b i t a t a s s o c i a t i o n s . M o o s e m a y n o t s e l e c t h a b i t a t o n t h e b a s i s o f p r e v a i l i n g c o n d i t i o n s , b u t r a t h e r may b e e x h i b i t i n g p a t t e r n s l e a r n e d f r o m t h e i r d a m s . M o o s e a v o i d a n c e o f l o g g e d a r e a s i n f a l l c o u l d b e c a u s e d b y t h e i r a v o i d a n c e o f h u m a n a c t i v i t y . F i n a l l y , t h e l a r g e p r o p o r t i o n o f m o o s e l o c a t i o n s i n h e a v y f o r e s t c o v e r s i n a l l s e a s o n s c o u l d be a p r e d a t o r a v o i d a n c e s t r a t e g y . I n g e n e r a l , t h e h y p o t h e s i s t h a t m o o s e r e q u i r e f o r e s t c o v e r t o p r o t e c t t h e m f r o m h e a t s t r e s s w a s m u c h m o r e s u c c e s s -f u l a t p r e d i c t i n g m o o s e h a b i t a t a s s o c i a t i o n s t h a n w a s t h e h y p o t h e s i s t h a t m o o s e s h o u l d s e e k h a b i t a t p r o d u c i n g l a r g e a m o u n t s o f f o r a g e c o n d i t i o n a l u p o n s n o w d e p t h b e i n g l e s s t h a n 9 0 c m . I n f a c t , f o r a g e a b u n d a n c e c o n d i t i o n a l u p o n s n o w d e p t h c o u l d n o t u n e q u i v o c a l l y e x p l a i n t h e p a t t e r n o f m o o s e h a b i t a t a s s o c i a t i o n i n a n y s e a s o n . T h i s r e s u l t s h o u l d be i n t e r p r e t e d a s s u p p o r t i n g t h e n e e d f o r m a t u r e t r e e s o n m o o s e r a n g e , t o a v o i d h e a t s t r e s s , a n d n o t t o m i n i m i z e c o n c e r n f o r m o o s e 166 r e q u i r e m e n t s o f p l e n t i f u l f o r a g e . I n a n y s e a s o n , g o o d m o o s e r a n g e w i l l c o n t a i n b o t h f o r a g e a n d m a t u r e t r e e s . A s i n m a n y s t u d i e s , t h i s t h e s i s c o n c l u d e s t h a t m o o s e r e q u i r e h e t e r o g e n e o u s h a b i t a t . I f m o o s e h a b i t a t i s t o b e c o n s e r v e d , t h e r e q u i r e m e n t s o f f o r a g e , t h e r m a l s h e l t e r , a c c e s s t o w a t e r , a n d e s c a p e c o v e r m u s t a l l be c o n s i d e r e d w h e n l o g g i n g p l a n s a r e m a d e . T h e n e e d f o r s o m e o f t h e s e h a b i t a t f e a t u r e s w i l l b e g r e a t e r o r l e s s e r , d e p e n d i n g u p o n t h e s e a s o n h a b i t a t i s o c c u p i e d , b u t t h e y a r e a l l r e q u i r e d t o s o m e d e g r e e i n a l l s e a s o n s . On t h e s t u d y a r e a , t h e h e t e r o g e n e o u s i n t e r m e d i a t e u t i l i z a t i o n l o g g e d a r e a s s h o u l d be c o n s e r v e d a s mOose h a b i t a t e s p e c i a l l y w h e r e i t o c c u r s o n m o o s e w i n t e r r a n g e . C l e a r c u t s s h o u l d b e k e p t s m a l l , 4 h a . W h e r e l o g g i n g e c o n o m i c s d e m a n d l a r g e c l e a r c u t s , t h e m a x i m u m d i s t a n c e f r o m w i t h i n a c l e a r c u t t o c o v e r s h o u l d be l i m i t e d . 167 V I I L I T E R A T U R E C I T E D A d d i s o n , R. B . , J . C . W i l l i a m s o n , B . P . S a u n d e r s , a n d D. F r a s e r . 1 9 8 0 . R a d i o - t r a c k i n g o f m o o s e i n t h e b o r e a l f o r e s t o f n o r t h - w e s t e r n O n t a r i o . C a n . F i e l d - N a t . 