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Winter habitat selection and use by moose in the West-Chilcotin region of British Columbia Baker, Bruce Garry 1990

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WINTER HABITAT SELECTION AND USE BY MOOSE IN THE WEST-CHILCOTIN REGION OF BRITISH COLUMBIA By BRUCE GARRY BAKER B . S c , The U n i v e r s i t y o f V i c t o r i a , 1986 THESIS SUBMITTED IN PARTIAL FULFILLMENT THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE i n THE FACULTY OF GRADUATE STUDIES (Department o f 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 t h e r e q u i r e d s t a n d a r d THE UNIVERSITY OF BRITISH COLUMBIA JULY 1990 © BRUCE GARRY BAKER, 1990 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 P/dnrt S<uer\C.z The University of British Columbia Vancouver, Canada D ^ e July 3A /99Q  DE-6 (2/88) Abstract Wetlands have been considered an important component of w i n t e r i n g 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 i n the West-Chilcotin Region of B r i t i s h Columbia. This study evaluates the importance of wetlands, p a r t i c u l a r l y the ecotone between f o r e s t s and wetlands and i d e n t i f i e s important cover types f o r wintering moose. Additional baseline data regarding food habits and home range sizes are included. Two hypotheses were tested i n t h i s study; that moose habitat use was independent of cover type, and that moose habitat use was random with respect to distance from forest/wetland borders. The data i n t h i s study led to r e j e c t i o n of both these hypotheses. Moose used spruce (Picea glauca) wetlands and spruce for e s t s more frequently than expected i f use were random. Moose concentrated p r i m a r i l y within 100 m of the forest/wetland edge and v i r t u a l l y d i d not use areas g r e a t e r than 200 m from the edge. The combination of food and cover i n areas of spruce and edge i s l i k e l y a major factor determining habitat use of wintering moose. Average home range sizes of radio-collared moose ranged from 20.7 to 45.2 km2. Bog b i r c h ( B e t u l a q l a n d u l o s a ) , l o d g e p o l e p i n e (Pinus c o n t o r t a ) , and willows ( S a l i x spp.) were the most frequently consumed forages. - i i -Table of Contents Abstract i i Table of Contents i i i L i s t of Tables v i L i s t of Figures i x Acknowledgments x 1. INTRODUCTION 1 1.1. Current Recommendations 2 1.2. Objectives 2 1.3. L i t e r a t u r e Review 3 1.3.1. History of Moose i n the C h i l c o t i n 3 1.3.2. Winter Habitat Use 4 1.3.3. Use of Edge Habitats 5 2. THEORY AND HYPOTHESES 6 3. STUDY AREA 7 3.1. Location 7 3.2. Climate 7 3.3. Vegetation 10 3.4. S o i l s 11 4. METHODS 12 4.1. Habitat Selection 12 4.2. Cover Type Descriptions 12 4.2.1. Forest Habitats 13 4.2.2. Wetland Habitats 14 4.3. Cover Type A v a i l a b i l i t y Assessment 15 4.4. Physical Structure 16 4.5. Habitat Use 18 4.5.1. Moose Capture 18 4.5.2. Moose Locations 18 4.6. S t a t i s t i c a l Analyses 19 4.7. Moose Home Ranges 21 4.8. Winter Diet 22 4.8.1. Composition 22 4.8.2. Quality 23 4.9. Snow Depth 23 5. RESULTS 24 5.1. Habitat Selection 24 5.1.1. Cover Type Classes 24 5.1.2. Distance to Edge 28 5.1.3. Distance to Wetland 30 5.1.4. Distance to Forest Edge 32 5.1.5. Forest Age 34 5.1.6. Forest Density 35 - i i i -5.1.7. Forest Height 37 5.1.8. Forest DBH 37 5.1.9. Forest Canopy Closure 39 5.1.10. Forest Shrub Cover 40 5.1.11. Wetland Shrub Cover 41 5.1.12. Wetland Shrub Height 42 5.2. Monthly Analyses of Use 44 5.2.1. Cover Type Classes 44 5.2.2. Distance to Wetland 50 5.2.3. Distance to Forest 51 5.2.4. Forest Age 52 5.2.5. Forest Height 53 5.2.6. Forest Density 54 5.2.7. Forest Canopy Closure 54 5.2.8. Forest DBH 55 5.2.9. Forest Shrub Cover 55 5.2.10. Wetland Shrub Cover 57 5.2.11. Wetland Shrub Height 57 5.3. Winter Home Range Size 58 5.4. Winter Diet 59 5.4.1. Composition 59 5.4.2. Monthly Analyses 60 5.4.3. Quality 61 5.5. Snow Depth 62 6. DISCUSSION 64 6.1. Selection Indices 64 6.1.1. Selection 64 6.1.2. Errors i n Interpretation 66 6.1.3. Assumptions 67 6.1.4. Selection and A v a i l a b i l i t y 68 6.1.5. Is Use Equivalent to Selection 71 6.1.6. Is the Logic Flawed? 72 6.1.7. The Unit i s I l l o g i c a l 74 6.1.8. When Do Habitat Selection Indices Work?.... 76 6.1.9. Recommendations 77 6.2. Selection Results 78 6.2.1. Cover Type Classes 78 6.2.2. Distance to Edge 80 6.2.3. Forest Age 81 6.2.4. Forest Density and DBH 81 6.2.5. Forest Height 82 6.2.6. Forest Canopy Closure 83 6.2.7. Forest Shrub Cover 83 6.2.8. Wetland Shrub Cover 84 6.2.9. Wetland Shrub Height 85 6.3. Importance of Wetlands 86 6.4. Monthly Analyses of Use 86 6.4.1. Forest Versus Wetland 87 6.4.2. Forests 88 - i v -6.5. Diet composition 90 6.6. Diet Quality 91 6.7. Summary 91 7. MANAGEMENT RECOMMENDATIONS 94 8. LITERATURE CITED 97 - V -L i s t of Tables Table l . Climate data for Anahim Lake B.C. Monthly Averages 1951-1980 9 Table 2. A v a i l a b i l i t y of forest cover type classes i n the West-Chilcotin 25 Table 3. A v a i l a b i l i t y and moose use of cover classes i n the West-Chilcotin during winter 1988 26 Table 4. Moose use of cover classes i n the West-Chilcotin during winter 1987 28 Table 5. A v a i l a b i l i t y and moose use of distance-to-edge classes i n the West-Chilcotin during winter 1988 29 Table 6. A v a i l a b i l i t y and moose use of distance-to-edge classes i n the West-Chilcotin during winter 1987 29 Table 7. A v a i l a b i l i t y and moose use of distance-to-wetland classes i n the West-Chilcotin during winter 1988 31 Table 8. A v a i l a b i l i t y and moose use of distance-to-wetland classes i n the West-Chilcotin during winter 1987 31 Table 9. A v a i l a b i l i t y and moose use of distance-to-forest classes i n the West-Chilcotin during winter 1988 33 Table 10. A v a i l a b i l i t y and moose use of distance-to-forest classes i n the West-Chilcotin during winter 1987 34 Table 11. A v a i l a b i l i t y and moose use of forest age classes i n the West-Chilcotin during winter 1988 35 Table 12. A v a i l a b i l i t y and moose use of forest density classes i n the West-Chilcotin during winter 1988 36 Table 13. A v a i l a b i l i t y and moose use of forest height classes i n the West-Chilcotin during winter 1988 38 - v i -Table 14. A v a i l a b i l i t y and moose use of forest DBH classes i n the West-Chilcotin during winter 1988 38 Table 15. A v a i l a b i l i t y and moose use of forest canopy closure classes i n the West-Chilcotin during winter 1988 40 Table 16. A v a i l a b i l i t y and moose use of forest shrub cover classes i n the West-Chilcotin during winter 1988 41 Table 17. A v a i l a b i l i t y and moose use of wetland shrub cover classes i n the West-Chilcotin during winter 1988 42 Table 18. A v a i l a b i l i t y and moose use of wetland shrub height classes i n the West-Chilcotin during winter 1988 43 Table 19. Moose use of cover types i n the West-Chilcotin during January 1987 and 1988 45 Table 20. Moose use of cover types i n the West-Chilcotin during February 1987 and 1988 46 Table 21. Moose use of cover types i n the West-Chilcotin during March 1987 and 1988 47 Table 22. Monthly moose use of distance-to-wetland classes i n the West-Chilcotin during winter 1987 and 1988 51 Table 23. Monthly moose use of distance-to-forest classes i n the West-Chilcotin during winter 1987 and 1988 52 Table 24. Monthly moose use of forest age classes i n the West-Chilcotin during winter 1988 53 Table 25. Monthly moose use of forest height classes i n the West-Chilcotin during winter 1988 53 Table 