9 4 : 2 6 9 -2 7 6 . B a n g s , E. E . , T . N . B a i l e y , a n d M . F . P o r t n e r . 1 9 8 4 . B u l l m o o s e b e h a v i o u r a n d m o v e m e n t s i n r e l a t i o n t o h a r v e s t o n t h e K e n a i N a t i o n a l W i l d l i f e R e f u g e . A l c e s 2 0 : 1 8 7 - 2 0 7 . B a r n e s , D. L . 1 9 7 6 . A r e v i e w o f p l a n t - b a s e d m e t h o d s o f e s t i m a t i n g f o o d c o n s u m p t i o n , p e r c e n t a g e u t i l i z a t i o n , s p e c i e s p r e f e r e n c e s a n d f e e d i n g p a t t e r n s o f g r a z i n g a n d b r o w s i n g a n i m a l s . P r o c . G r a s s l d . S o c . 5 t h A f r . 1 1 : 6 5 - 7 1 . B a s k e r v i l l e , G. L . 1 9 6 5 . E s t i m a t i o n o f d r y w e i g h t o f t r e e c o m p o n e n t s a n d t o t a l s t a n d i n g c r o p i n c o n i f e r s t a n d s . E c o l o g y 4 6 : 8 6 7 - 8 6 9 . B a t c h e l e r , C . L . 1 9 7 1 . E s t i m a t i o n o f d e n s i t y f r o m a s a m p l e o f j o i n t p o i n t a n d n e a r e s t - n e i g h b o u r d i s t a n c e s . E c o l o g y 5 2 : 7 0 3 - 7 0 9 . . 1 9 7 5 . P r o b a b l e l i m i t o f e r r o r o f t h e p o i n t d i s t a n c e e s t i m a t e o f d e n s i t y . P r o c . N . Z . E c o l . S o c . 2 2 : 2 8 - 3 3 . B e a u c h a m p , J . J . , a n d J . S . O l s o n . 1 9 7 3 . C o r r e c t i o n s f o r b i a s i n r e g r e s s i o n e s t i m a t e s a f t e r l o g a r i t h m i c t r a n s f o r m a t i o n . E c o l o g y 5 4 : 1 4 0 3 - 1 4 0 7 B e l o v s k y , G. E . 1 9 7 8 . D i e t o p t i m i z a t i o n i n a g e n e r a l i s t h e r b i v o r e : t h e m o o s e . T h e o r . P o p . B i o l . 1 4 : 1 0 5 - 1 3 4 . . 1 9 8 1 . O p t i m a l a c t i v i t y t i m e s a n d h a b i t a t c h o i c e o f m o o s e . O e c o l o g i a 4 8 : 2 2 - 3 0 . , a n d P . A . J o r d a n . 1 9 7 8 . T h e t i m e - e n e r g y b u d g e t o f a m o o s e . T h e o r . P o p . B i o l . 1 4 : 7 6 - 1 0 4 . B e r g , W. E . , a n d R. L . P h i l l i p s . 1 9 7 2 . W i n t e r s p a c i n g o f m o o s e i n n o r t h w e s t e r n M i n n e s o t a . P r o c . N o r t h A m . M o o s e C o n f . 8 : 1 6 6 - 1 7 6 . _, a n d w e s t e r n w i l l o w e d . 1 9 7 4 . H a b i t a t u s e b y m o o s e i n n o r t h -M i n n e s o t a w i t h r e f e r e n c e t o o t h e r h e a v i l y a r e a s . N a t u r a l i s t e C a n . 1 0 1 : 1 0 1 - 1 1 6 . 1 6 8 B l a i r , R . M . , R . A l c a n i z , a n d A . H a r r e l l . 1 9 8 3 . S h a d e i n t e n s i t y i n f l u e n c e s t h e n u t r i e n t q u a l i t y a n d d i g e s t i b i -l i t y o f s o u t h e r n d e e r b r o w s e l e a v e s . J . R a n g e M a n a g e . 3 6 : 2 5 7 - 2 6 4 . B o b e k , B . , K. P e r z a n o w s k i , J . S i w a n o w i c z , a n d J . Z i e l i n s k i . 1 9 7 9 . D e e r p r e s s u r e o n f o r a g e i n a d e c i d u o u s f o r e s t . O i k o s 3 2 : 3 7 3 - 3 8 0 . B o n a r , R . , R. S m i t h , a n d D. S . E a s t m a n . 1 9 7 5 . T h e S a l m o n R i v e r m o o s e p r o j e c t : h a b i t a t u t i l i z a t i o n by m o o s e o f a n u n l o g g e d s u b - b o r e a l f o r e s t . B . C . F i s h a n d W i l d l . B r a n c h , V i c t o r i a , 7 3 p p . B o y d , M . 