26. Monthly moose use of forest density classes i n the West-Chilcotin during winter 1988 54 - v i i -Table 27. Monthly moose use of forest canopy closure classes i n the West-Chilcotin during winter 1988 55 Table 28. Monthly moose use of forest DBH classes i n the West-Chilcotin during winter 1988 56 Table 29. Monthly moose use of forest shrub cover classes i n the West-Chilcotin during winter 1988 56 Table 30. Monthly moose use of wetland shrub cover classes i n the West-Chilcotin during winter 1988 57 Table 31. Monthly moose use of wetland shrub height classes i n the West-Chilcotin during winter 1988 58 Table 32. Winter home range sizes of radio-collared moose i n the West-Chilcotin during 1987 and 1988 59 Table 33. Winter di e t composition of radio-collared moose i n the West-Chilcotin during 1987 and 1988 60 Table 34. Monthly n u t r i t i o n a l parameters of major forage species consumed by wintering moose i n the West-Chilcotin, 1988 62 Table 35. Winter 1987 and 1988 d a i l y snow depths (cm) recorded at 5 Mile Ranch i n the West-Chilcotin 63 Table 36. Example i l l u s t r a t i n g r e s u l t s of comparing usage and a v a i l a b i l i t y when a common but seldom-used item i s included (A) and when excluded (B) from consideration 69 - v i i i -L i s t of Figures Figure l . Location of the study area within B r i t i s h Columbia 8 - i x -Acknowledgments I would l i k e to express my h e a r t f e l t thanks to a l l the i n d i v i d u a l s and agencies who made completion of t h i s p r o j e c t possible. F i r s t and foremost, I want to thank my thesis advisor Dr. Michael P i t t f o r h i s endless support, encouragement, and generosity. Thanks go to Herb Langin f o r h i s r o l e i n i n i t i a t i n g the pr o j e c t and who, along with Randy Wright, were instrumental i n r a d i o - c o l l a r i n g the moose. Also thanks to Debbie Cichowski who a s s i s t e d with data c o l l e c t i o n and analyses, p i l o t s Wayne Escott and Floyd Vaughan whose s k i l l s made short work of lo c a t i n g moose, Mary R a t z l a f f who provided a home away from home, and Br i a n Nyberg who helped obtain funding and read the manuscript. I would a l s o l i k e to thank my committee members Dr. Fred Bunnell and Dr. Brian H o l l . Agencies that provided funding were the B r i t i s h Columbia M i n i s t r y of Environment, the B r i t i s h Columbia M i n i s t r y of Forests, the B r i t i s h Columbia Habitat Conservation Foundation, the Canadian W i l d l i f e S ervice and the U n i v e r s i t y of B r i t i s h Columbia. -x-1. INTRODUCTION Moose (Alces a l c e s andersonii) are an important big-game resource f o r the C a r i b o o - C h i l c o t i n region of B r i t i s h Columbia (B.C.), providing recreational opportunities for both consumptive and non-consumptive users. In 1981, 11,430 people hunted a t o t a l of 105,000 days i n t h i s r e g i o n , g e n e r a t i n g $3,160,500 i n r e c r e a t i o n a l resource value (Reid 1985). Recent outbreaks of mountain pine b e e t l e (Dendroctinus ponderosae) have reached epidemic proportions throughout the mature lodgepole pine (Pinus c o n t o r t a ) f o r e s t s i n the C h i l c o t i n r e g i o n , r e s u l t i n g i n accelerated harvest of pine, as well as mixed stands of pine and spruce (Picea spp.). Harvesting these f o r e s t s may adversely a f f e c t moose populations on the p l a t e a u , although p o t e n t i a l e x i s t s to use logging to improve moose habitat. Despite the value of moose to t h i s region, l i t t l e research has been conducted to determine t h e i r h a b i t a t requirements. Whereas studies i n other regions of B.C. document p a t t e r n s of h a b i t a t use by moose, they are not applicable to the C h i l c o t i n region where the harsh environment l i m i t s forage p r o d u c t i v i t y , e s p e c i a l l y i n forested areas (Annas and Coupe 1979). Presently, the M i n i s t r y of Environment (MOE) i s providing recommendations f o r f o r e s t h a r v e s t i n g on c r i t i c a l moose wintering ranges with l i t t l e information on which to base t h e i r recommendations. This project was i n i t i a t e d by the MOE to provide an improved assessment of habitat requirements f o r w i n t e r i n g moose i n the C h i l c o t i n region and to provide guidelines f o r integrating - l -habitat management with forest harvesting a c t i v i t i e s . 1.1 Current Recommendations Winter a e r i a l surveys of moose i n t h i s area have led w i l d l i f e b i o l o g i s t s t o b e l i e v e t h a t moose c o n c e n t r a t e i n or near wetlands, u t i l i z i n g the border between wetlands and surrounding c o n i f e r stands. Logging g u i d e l i n e s have been based on t h i s b e l i e f and logging within 100-200 m of a wetland edge has been r e s t r i c t e d . 1.2 Objectives The primary aim of t h i s research i s to document the patterns of w i n t e r h a b i t a t use by moose i n the C h i l c o t i n . S p e c i f i c o b j e c t i v e s are 1) to determine i f moose c o n c e n t r a t e around w e t l a n d / f o r e s t borders or i f t h i s o b s e r v a t i o n r e s u l t s from d i f f e r e n t i a l s i g h t a b i l i t y of moose i n the open compared to moose i n f o r e s t s ; 2) i f moose do concentrate around wetland/forest borders, to determine the distance from the border where use becomes minimal; and 3) to determine s p e c i f i c types of fo r e s t and wetland cover that moose use. Meeting these o b j e c t i v e s w i l l a s s i s t managers i n designing logging p r a c t i c e s that w i l l have minimal detrimental e f f e c t s on wintering moose, and which may improve habitat q u a l i t y for moose. A f u r t h e r o b j e c t i v e of t h i s p roject i s to gather b a s e l i n e d a t a r e g a r d i n g t h e g e n e r a l e c o l o g y o f moose i n t h e West-Chilcotin. S p e c i f i c a l l y , data regarding winter home range sizes and winter d i e t composition and q u a l i t y are sought. 2-1.3 L i t e r a t u r e Review 1.3.1 History of Moose i n the C h i l c o t i n Moose have inhabited the C h i l c o t i n only since the turn of the century. Hatter (1950) reported that before 1900 moose were absent from the i n t e r i o r of the province. For the most part moose were present only i n the north-eastern s e c t i o n of the province adjacent to the Rocky Mountains; however, since 1900 moose have dispersed west and south of t h i s area and are now abundant throughout the central i n t e r i o r . The most important f a c t o r i n i t i a t i n g the s h i f t of moose populations into previously unoccupied areas was probably forest f i r e s caused by human settlement and i n d u s t r y (Hatter 1950). These f i r e s removed climax f o r e s t s which acted as b a r r i e r s to d i s p e r s a l . Once moose reached the i n t e r i o r p l a t e a u t h e i r d i s p e r s a l was favored due to the topography of the plateau which offered few b a r r i e r s to dispersal south and west (Hatter 1950). Once moose reached the central i n t e r i o r of B r i t i s h Columbia t h e i r numbers increased r a p i d l y , l i k e l y due to the f a v o r a b l e habitat provided by the numerous wetlands of the region. Hatter (1950) i d e n t i f i e d that peak populations l i k e l y occurred i n the 1920's and 1930's. Since then, moose populations have declined, but how present day populations compare to those peak populations i s not known. -3-1.3.2 Winter Habitat Use Studies of moose i n North America have indicated that moose s e l e c t areas of high forage production i n winter (Telfer 1978); however, factors such as snow depth, cold stress and heat stress may cause moose to seek more sheltered environments. Deep snow may cause moose to seek s h e l t e r e d environments e i t h e r f o r eas i e r locomotion or increased browse a v a i l a b i l i t y . Moose movements become r e s t r i c t e d when snow exceeds 65 cm (Des Meules 1964, T e l f e r 1970, K e l s a l l and Prescott 1971, P h i l l i p s et a l . 1973, Coady 1974). Increased metabolic demands of impeded locomotion associated with open feeding areas may make feeding there e n e r g e t i c a l l y uneconomical. Forested areas with reduced snow depths may make foraging i n forests more b e n e f i c i a l . Protection from cold stress has been c i t e d by several authors as a reason f o r moose using f o r e s t e d areas during the winter ( P h i l l i p s et a l . 1973, T e l f e r 1978, Brusnyk and G i l b e r t 1983). This explanation i s often used when moose use fo r e s t stands even though snow depths i n the open are not r e s t r i c t i n g . H a t t e r (1950), Knowlton (I960), Van Ballenberghe and Peek (1971), and Eastman (1978) speculated that f o r e s t s e l e c t i o n i s r e l a t e d to thermal protection and/or ease of locomotion. Schwab (1985) speculated that heat s t r e s s may a l s o cause moose to seek shady f o r e s t e d areas i n winter. His idea i s supported by Renecker and Hudson (1986) who found that moose have no metabolic response to temperatures as low as -30 C, but have increased metabolic rates at temperatures of -5 C, i n d i c a t i n g that moose are e a s i l y heat-stressed, even at low temperatures. -4-1.3.3 Use of Edge Habitats Since the advent of cl e a r - c u t logging, w i l d l i f e b i o l o g i s t s have been interested i n the ef f e c t s that the r e s u l t i n g increase i n edge have on b i g game (Lyon and Jensen 1980). Numerous studies have shown that wild ungulates prefer edge habitats over other a v a i l a b l e h a b i t a t s (Lyon and Jenson 1980, McNicol and G i l b e r t 1980, and Brusnyk and Gi l b e r t 1983). Edge habitats have the advantage of providing abundant browse i n close proximity to cover. While these s t u d i e s suggest t h a t edge h a b i t a t s are important, they do not provide any i n f o r m a t i o n on how wide c l e a r - c u t s should be nor how wide reserve timber s t r i p s between open areas should be. - 5 -2. THEORY AND HYPOTHESES To achieve the objectives, a general theory was established and then tested with s p e c i f i c research hypotheses. The general theory i s that: Moose i n the West-Chilcotin Region use wetlands f o r feeding and forests for protective cover. The s p e c i f i c hypotheses tested were: 1. Ho: Moose are randomly d i s t r i b u t e d with r e s p e c t to cover types. Ha: Moose are not randomly d i s t r i b u t e d with respect to cover types. 2. Ho: Moose use of f o r e s t s and wetlands i s random with respect to distance to forest/wetland border. Ha: Moose use of f o r e s t s and wetlands i s not random with respect to distance to forest/wetland border. -6-3. STUDY AREA 3.1 LOCATION The 40,000-ha study area occurs i n the western p o r t i o n of the C h i l c o t i n Region of B.C., east of Tweedsmuir P r o v i n c i a l Park and south of the Itcha and Ilgatchuz mountain ranges (Figure 1). The study area occurs e n t i r e l y i n what was formerly considered the Sub-Boreal Spruce Biogeoclimatic (SBS) Zone (Annas and Coupe 1979) . More recently, however, t h i s area was r e c l a s s i f i e d as a zone d i s t i n c t from the SBS Zone and i s now c l a s s i f i e d as the Sub-Boreal Pine Spruce (SBPS) Zone ( B r i t i s h Columbia Ministry of Forests (BCMOF) 1989). 3.2 CLIMATE The c l i m a t e o f t h e s t u d y a r e a i s c o n t i n e n t a l and characterized by cold, dry winters and cool dry summers. Climate data accumulated between 1951 and 1980 are summarized i n Table 1. The number of days between the l a s t spring f r o s t and f i r s t f a l l f r o s t ranges from 2 to 19 days. The average l a s t spring f r o s t i s J u l y 6, w h i l e t h e a v e r a g e f i r s t f a l l f r o s t i s J u l y 22 (Atmospheric Environment S e r v i c e ) . The c o o l , dry climate i s l a r g e l y a r e s u l t of the area's po s i t i o n i n the rainshadow of the Coast Mountains and i t s r e l a t i v e l y high elevation (1100-1500 m) (BCMOF 1989) . -7-Table 1. C l i m a t e Data f o r Anahim Lake B.C. Monthly Averages 1951-1980. January February March A p r i l May June J u l y August September October November December Mean Snowfall 2 (cm) 31.2 17.7 21.1 10.1 3.0 0.5 N/A N/A 1.0 9.0 25.2 29.4 Mean R a i n f a l l (mm) 4.3 1.6 0.5 2.9 14.6 25.9 N/A N/A 14.9 20.0 10.8 6.4 To t a l P r e c i p i t a t i o n 33.8 23.1 20.3 16.1 15.6 30.8 N/A N/A 17.6 25.0 36.4 32.8 (mm) Number of days w i t h measurable 0 0 0 1 6 8 N/A N/A 5 4 2 2 r a i n f a l l Number of days w i t h measurable 8 7 6 4 1 0 N/A N/A 0 3 6 9 snowfal1 TEMPERATURE (C) Mean D a i l y Maximum -6.2 -0.3 3.1 7.8 13.5 17.2 N/A N/A 15.5 9.3 0.9 -4.4 Mean D a i l y Minimum -21.1 -16.7 -13.0 -6.0 -2.2 1.3 N/A N/A -1.4 -4.2 -11.6 -16.9 Mean D a i l y -13.7 -8.5 -5.0 1.0 5.7 9.2 N/A N/A 7.1 2.6 -5.4 -10.7 Extreme Maximum 13.3 11.7 15.6 18.3 28.3 29.4 30.6 31.1 26.7 23.3 17.8 10.0 Extreme Minimum -44.4 -46.1 -41.7 -16.1 -11.7 -7.8 -2.2 -6.7 -12.2 -20.0 -43.9 -42.2 Atmospheric Environment S e r v i c e (1981). Data not a v a i l a b l e . The remainder of the description of the study area i s taken from BCMOF's (1989) description of the SBPS Zone. 3.3 VEGETATION Upland coniferous f o r e s t s are dominated by pure stands of lodgepole pine. Due to extensive f i r e h i s t o r y , the lodgepole pine stands are generally young, even aged, and often dense. The only other common tree species are white spruce (Picea glauca) a n d t r e m b l i n g a s p e n ( P o p u l u s t r e m u l o i d e s ) . W h i t e spruce-dominated f o r e s t s occur on many of the mo i s t s i t e s throughout the SBPS Zone, but these spruce f o r e s t s are usually small and border only non-forested wetlands (BCMOF 1989). The t h e o r e t i c a l c l i m a t i c climax t r e e s p e c i e s i s white spruce, but the dominance of lodgepole pine on mesic s i t e s has been maintained by recurrent w i l d f i r e s . In the portion of the zone where t h e s t u d y a r e a l i e s , t he abundance o f p i n e r e g e n e r a t i o n and lack of spruce regeneration on mesic s i t e s , suggests that lodgepole pine i s the c l i m a t i c climax s p e c i e s (BCMOF 1989). The undergrowth vegetation of lodgepole pine f o r e s t s i s g e n e r a l l y low growing and s p a r s e l y d i s t r i b u t e d , c o n s i s t i n g p r i m a r i l y of b u f f a l o - b e r r y (Shepherdia canadensis) , common ju n i p e r (Juniperus communis), and k i n n i k i n n i c k (Arctostaphylos  uva-ursi). Cover of the former two plants i s generally less than 1%. Other vegetation i s r e s t r i c t e d to lichens (primarily Cladina spp. and Cladonia spp.) and mosses. -10-Non-forested wetlands of v a r y i n g s i z e are i n t e r s p e r s e d throughout the study area. Shrub-carrs dominated by bog-birch (Betula qlandulosa) and willows (Salix spp.), shrub-fens with various willow and sedge (Carex spp.) species and various types of sedge-fens are common wetland communities. Grass-dominated or sedge-dominated meadows are also abundant (BCMOF 1989). 