1 9 7 9 . A c o n s i d e r a t i o n o f t h e c o r r e c t e d p o i n t d i s t a n c e m e t h o d o f v e g e t a t i o n s a m p l i n g . A l b e r t a R e c . a n d P a r k s . 1 7 p p . B r a s s a r d , J . M . , E . A u d y , M . C r e t e , a n d P . G r e n i e r . 1 9 7 4 . D i s t r i b u t i o n a n d w i n t e r h a b i t a t o f m o o s e i n Q u e b e c . N a t u r a l i s t e C a n . 1 0 1 : 6 7 - 8 0 . B r i t i s h C o l u m b i a M i n i s t r y o f F o r e s t s . 1 9 8 0 . T h e b i o g e o -c l i m a t i c z o n e s o f B r i t i s h C o l u m b i a . u n p u b l i s h e d m a p . B r u s n y k , L . M . , a n d F . F . G i l b e r t . 1 9 8 3 . U s e o f s h o r e l i n e t i m b e r r e s e r v e s by m o o s e . J . W i l d l . M a n a g e . 4 7 : 6 7 3 - 6 8 5 . C a m p b e l l , G. S . 1 9 7 7 . A n i n t r o d u c t i o n t o e n v i r o n m e n t a l b i o p h y s i c s . S p r i n g e r - V e r l a g I n c . , New Y o r k . 1 5 9 p p . C h a m b e r l i n , L . C . 1 9 7 2 . S o m e a s p e c t s o f p r e f e r r e d w i n t e r m o o s e r a n g e . P r o c . N o r t h Am. M o o s e C o n f . 8 : 1 3 8 - 1 6 5 . . 1 9 8 1 . M a n a g i n g f o r f i s h a n d w i l d l i f e v a l u e s w i t h i n t h e f o r e s t m a n a g e m e n t p l a n n i n g p r o c e s s . A l c e s 1 7 : 1 9 3 -2 2 8 . C a n a d a , A t m o s p h e r i c E n v i r o n m e n t S e r v i c e . 1 9 8 0 . M o n t h l y r e c o r d , m e t e r o 1 o g i c a 1 o b s e r v a t i o n s i n W e s t e r n C a n a d a . E n v i r o n m e n t C a n a d a . D o w n s v i e w , O n t a r i o . C a n a d a , A t m o s p h e r i c E n v i r o n m e n t S e r v i c e . 1 9 8 1 . 1 9 8 2 . 1 9 8 3 . M o n t h l y r a d i a t i o n s u m m a r y . E n v i r o n m e n t C a n a d a . T o r o n t o , O n t a r i o . C o a d y , J . W. 1 9 7 4 . I n f l u e n c e o f s n o w N a t u r a l i s t e C a n . 1 0 1 : 4 1 7 - 4 3 6 . o n b e h a v i o u r o f m o o s e . 169 C o c h r a n , W.G. 1 9 6 3 . S a m p l i n g t e c h n i q u e s , 2 n d e d . J o h n W i l e y a n d S o n s , New Y o r k . 4 1 3 p p . C o l e , L. C. 1 9 4 9 . T h e m e a s u r e m e n t o f i n t e r s p e c i f i c a s s o c i a -t i o n . E c o l o g y 3 0 : 4 1 1 - 4 2 4 . C o w a n , I . M . , W. S . H o a r , a n d J . H a t t e r . 1 9 5 0 . T h e e f f e c t o f f o r e s t s u c c e s s i o n u p o n t h e q u a n t i t y a n d u p o n t h e n u t r i t i v e v a l u e s o f w o o d y p l a n t s u s e d a s f o o d b y m o o s e . C a n . J . R e s . S e c t . D 2 8 : 2 4 9 - 2 7 1 . D e a n , S . , J . W. B u r k h a r d t , a n d R. 0 . M e e u w i g . 1 9 8 1 . E s t i m a -t i n g t w i g a n d f o l i a g e b i o m a s s o f s a g e b r u s h , b i t t e r b r u s h , a n d r a b b i t b r u s h i n t h e G r e a t B a s i n . J . R a n g e M a n a g e . 3 4 : 2 2 4 - 2 2 7 . D o d d s , D. G. 1 9 7 4 . D i s t r i b u t i o n , h a b i t a t , a n d s t a t u s o f m o o s e i n t h e A t l a n t i c P r o v i n c e s o f C a n a d a a n d n o r t h e a s t -e r n U n i t e d S t a t e s . N a t u r a l i s t e C a n . 1 0 1 : 5 1 - 6 5 . D o r n , R. D. 1 9 7 0 . M o o s e a n d c a t t l e f o o d h a b i t s i n s o u t h w e s t M o n t a n a . J . W i l d l . M a n a g e . 