3.4 SOILS S o i l development i n the SBSP Zone i s r e l a t i v e l y weak and s o i l s on z o n a l s i t e s are p r i m a r i l y B r u n i s o l s and L u v i s o l s . B r u n i s o l i c Gray L u v i s o l s and Orthic D y s t r i c B r u n i s o l s are the most common s o i l s on the dominant morainal deposits. The surface organic layer (humus form) i s t y p i c a l l y <4 cm and has very slow rates of decomposition. On imperfectly and poorly drained s i t e s , common s o i l s are gleyed subgroups of B r u n i s o l i c and L u v i s o l i c s o i l s or Gleysols. (BCMOF 1989). -11 4. METHODS 4.1 Habitat Selection To t e s t the hypotheses of random use of cover types, a s e l e c t i o n index was used. S e l e c t i o n of a cover type c l a s s occurred when the frequency of use i n that c l a s s exceeded the f r e q u e n c y of use expected with random use. The technique involved defining cover type classes, determining a v a i l a b i l i t y of each cover type class, determining use of each cover type cl a s s , and then comparing use and a v a i l a b i l i t y w i t h a c h i - s q u a r e goodness of f i t t e s t (Marcum and Loftsgaarden 1980). I f t h i s t e s t was s i g n i f i c a n t , B o n f e r r o n i s i m u l t a n e o u s c o n f i d e n c e i n t e r v a l s were constructed to determine which cover types were used d i f f e r e n t l y from a v a i l a b i l i t y (Marcum and L o f t s g a a r d e n 1980). 4.2 Cover Type Descriptions Cover types a v a i l a b l e to moose i n t h i s study area were divided into forest and wetland. For purposes of t h i s study a 2 fo r e s t habitat had >5 trees/100 m . A wetland was defined as a 2 s i t e with <5 trees/100 m (unless the trees were mature) and did not n e c e s s a r i l y r e f l e c t the amount of water present. Wetland c l a s s e s included grass, sedge, spruce wetland, low b i r c h , low willow, low shrub, t a l l willow, and sedge/shrub. The f o r e s t category was divided into age and species classes. F o r e s t c l a s s e s i n c l u d e d immature, i n t e r m e d i a t e , mature and over-mature lodgepole pine, immature, intermediate, and mature -12-spruce, mature lodgepole pine/spruce mix, and trembling aspen. Because moose l i k e l y select areas based on physical structure and not because of tree species, classes of diameter-at-breast height (dbh), canopy c l o s u r e , d e n s i t y , h e i g h t , shrub c o v e r and distance-to-wetland edge were also measured. 4.2.1 Forest Habitats Except f o r immature lodgepole p i n e , f o r e s t c over types t y p i c a l l y provide l i t t l e or no browse, but may serve as shelter from predators and environmental extremes. i) Immature Lodgepole Pine s i t e s are dominated by lodgepole pine trees <5 m t a l l . The proximity of the branches to the ground allow them to serve as a food source f o r moose. These s i t e s are t y p i c a l l y dense (average = 25-29 stems/100 m 2). i i ) I n t e r m e d i a t e Lodgepole Pine s i t e s a re dominated by lodgepole pine trees >5 m, but <12 cm dbh. i i i ) Mature Lodgepole Pine s i t e s are dominated by trees >5 m t a l l and >12 cm dbh. iv) Overmature Lodgepole Pine s i t e s consist of well-spaced 2 lodgepole pine trees (<5/100 m ) usually exceeding 25 cm dbh and i n poor condition. Canopy closure values are <5% and l o d g e p o l e p i n e s e e d l i n g s a r e p r e s e n t i n t h e understory. v) Immature Spruce s i t e s are dominated by spruce trees <5 m t a l l . 13 vi) Intermediate Spruce s i t e s are dominated by spruce trees >5 m t a l l and <12 cm dbh. v i i ) Mature Spruce s i t e s are dominated by spruce trees >5 m t a l l and >12 cm dbh. v i i i ) Mixed Lodgepole Pine/Spruce s i t e s have a nearly equal d e n s i t y of l o d g e p o l e p i n e and s p r u c e . Only mature lodgepole pine/spruce mixes were observed. 2 ix) Aspen s i t e s contain >5 aspen trees/100 m . 4.2.2 Wetland Habitats These cover types t y p i c a l l y p r o v i d e food, but l i t t l e s h e l t e r . Their names are d e s c r i p t i v e , r e f l e c t i n g the dominant vegetation within the p l o t . i) Grass s i t e s are dry, with Poa spp., and pinegrass (Calamoqrostis rubescens) dominating. i i ) Sedge s i t e s are moist and may or may not contain standing water. Sedges (Carex spp.) and sloughgrass (Beckmannia  syzigachne) dominate. i i i ) Spruce Wetland s i t e s contain <5 trees (spruce)/100 m . These s i t e s are usually moist, and contain abundant browse and sedge. iv) Low Birch s i t e s contain bog bir c h (Betula qlandulosa) <2 m i n height. v) Low Willow s i t e s contain willows ( S a l i x spp.) <2 m i n height. v i ) Low Shrub s i t e s contain a mix of low b i r c h and low willow. -14-v i i ) T a l l Willow s i t e s contain willows >2 m i n height. v i i i ) Hay Meadow s i t e s are grass or sedge meadows which are harvested. ix) Sedge/Shrub s i t e s are predominantly sedge but c o n t a i n b i r c h /willow shrubs (<0.5 m t a l l ) . These s i t e s are usually wet. 4.3 Cover Type A v a i l a b i l i t y Assessment The method of Mar cum and Loftsgaarden (1980) was used to assess the a v a i l a b i l i t y of various wetland and f o r e s t h a b i t a t parameters w i t h i n the study area. - P l o t s (254) were randomly al l o c a t e d throughout the 40,000-ha study area. This number of pl o t s was the minimum required to estimate the proportion of each cover type/attribute with 90% confidence and a maximum er r o r of 5%, based on a binomial d i s t r i b u t i o n (Mendenhall 1971:195). Randomization was ac h i e v e d by f i r s t c o n s t r u c t i n g a 1:64,000-scale b l a c k - a n d - w h i t e a i r photograph mosaic and overlaying i t with a g r i d . The gr i d , i n turn, was ov e r l a i n with a sheet of c l e a r acetate. Computer generated random x and y values were plotted on the acetate, using the g r i d beneath as a guide. The g r i d was removed, r e v e a l i n g the l o c a t i o n of the p l o t s . These locations were transferred to 1:20,000-scale color a i r photographs to f a c i l i t a t e p l o t location i n the f i e l d . At each l o c a t i o n , a 10 x 10 m p l o t was e s t a b l i s h e d . Measured habitat parameters of forest cover a t t r i b u t e s included tree species, age, overstory height, stem density, canopy cover, average dbh, distance-to-wetland edge, and cover (%) and height -15-of shrub species. Wetland habitat a t t r i b u t e s included cover (%) and height of browse species and distance-to-forest edge. 4.4 Physical Structure Average dbh was determined by measuring with a D-tape, to the nearest 1.0 cm, three representative trees ( o c u l a r l y estimated) i n each p l o t and then a s s i g n i n g the r e s u l t t o one of the following eight categories (cm): Canopy closure of each plot was ocularly estimated and assigned to one of the following classes: l , 3, 5, 7, 10, 15, 20, or >25%. Percent cover charts obtained from Walmsley et. a l . (1980) were used as reference to f a c i l i t a t e v i s u a l estimations. Density of each p l o t was measured by counting the number of trees 2 i n the 100 m p l o t . Values were then assigned to one of the 2 following eight classes (number of trees/100 m ): 0-5 5-7 7-10 10-12 13-16 17-20 21-24 >25 0-4 5-8 9-11 12-14 15-19 20-24 25-29 >30 -16-Average tree height of each plot was measured to the nearest 0.1 meter with a clinometer, and values then assigned to one of the following eight classes (m): 0.0-2.0 2.1-5.0 5.1-10.0 10.1-12.0 12.1-14.0 14.1-16.0 16.1-18.0 >18.0 Shrub cover was o c u l a r l y estimated using percent cover charts obtained from Walmsley et a l . (1980) and then assigned to one of the following classes: 1, 3, 5, 7, 10, 15, 20, >25%. Distance-to-wetland/forest edge was measured to the nearest 5 m on an a i r photograph or measured i n the f i e l d with a hip chain and then assigned to one of the following eight classes (m): Forest age was determined by counting core rings taken at breast height of the three representative trees. Each r i n g was assumed to represent one year. Values were assigned to one of the following classes (years): Percent cover of shrubs was o c u l a r l y estimated u s i n g percent cover charts from Walmsley et a l . (1980). Values were assigned to one of the following classes: 1, 3, 5, 7, 10, 15, 20, >25%. 0-20 21-40 41-60 61-80 81-100 101-149 150-200 >200 0-20 21-40 41-60 61-80 81-100 101-120 121-140 >141 -17-Height of shrubs was determined by measuring the height of a r e p r e s e n t a t i v e shrub of each species to the n e a r e s t 1.0 cm. Values were assigned to one of the following classes (m): 0.0-0.5 0.6-1.0 1.1-1.5 1.6-2.0 2.1-3.0 >3.1 4.5 Habitat Use To evaluate which habitat classes moose used, 10 moose were r a d i o - c o l l a r e d and monitored throughout the w i n t e r months. Habitat variables associated with each location were recorded. 4.5.1 Moose Capture On 12-13 January 1987, 10 adult female moose were chemically immobilized and f i t t e d with r a d i o - c o l l a r s . Of these 10 c o l l a r s , 2 became n o n - f u n c t i o n a l w i t h i n 2 weeks. A l l other c o l l a r s functioned normally f o r the duration of study. On 2 5 January 1988, 2 ad d i t i o n a l adult female moose were chemically immobilized and f i t t e d with r a d i o - c o l l a r s . 4.5.2 Moose Locations During the 1987 winter season, preliminary data on home range s i z e and g e n e r a l h a b i t a t use were r e c o r d e d f o r e i g h t r a d i o - c o l l a r e d moose. Five relocation flights/month were flown on average between January and A p r i l . Location, a c t i v i t y , forest 18 o r wetland cover type o c c u p i e d , as w e l l as d i s t a n c e from f o r e s t / w e t l a n d edge were r e c o r d e d f o r each o b s e r v a t i o n . Approximate locations were marked on 1:20 000 scale topographical maps. Other information recorded with each l o c a t i o n included date, time, temperature, and cloud cover. During the 1988 winter season more detailed habitat use data were c o l l e c t e d . Approximately ten f l i g h t s per month were flown between January and March. In addition to the above information, each l o c a t i o n was marked on 1:15 000 scale color a i r photographs. The large scale photographs allowed exact moose l o c a t i o n s to be i d e n t i f i e d . Subsequent sampling i n A p r i l documented s p e c i f i c f o r e s t and wetland attributes at these locations. The variables measured include d a l l those measured during determination of habitat a v a i l a b i l i t y . Moose were located using a Telonics receiver with a 4 element Yagi antenna mounted on each wing s t r u t of a S k i equipped airplane. In 1987, a Cessna 172 was used, while i n 1988 a Piper Pacer PA22 was used. In both seasons, over 9 0% of r e l o c a t i o n s resulted i n v i s u a l i d e n t i f i c a t i o n of the c o l l a r e d animal. 4.6 S t a t i s t i c a l Analyses The Chi-square t e s t was used to determine i f use d i f f e r e d from a v a i l a b i l i t y . This test can function properly providing no more than 20% of expected o b s e r v a t i o n s i n each c e l l are <5 (Roscoe and Byars 1971) . The n u l l h y p o t h e s i s of random d i s t r i b u t i o n with respect to each habitat variable was tested at P = 0.05. -19-I f the n u l l hypothesis was rejected, the next step was to determine which of the classes the moose p r e f e r r e d . This was done by obtaining 100(1-alpha)% simultaneous confidence i n t e r v a l s f o r the difference between use and a v a i l a b i l i t y f o r each category (Marcum and Loftsgaarden 1980). The confidence i n t e r v a l around the difference between use and a v a i l a b i l i t y equals: <W ± Zl-alpha/2k * [Vl-V/». + V^W*'* where: P = proportion of the randomly located p l o t s that l i e i n the category i n question, P u = proportion of moose locations that l i e i n the category i n question, k = the number of comparisons, n = number of plots randomly d i s t r i b u t e d over the study c l area, n u = t o t a l number of moose locations. I f the i n t e r v a l contains zero, then use equalled a v a i l a b i l i t y . I f both s i d e s of the i n t e r v a l are negative then use exceeded a v a i l a b i l i t y ; i f both sides of the i n t e r v a l are p o s i t i v e then use was less than a v a i l a b i l i t y . Alpha r e f e r s to the desired type l error f o r the simultaneous set of confidence i n t e r v a l s . Using the Bonferroni approach, the alph a l e v e l f o r each of the k c o n f i d e n c e i n t e r v a l s e quals alpha/k; therefore, i f a large number of simultaneous confidence i n t e r v a l s are being formed, a larger alpha should be chosen so that i n d i v i d u a l i n t e r v a l s are not so wide that they become -20-meaningless. (1-alpha/k can become very large) (Marcum and Loftsgaarden, 1980). Setting the confidence lev e l s for the i n d i v i d u a l comparison to 95% required alpha for the simultaneous set of i n t e r v a l s to be set at 0.4 ( i . e . , 1-alpha/k for 8 comparisons = 1 - 0.4/8 = 95%). Obviously a type l error rate of t h i s magnitude i s unacceptable; however, the two-stage design of t h i s t e s t negates t h i s problem. The Chi-square t e s t performed i n i t i a l l y provided a simultaneous c o n f i d e n c e i n t e r v a l c a l c u l a t e d a t an a l p h a of 0.05. The in d i v i d u a l comparison was not performed unless the Chi-square was f i r s t rejected; therefore, both the simultaneous and i n d i v i d u a l i n t e r v a l s were calculated at an alpha of 0.05. This approach i s equivalent procedurally to the Fisher's protected LSD. Data were analysed on a seasonal basis to obtain a sample s i z e s u f f i c i e n t to conduct chi-square t e s t s . These data were also summarized on a monthly basis, although the sample s i z e was not large enough to s a t i s f y chi-square requirements. 4.7 Moose Home Ranges A l l winter moose locations were entered into the HOME RANGE program on the UBC main frame computer. This program calculates home range s i z e and also provides a p l o t of movements. These p l o t s can be overlaid onto vegetation maps of desired scales. Because the 1987 l o c a t i o n s were only approximates, home ranges based on these data cannot be compared with the 1988 home ranges. These data were useful i n demonstrating whether the same -21-moose wintered i n the same areas. 4.8 Winter Diet A minimum of 30 moose p e l l e t groups were c o l l e c t e d monthly i n the 1987 and 1988 winter f i e l d seasons. These monthly p e l l e t groups were composited and analysed f o r d i e t composition and q u a l i t y . In 1988, at the end of each month, 5 i n d i v i d u a l moose were tracked and fresh p e l l e t groups encountered were c o l l e c t e d . A d d i t i o n a l l y , samples were c o l l e c t e d from browsed shrubs along the moose t r a i l . Care was taken to ensure that the sampled plant part had also been browsed. I f only the shoots of the year were browsed, then only the shoots of the year were c o l l e c t e d . C o l l e c t i n g f e c a l and browse samples by t h i s means enhanced the p r o b a b i l i t y that f e c a l samples correlated with d i e t intake. 