3 4 : 5 5 9 - 5 6 4 . E a s t m a n , D. S . 1 9 7 8 . H a b i t a t s e l e c t i o n a n d u s e i n w i n t e r b y m o o s e i n s u b - b o r e a l f o r e s t s o f n o r t h c e n t r a l B r i t i s h C o l u m b i a , a n d r e l a t i o n s h i p s t o f o r e s t r y . P h . D . t h e s i s , U n i v e r s i t y o f B r i t i s h C o l u m b i a , V a n c o u v e r . E d w a r d s , R. Y . 1 9 5 6 . S n o w d e p t h s a n d u n g u l a t e a b u n d a n c e i n t h e m o u n t a i n s o f w e s t e r n C a n a d a . J . W i l d l . M a n a g e . 2 0 : 1 5 9 - 1 6 8 . , a n d R. W. R i t c e y . 1 9 5 6 . T h e m i g r a t i o n s o f a m o o s e h e r d . J . M a m m a l . 3 7 : 4 8 6 - 4 9 4 . E u l e r , D. 1 9 8 1 . A m o o s e h a b i t a t s t r a t e g y f o r O n t a r i o . A l c e s 1 7 : 1 8 0 - 1 9 2 . F i n c h , V . A . 1 9 7 2 . T h e r m o r e g u l a t i o n a n d h e a t b a l a n c e o f t h e E a s t A f r i c a n e l a n d a n d h a r t e b e e s t . A m . J . P h y s i o l . 2 2 2 : 1 3 7 4 - 1 3 7 9 . F r a n e , J . 1 9 8 1 . A l l p o s s i b l e s u b s e t s r e g r e s s i o n . p 2 6 4 - 2 7 7 i n W . J . D i x i o n e d . B M D P s t a t i s t i c a l s o f t w a r e . 1 9 8 1 . U n i v e r s i t y o f C a l i f o r n i a P r e s s , B e r k e l e y . 7 2 6 p p . 170 F r a s e r , D . , E . R. C h a v e z , a n d J . E . P a l o h e i m o . 1 9 8 4 . A q u a t i c f e e d i n g b y m o o s e : s e l e c t i o n o f p l a n t a n d f e e d i n g a r e a s i n r e l a t i o n t o p l a n t c h e m i c a l c o m p o s i t i o n a n d c h a r a c t e r -i s t i c s o f l a k e s . C a n . J . Z o o l . 6 2 : 8 0 - 8 7 . G a s a w a y , W. C , S . D. D u B o i s , a n d K. L . B r i n k . 1 9 8 0 . 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B i o m e t r y - t h e p r i n c i p -l e s a n d p r a c t i c e o f s t a t i s t i c s i n b i o l o g i c a l r e s e a r c h 2 n d e d . W . H . F r e e m a n a n d C o . , S a n F r a n c i s c o , C a l i f o r n i a . 8 5 9 p p . 175 S t e v e n s o n , D. G. 1 9 8 4 . H a b i t a t P r o t e c t i o n T e c h n i c i a n , M i n i s -t r y o f E n v i r o n m e n t , P r i n c e G e o r g e , B r i t i s h C o l u m b i a , p e r s . c o m m . S t e y n , D. G. 1 9 8 0 . T h e c a l c u l a t i o n o f v i e w f a c t o r s f r o m f i s h e y e - l e n s p h o t o g r a p h s . A t m o s p h e r e - O c e a n 1 8 : 2 4 8 - 2 5 8 . . 1 9 8 2 . A s s i s t a n t P r o f e s s o r , D e p a r t m e n t o f G e o g r a p h y , U n i v e r s i t y o f B r i t i s h C o l u m b i a . p e r s . c o m m . S t o n e , T . L . , a n d H . S . C r a w f o r d . 1 9 8 1 . E s t i m a t i n g f o l i a g e a n d t w i g w e i g h t o f s p r u c e a n d f i r . J . W i l d l . 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J . W i l d l . M a n a g e . 3 8 : 9 4 4 - 9 4 6 . . 1 9 7 8 a . C e r v i d d i s t r i b u t i o n , b r o w s e a n d s n o w c o v e r i n A l b e r t a . J . W i l d l . M a n a g e . 4 2 : 3 5 2 - 3 8 1 . . 1 9 7 8 b . H a b i t a t r e q u i r e m e n t s o f m o o s e — t h e p r i n c i p a l t a i g a r a n g e a n i m a l . P r o c e e d i n g s I n t e r n a t i o n a l R a n g e l a n d C o n g r e s s . 1 : 4 6 2 - 4 6 5 . . 