4.8.1 Composition Forage composition of moose diets was determined f o r January, February and March of both winters. Samples c o l l e c t e d at the end of each month were frozen u n t i l they could be oven-dried. At the end of each winter the samples were thawed and then oven-dried at 55 C. The samples were subsequently sent to the W i l d l i f e Habitat Laboratory (WHL) at the University of Washington i n Pullman f o r f e c a l fragment analyses. -22-4.8.2 Quality In addition to f e c a l fragment analyses, the WHL also measured crude protein (%), gross energy (c a l / g ) , and % detergent f i b e r (NDF) on the 1988 samples. 4.9 Snow Depth A weather observation station operated by Environment Canada was l o c a t e d at 5 M i l e Ranch i n the center of the study area. This s t a t i o n provided d a i l y snow depth for a l l winter months of 1987 and 1988. -23-5. RESULTS 5.1 HABITAT SELECTION 5.1.1 Cover Type Classes A v a i l a b i l i t y Of the 254 p l o t s , 148 occurred i n forested s i t e s and 105 i n wetland s i t e s (Table 2). One of the forest c l a s s p l o t s was not c l a s s i f i e d and i s excluded from t h i s t a b l e . The t o t a l f o r e s t a v a i l a b l e was 58.5%, while the t o t a l wetland a v a i l a b l e was 41.5%. The f o r e s t category consisted mainly of lodgepole pine (73.6%), p r i m a r i l y mature stands (50%). Spruce stands comprised 25.1% of the f o r e s t , while aspen stands accounted f o r 1.4%. Within wetlands, the low willow c l a s s was most pr e v a l e n t (19%), while the low b i r c h class was the least prevalent (6.7%). Use January - March 1988 A e r i a l t e l e m e t r y of 8 r a d i o - c o l l a r e d moose provided 194 observations of which 49.5% were i n forest cover types and 50.5 % were i n wetland cover types (Table 3). Mature spruce and mature pine were used to a greater extent than any other f o r e s t c l a s s , making up 37.5 and 25.0% of f o r e s t observations, r e s p e c t i v e l y (18.6 and 12.4% of t o t a l observations, r e s p e c t i v e l y ) . L i t t l e use -24-T a b l e 2. A v a i l a b i l i t y of f o r e s t cover type c l a s s e s i n the West-Chilcotin. Class # of Plots % of Study Area % of Category Lodgepole Pine immature 9 3.6 6.1 intermediate 19 7.5 12.8 mature 74 29.2 50.0 overmature 7 2.8 4.7 Pine Total 109 43.1 73.6 Spruce immature 1 0.4 0.7 intermediate 9 3.6 6.1 mature 13 5.1 8.8 pine/spruce 14 5.5 9.5 Spruce Total 37 14.6 25.1 Trembling Aspen 2 0.8 1.4 Forest Total 148 58.5 100.1 Wetland grass 13 5.1 12.4 sedge 17 6.7 16.2 spruce wetland 14 5.5 13.3 low b i r c h 7 2.8 6.7 low willow 20 7.9 19.0 low shrub 16 6.3 15.2 t a l l willow 9 3.6 8.6 sedge/shrub 13 3.6 8.6 Wetland Total 105 41.5 100.0 Tot a l 253 100.0 Forest or Wetland -25-Table 3. A v a i l a b i l i t y and moose use of cover c l a s s e s i n the W e s t - C h i l c o t i n d u r i n g w i n t e r 1988. C l a s s A v a i l a b i l i t y Use Chi square s.c .1. 1 S e l e c t i o n p l o t s 5 i s i g h t i n g s 5 i upper lower Pine * immature 9 3. .6 4 2. .1 50.67 -0. .036 0. ,075 0 interm. 19 7. .5 10 5. ,1 -0. ,057 0. 106 0 mature 74 29.2 24 12, ,4 0, ,132 0. ,368 -overmat. 7 2, .8 6 3. .1 -0, .074 0. ,044 0 Spruce immature 1 0, .4 9 4. .6 -0. ,147 -0. 027 + interm. 9 3. .6 1 0. ,5 0. ,007 0. ,094 -mature 13 5, .1 36 18. ,6 -0. ,394 • -0. ,180 + p/s 14 5, .5 4 2. ,1 -0, ,009 0. ,115 0 Aspen 2 0, .8 2 1. .0 -0, ,041 0. .027 0 F r s t T o t a l 148 58. .5 96 49. ,5 Wetland * grass 13 5. .1 0 0. .0 49.19 0, .061 0. ,187 -sedge 17 6 .7 3 1. .5 0. .053 0, ,210 -spruce 14 5. .5 44 22. .7 -0. .434 • -0. ,198 + low b i r c h 7 2. .8 7 3. ,6 -0. ,075 0. 065 0 l.w. 20 7, .9 8 4. ,1 0. ,016 0. ,201 -low shrub 16 6, .3 23 11, ,9 -0, .191 0. ,026 0 4 t.w. 9 3. .6 11 5, .7 -0, .109 0. ,056 0 sdg/shrb 9 3. .6 2 1. .0 0. .005 0. ,126 0 Wtld T o t a l 105 41. .5 98 50. .5 Total 253 100. .0 194 100. ,0 * S i g n i f i c a n t a t p. < 0.05. Refers t o the simultaneous comparison. ^ Simultaneous con f i d e n c e i n t e r v a l f o r d i f f e r e n c e between a v a i l a b i l i t y and use. pine/spruce 1ow wi11ow 4 t a l l w i l l o w -26-occurred i n immature pine, pine/spruce, or aspen. Spruce wetlands were the most o f t e n - u s e d cover c l a s s , accounting f o r 22.7% of t o t a l observations and 4 5% of wetland observations. The low shrub class was used next most frequently, accounting f o r 11.9% of observations. This c l a s s , combined with the low b i r c h and low w i l l o w , accounted f o r 19.6% of a l l observations or 40% of a l l wetland observations. Use of grass, sedge, and sedge shrub classes was low. January - March 1987 A e r i a l t e l e m e t r y of 8 r a d i o - c o l l a r e d moose provided 134 observations, of which 56.0% occurred i n forest cover types, and 44.0% occurred i n wetland cover types (Table 4). Of the forest cover types, intermediate pine was used most exte n s i v e l y (46.7% of f o r e s t e d observations) . Aspen was used le a s t of a l l forest cover classes. Selection Of the 17 cover c l a s s e s a v a i l a b l e , only immature spruce, mature spruce, and spruce wetlands were used i n p r o p o r t i o n s greater than a v a i l a b i l i t y (Table 4). Mature pine, intermediate spruce, and grass, sedge, and t a l l willow wetlands were a l l used les s than expected from random d i s t r i b u t i o n . -27-Table 4. Moose use of cover c l a s s e s i n the W e s t - C h i l c o t i n during w i n t e r 1987. C l a s s # of Observati ons % of Total Observations Pine immature 4 3. .0 i n t e r m e d i a t e 35 26. .1 mature 16 11, .9 Spruce 6 4. .5 Mature pine/spruce 12 . 9, .0 Aspen 2 . 1, .5 Fo r e s t T o t a l 75 56. ,0 Wetland 59 44. .0 Total 134 100. ,0 5.1.2 Distance-to-Edge A v a i l a b i l i t y The amount of area w i t h i n each d i s t a n c e - t o - e d g e c l a s s generally decreased as distance from edge increased (Table 5) . T h i r t y - o n e p e r c e n t of the study area i s w i t h i n 20 m of a forest/wetland edge, whereas only 6.7% of the study area i s between 150 and 2 00 m. Because of the open-ended nature of the cl a s s , 14.6% of the study area was >200 m away from the edge. Use In both winters, use was concentrated near the wetland/forest edge. In 1988, 91.7% of a l l observations were within 100 m of t h e f o r e s t / w e t l a n d edge ( T a b l e 5 ) . In 1987, 90.6% o f observations were within 100 m of the forest-wetland edge (Table 6). In both years, >50% of a l l observations were within 20 m of -28-the edge (p_ < 0.05). In 1987, 64.1% of observations were within 20 m while only 8.5% were >200 m from the edge. In 1988, 54% were within 20 m, while only 1.4% were >200 m away from the edge. Table 5. A v a i l a b i l i t y and moose use of distance-to-edge c l a s s e s i n the West C h i l c o t i n d u r i n g w i n t e r 1988. D i s t . t o A v a i 1 a b i 1 i t y Use Chi S.C .1 1 S e l e c t i o n edge (m) p l o t s 0 i s i g h t i n g s % square upper lower 0-20 79 31. .1 112 54.1 54.75* -0 .319 • -0 .141 + 21-40 30 11. .8 35 16.9 -0 .116 0 .014 0 41-60 28 11. .0 23 11.1 -0 .058 0 .057 0 61-80 25 9. .8 15 7.2 -0 .025 0 .077 0 81-100 20 7. .8 5 2.4 0 .015 0 .094 -101-149 18 7. .1 3 1.4 0 .021 0 .092 -150-200 17 6. .7 11 5.3 -0 .030 0 .057 0 >200 37 14. .6 3 1.4 0 .085 0 .178 -Total 254 99. .9 207 99.8 * S i g n i f i c a n t at j) < 0. ,05. Refers to simultaneous comparison. Simultaneous confidence i n t e r v a l f o r d i f f e r e n c e between a v a i l a b i l i t y and use. Table 6. A v a i 1 a b i 1 i t y and moose use of distance-to-edge c l a s s e s i n the W e s t - C h i l c o t i n d u r i n g w i n t e r 1987. D i s t . t o A v a i l a b i l i t y Use Chi S.C.I S e l e c t i o n edge (m) p l o t s °/ 0 s i g h t i n g s % square upper lower 0-20 79 31. 1 75 64.1 49.23 0.434 -0. .226 + 21-40 30 11. 8 8 6.8 -0. .011 0, .110 0 41-60 28 11. 0 4 3.4 0. .025 0. .127 -61-80 25 9. 8 16 13.7 -0. .111 0. .034 0 81-100 20 7. 8 3 2.6 0. .009 0. .097 -101-149 18 7. 1 0 0.0 0. .039 0. .102 -150-200 17 6. 7 1 0.9 0. ,023 0. ,093 ->200 37 14. 6 10 8.5 -0. ,006 0. .127 0 Total 254 99. 9 117 100.0 * j S i g n i f i c a n t a t p. < 0.05. Refers t o simultaneous comparison. Simultaneous confidence i n t e r v a l f o r d i f f e r e n c e between a v a i l a b i l i t y and use. -29-Selection 1988 Areas within 20 m of the edge were selected (p. < 0.05), while areas between 21 and 80 m were used i n p r o p o r t i o n to t h e i r a v a i l a b i l i t y . With the exception of the 150-200 m c l a s s , a l l areas >100 m from the edge were selected against (p_ < 0.05). 1987 Areas within 20 m of an edge were used i n greater proportion t h a n t h e i r a v a i l a b i l i t y (p_ < 0.05). Each of t h e o t h e r distance-to-edge classes were used either equal to or l e s s than t h e i r a v a i l a b i l i t y . 5.1.3 Distance-to-Wetland A v a i l a b i l i t y The 0-2 0 m c l a s s i s most abundant, comprising 25.5% of the f o r e s t (Table 7 ) . The 21-40 m c l a s s i s l e a s t abundant, comprising 6.7% of the forest. Use S i m i l a r t o the d i s t a n c e - t o - e d g e c l a s s e s , use of the distance-to-wetland classes dropped as the distance to wetland increased beyond 20 m (Tables 7,8). In both 1987 and 1988 use of fore s t s was concentrated near the forest/wetland border. In 1987 and 1988, 86.6% and 87.0% of f o r e s t e d observations occurred within 100 m of the wetland, respectively. In both years, more than 50% of f o r e s t e d o b s e r v a t i o n s were w i t h i n 20 m of the wetland. Beyond 200 m from the wetland edge, use was noteable i n 1987, but was n e g l i g i b l e i n 1988. -30-Table 7. A v a i l a b i l i t y and moose use of d i s t a n c e - t o - w e t l a n d c l a s s e s i n the W e s t - C h i l c o t i n d u r i n g w i n t e r 1988. D i s t . t o A v a i l a b i l i t y Use Chi S.C.I. S e l e c t i o n w t l n d (m) p l o t s % s i g h t i n g s % square upper lower 0-20 38 25. .5 51 21-40 10 6. .7 18 41-60 17 11. .4 8 61-80 18 12. .1 8 81-100 14 9. .4 2 101-149 14 9. .4 2 150-200 11 7 .4 9 >200 27 18, .1 2 Total 149 100. .0 100 51, .0 43.05 -0. 375 -0. ,135 + 18, .0 -0. ,198 -0. ,028 + 8, .0 -0. 040 0. ,108 0 8, .0 -0. ,034 0. ,115 0 2, ,0 0. ,020 0. ,128 -2, .0 0. ,020 0. ,128 -9, ,0 -0. ,086 0. ,054 0 2, .0 0, ,094 0, .229 -100.0 * S i g n i f i c a n t at p. < 0.05. Refers t o simultaneous comparison. Simultaneous con f i d e n c e i n t e r v a l f o r d i f f e r e n c e between a v a i l a b i l i t y and use. Table 8. A v a i l a b i l i t y and moose use of di s t a n c e - t o - w e t l a n d c l a s s e s i n the W e s t - C h i l c o t i n d u r i n g w i n t e r 1987. D i s t . t o A v a i l a b i l i t y Use Chi S.C.I. S e l e c t i o n w t l n d (m) p l o t s % s i g h t i n g s % square upper lower * 0-20 38 25. ,5 39 52. ,7 26.79 -0. ,406 -0. 138 + 21-40 10 6. .7 5 6. ,8 -0. ,070 0. 069 0 41-60 17 11. .4 4 5. ,4 -0. ,012 0. .133 0 61-80 18 12. ,1 13 17. ,6 -0. ,156 0. 046 0 81-100 14 9. .4 3 4. ,1 -0. ,011 0. ,118 0 101-149 14 9, .4 0 0. ,0 0. ,047 0. ,141 -150-200 11 7. .4 1 1, .4 0. O i l 0. ,110 ->200 27 18, .1 9 12, .1 -0, .037 0. ,156 0 To t a l 149 100. .0 74 100. ,1 * S i g n i f i c a n t at Q < 0.05. Refers t o simultaneous comparison. Simultaneous con f i d e n c e i n t e r v a l f o r d i f f e r e n c e between a v a i l a b i l i t y and use. -31-Selection 1988 Selection of distance-to-wetland classes generally decreased as distance from wetland increased. Within 40 m, s e l e c t i o n was p o s i t i v e . Between 40 and 80 m, use equalled a v a i l a b i l i t y . With the e x c e p t i o n of the 150-200 m c l a s s , areas >80 m from the wetland were used less than a v a i l a b i l i t y . 1987 Selection of distance-to-wetland classes also shows a general decreasing trend as distance from wetland increases. Use within 20 m o f t h e w e t l a n d was p r o p o r t i o n a t e l y g r e a t e r t h a n a v a i l a b i l i t y . Between 20 and 100 m use was e q u a l t o a v a i l a b i l i t y . With the exception of the >200 m c l a s s , use beyond 100m was less than a v a i l a b i l i t y . 5.1.4 Distance-to-Forest Edge A v a i l a b i l i t y T h i r t y - n i n e percent of the area occupied by wetlands i s wit h i n 20 m of a f o r e s t edge, while 80% i s w i t h i n 100 m of a for e s t edge. (Table 9). Use As with other edge measurements, use was concentrated near the edge i n both 1988 and 1987, when use within 2 0 m was 57.0% and 83.7% respectively (Tables 9,10). In 1988, use declined as distance from edge increased. The few numbers of observations i n -32-1987 obscure t h i s trend, but i n both cases use of wetlands beyond 100 m from the forest edge was n e g l i g i b l e . Selection 1988 Selection of distance-to-forest classes was s i m i l a r to other d i s t a n c e - t o - e d g e v a r i a b l e s . As d i s t a n c e from f o r e s t edge increased, the degree of selection decreased. P o s i t i v e s e l e c t i o n o c c u r r e d i n the 0-20 m c l a s s . Use was p r o p o r t i o n a l t o a v a i l a b i l i t y f o r classes between 21 and 200 m, whereas use was l e s s than a v a i l a b i l i t y for the >200 m c l a s s . Table 9. A v a i l a b i l i t y and moose use of distance-to-forest classes in the West-Chilcotin during winter 1988. Dist. to forest (m) A v a i l a b i l i t y Use Chi S .C. , I.~ Selection plots 5 i sightings % square upper 1 ower 0-20 41 39. .0 61 57, .0 16.93 -0. ,312 --0. ,047 + 21-40 20 19, .0 17 15, .9 -0. ,071 0. ,134 0 41-60 11 10. .5 15 14, .0 -0. ,124 0. ,053 0 61-80 7 6, .7 7 6, .5 -0. ,066 0. ,068 0 81-100 6 5. .7 3 2, .8 -0. ,025 0. ,083 0 101-149 4 3, .8 1 0, .9 -0. ,012 0. ,070 0 150-200 6 5, .7 2 1, .9 -0. ,013 0. ,090 0 >200 10 9, .5 1 0, .9 0. ,027 0. ,145 -Total 105 99, .9 107 99, .9 Significant at p_ < 0.05. Refers to simultaneous comparison. Simultaneous confidence interval for difference between a v a i l a b i l i t y and use. -33-Table 10. A v a i l a b i l i t y and moose use of d i s t a n c e - t o - f o r e s t c l a s s e s i n the W e s t - C h i l c o t i n d u r i n g w i n t e r 1987. D i s t . t o f o r e s t (m) A v a i l a b i l i t y Use Chi square S.C A1- S e l e c t i o n p l o t s % s i g h t i n g s % upper lower 0-20 41 39.0 36 83. .7 27.75 -0. 591 --0.302 + 21-40 20 19.0 3 7. .0 0. .014 0.228 -41-60 11 10.5 0 0. ,0 0. .046 0.163 -61-80 7 6.7 3 7. .0 -0. .093 0.087 0 81-100 6 5.7 0 0. .0 0. 013 0.102 -101-149 4 3.8 0 0. ,0 0. .001 0.075 -150-200 6 5.7 0 0. ,0 0. .013 0.102 ->200 10 9.5 1 2. ,3 -0. ,000 0.144 0 To t a l 105 99.9 43 100. 