1 9 8 1 . B r o w s e i n v e n t o r i e s : t e c h n i q u e s a n d e v a l u a t i o n . I n : M i l l e r , F . L . , A . G u n n a n d S . R . H i e b ( e d s ) . C e n s u s a n d i n v e n t o r y m e t h o d s f o r p o p u l a t i o n a n d h a b i t a t . F o r e s t W i l d l i f e a n d R a n g e E x p e r i m e n t a l S t a t i o n , U n i v e r s i t y o f I d a h o , 2 2 0 p p . T e l f e r , E. S . a n d J . P . K e l s a l l . 1 9 7 9 . S t u d i e s o f m o r p h o l o g i -c a l p a r a m e t e r s a f f e c t i n g u n g u l a t e l o c o m o t i o n i n s n o w . C a n . J . Z o o l . 5 7 : 2 1 5 3 - 2 1 5 9 . 176 T h o m p s o n , I . D . , D. A . W e l s h , a n d M . F . V u k e l i c h . 1 9 8 1 . T r a d i t i o n a l u s e o f e a r l y w i n t e r c o n c e n t r a t i o n a r e a s b y m o o s e i n n o r t h e a s t e r n O n t a r i o . A l c e s 1 7 : 1 - 1 4 . Tomm , H . 0 . 1 9 7 8 . R e s p o n s e o f w i l d u n g u l a t e s t o l o g g i n g p r a c t i c e s i n A l b e r t a . M . S c . t h e s i s , U n i v e r s i t y o f A l b e r -t a , E d m o n t o n . T r e m b l a y , J . 1 9 8 1 . Q u a l i t e d e l a n o u r i t u r e d ' e t e d u c e r f d e V i r g i n i e a l ' l l e d ' A n t i c o s t i . M . S c . t h e s e , U n i v e r s i t e de L a v a l , S t . F o y . ( i n e n g l i s h ) V a n B a l l e n b e r g h e , V . , a n d J . M . P e e k . 1 9 7 1 . R a d i o t e l e m e t r y s t u d i e s o f m o o s e i n n o r t h e a s t e r n M i n n e s o t a . J . W i l d l . M a n a g e . 3 5 : 6 3 - 7 1 . W e i n a r d , R. M. 1 9 8 0 . 1 9 8 3 . R e s o u r c e C o o r d i n a t o r , N o r t h w o o d P u l p a n d T i m b e r L t d . p e r s . c o m m . W e l s h , D. A . , K. P . M o r r i s o n , 1 9 8 0 . W i n t e r u t i l i z a t i o n t o f o r e s t h a r v e s t i n g . 1 6 : 3 9 8 - 4 2 8 . K . O s w a l d , a n d E. R. T h o m p s o n , o f h a b i t a t b y m o o s e i n r e l a t i o n P r o c . N o r t h A m . M o o s e C o n f . - 2 -POSITIONS HELD: B . C . M i n i s t r y o f E n v i r o n m e n t W i l d l i f e I n v e n t o r y T e c h n i c i a n 1 9 7 5 - 1 9 7 6 PUBLICATIONS: S c h w a b , F . E . , S . W . S c h w a b , a n d M . D . P i t t . 1 9 8 4 . Moose i m m o b i l i z a t i o n p r o g r a m i n n o r t h c e n t r a l B r i t i s h C o l u m b i a . A l c e s 2 0 : 2 0 9 - 2 2 1 . S c h w a b , F . E . 1 9 8 2 . Use o f m a r k - r e c a p t u r e i n d i c e s t o c e n s u s a moose p o p u l a t i o n : R e p o r t t o t h e P r i n c e G e o r g e o f f i c e o f t h e B . C . F i s h and W i l d l i f e B r a n c h . 5 p p . S c h w a b , F . E . 1 9 8 0 . O m e n i c a c o w / c a l f moose s u r v e y s p r i n g 1 9 8 0 : R e p o r t t o t h e P r i n c e G e o r g e o f f i c e o f t h e B . C . F i s h and W i l d l i f e B r a n c h . 7 p p . S c h w a b , F . E . 1 9 7 9 . E f f e c t o f v e g e t a t i o n s t r u c t u r e on b r e e d i n g b i r d c o m u n i t i e s i n t h e d r y z o n e D o u g l a s f i r f o r e s t o f s o u t h e a s t e r n B r i t i s h C o l u m b i a . M . S c . t h e s i s , U n i v e r s i t y o f B r i t i s h C o l u m b i a , V a n c o u v e r . 135 p p . 

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