0 S i g n i f i c a n t a t p_ < 0.05. Refers to simultaneous comparison. Simultaneous confidence i n t e r v a l f o r d i f f e r e n c e between a v a i l a b i l i t y and use. 5.1.5 Forest Age A v a i l a b i l i t y The m a j o r i t y of f o r e s t s are between 41 and 100 years o l d , with low proportions i n both younger and older c l a s s e s (Table l l ) • use Although the 21-40 year old class was the second most highly used c l a s s , most use occurred i n o l d e r f o r e s t s . S i x t y - s i x percent of forest observations occurred i n forests >60 years old. The 61-80 year o l d c l a s s was used more o f t e n than any other c l a s s , while the 0-20 year old class was used l e a s t . -34-Table 11. A v a i l a b i l i t y and moose use of f o r e s t age c l a s s e s i n the W e s t - C h i l c o t i n d u r i n g w i n t e r 1988. Age (years) A v a i l a b i l i t y Use Chi square S .C. I . ~ S e l e c t i on p l o t s % s i g h t i n g s % upper lower 0-20 4 2.8 6 8. ,2 20.10* -0. ,123 0. 014 0 21-40 19 13.1 12 16. ,4 -0. ,135 0. 068 0 41-60 34 23.4 7 9. ,6 0. .042 0. 235 -61-80 39 26.9 14 19. ,2 -0. ,038 0. 193 0 81-100 29 20.0 12 16. ,4 -0. ,071 0. 143 0 101-120 9 6.2 7 9, ,6 -0, .112 0. ,044 0 121-140 2 1.4 7 9, .6 -0, .152 • -0. ,012 + >141 9 6.2 8 11, .0 -0, ,129 0. ,034 + Total 145 100.0 73 100, .0 * S i g n i f i c a n t at p. < 0.05. Refers t o simultaneous comparison. Simultaneous con f i d e n c e i n t e r v a l f o r d i f f e r e n c e between a v a i l a b i l i t y and use. Selection With the exception of the 41-60, 120-140, and >140 cl a s s e s , f o r e s t age classes were used i n proportion to t h e i r a v a i l a b i l i t y . The 41-60 year old class was used le s s , while the l a t t e r two were used more than expected from random use. 5.1.6 Forest Density A v a i l a b i l i t y 2 The 20-24 stems/100 m c l a s s i s l e a s t p r e v a l e n t at 7.5%, while the >30 class i s most highly represented at 19.9% (Table 12). -35-Use S i x t y - f i v e percent of forest observations occurred i n forests 2 w i t h d e n s i t i e s between 9 and 19 stems/100 m . Outside t h i s range, use dropped o f f dramatically. Selection Use generally equalled a v a i l a b i l i t y . In the 9-11 stems/100 m c l a s s , however, use exceeded a v a i l a b i l i t y , while i n the >30 cla s s use was less than a v a i l a b i l i t y . Table 12. A v a i l a b i l i t y and moose use of f o r e s t d e n s i t y c l a s s e s i n the W e s t - C h i l c o t i n d u r i n g w i n t e r 1988. Densi t y (stems/100 m ) A v a i 1 a b i 1 i t y Use Chi square S . C . I . 1 S e l e c t i o n p l o t s % s i g h t i n g s % upper lower 0-4 13 8.9 4 5.2 18.24* -0. .030 0.106 0 5-8 23 15.8 7 9.1 -0, .020 0.155 0 9-11 21 14.4 22 28.6 -0. .257 -0.025 + 12-14 18 12.3 15 19.5 -0. .174 0.033 0 15-19 13 8.9 13 16.9 -0. ,175 0.017 0 20-24 11 7.5 5 6.5 -0. .059 0.081 0 25-29 18 12.3 4 5.2 -0. .001 0.145 0 >30 28 19.9 7 9.1 0. .011 0.193 -T o t a l 145 100.0 77 100.1 S i g n i f i c a n t a t p. < 0.05. Refers to simultaneous comparison. Simultaneous confidence i n t e r v a l f o r d i f f e r e n c e between a v a i l a b i l i t y and use. -36-5.1.7 Forest Height A v a i l a b i l i t y The majority of forests were >5 m i n height with 79% between 5 and 16 m (Table 13). Use With the exception of the 0-2 m-class, use occurred i n a l l f o r e s t height c l a s s e s , ranging from 6.6% f o r the 16.1-18.0-m clas s to 21.1% for the 10.1-12.0 m-class. The majority of use occurred i n forests between 5 and 16 m i n height which accounted f o r 72.4% of forest observations. Selection Use of each f o r e s t height c l a s s was p r o p o r t i o n a l to the a v a i l a b i l i t y of each cl a s s . 5.1.8 Forest DBH A v a i l a b i l i t y Eighty percent of forests were >12 cm dbh. The 16.1-2 0.0-cm cl a s s was most highly represented at 26.9% (Table 14). Use Use of large diameter trees was greater than small diameter trees. Seventy two percent of observations occurred i n forests with a dbh >16 cm. -37-Table 13. A v a i l a b i l i t y and moose use of f o r e s t height c l a s s e s i n the W e s t - C h i l c o t i n d u r i n g w i n t e r 1988. Height A v a i l a b i l i t y Use Chi (meters) p l o t s % s i g h t i n g s % square 0.0-2.0 3 2, .1 0 0, .0 2.1-5.0 5 3, .5 9 11. .8 5.1-10.0 33 23, .2 14 18, .4 10.1-12.0 27 19, .0 16 21, .1 12.1-14.0 31 21, .8 10 13. .2 14.1-16.0 22 15, .4 15 19, .7 16.1-18.0 13 9, .2 5 6. .6 >18.0 8 5, .6 7 9, ,2 Total 142 99, .8 76 100. .0 11.40 Table 14. A v a i l a b i l i t y and moose use of f o r e s t DBH c l a s s e s i n the W e s t - C h i l c o t i n d u r i n g w i n t e r 1988. DBH A v a i l a b i l i t y Use Chi S.C.I." S e l e c t i o n (cm) p l o t s 0 i s i g h t i n g s % square upper lower 0-5.0 3 2. .1 5 6. .7 17.84 -0. ,107 0. ,015 0 5.1-7.0 4 2, .8 2 2, .7 -0. ,044 0. ,046 0 7.1-10.0 12 8, .3 7 9, .3 -0. ,090 0. ,069 0 10.1-12.0 9 6, .2 1 1, .3 0. ,002 0. ,096 -12.1-16.0 34 23, .4 6 8, .0 0. ,062 0. ,247 -16.1-20.0 39 26, .9 20 26, .7 -0, ,121 0. ,126 0 20.1-24.0 26 17, .9 14 18, .7 -0. ,115 0. ,101 0 >24.0 18 12, .4 20 26, .7 -0. ,256 --0. ,029 + T o t a l 145 100 .0 75 100, .1 * S i g n i f i c a n t at p. < 0.05. Refers t o simultaneous comparison. Simultaneous con f i d e n c e i n t e r v a l f o r d i f f e r e n c e between a v a i l a b i l i t y and use. -38-Selection The >24-cm c l a s s was the only c l a s s i n which use exceeded a v a i l a b i l i t y . Forests with DBH between 10 and 16 cm were used les s than expected. A l l other classes were used i n proportion to t h e i r a v a i l a b i l i t y . 5.1.9 Forest Canopy Closure A v a i l a b i l i t y The 3% canopy closure c l a s s was most highly represented at 21.1%. The majority of forests (58%) had canopy closures between 3% and 7% (Table 15). Use Eighty-eight percent of observations were i n f o r e s t s with canopy c l o s u r e values >5%, with the 5% canopy c l o s u r e c l a s s r e c e i v i n g more use than any other single c l a s s . Selection Most canopy closure classes were used i n proportion to t h e i r a v a i l a b i l i t y . The 3% class was used le s s , and the >25% c l a s s was used more. 39-Table 15. Availability and moose use of forest canopy closure classes in the West-Chilcotin during winter 1988. Canopy clsr Availability Use Chi S.C.I. Selection (%) plots 5 i sightings % square upper lower 1 12 9. .8 3 3, .9 19.77 -0. .009 0. ,127 0 3 26 21. ,1 6 7, .8 0. ,040 0. ,227 -5 23 18. ,7 19 24, .7 -0. ,178 0, ,059 0 7 22 17, .9 9 11. .7 -0. ,037 0. ,161 0 10 12 9, .8 11 14, .3 -0. ,139 0. ,049 0 15 17 13, ,8 9 11, .7 -0. ,073 0. ,116 0 20 8 6. .5 12 15, .6 -0. ,183 0. ,001 0 >25 3 2, .4 8 10, .4 -0. ,153 • -0. .006 + Total 123 100. ,0 77 100. .1 * Significant at Q < 0.05. Refers to simultaneous comparison. Simultaneous confidence interval for difference between availability and use. 5.1.10 Forest Shrub Cover A v a i l a b i l i t y Forests have low shrub cover. Sixty four percent of forests have <1% shrub cover, while 81.2% of forests have <5% shrub cover (Table 16). Use No pattern of use of shrub cover classes i s obvious. The 1% c l a s s received the most use at 37.7%. The le a s t used class was the >2 5% class which accounted for 3.9% of observations. Selection Most shrub cover classes were used i n proportion s i m i l a r to a v a i l a b i l i t y . The 1% class was used less, and the 20% class was used more. -40-Table 16. Availability and moose use of forest shrub cover classes in the West-Chilcotin during winter 1988. Shrub Availability Use Chi s.c.i .- Selection cover (%) plots % sightings % square upper lower 1 92 63.9 29 37.7 22.86* 0. 129 0.396 -3 12 8.3 6 7.8 -0. .070 0.080 0 5 13 9.0 14 18.2 -0. ,190 0.007 0 7 9 6.3 4 5.2 -0. ,053 0.074 0 10 6 4.2 8 10.4 -0. ,138 0.013 0 15 9 6.3 7 9.1 -0. ,104 0.047 0 20 1 0.7 6 7.8 -0. ,132 -0.010 + >25 2 1.4 3 3.9 -0. ,072 0.022 0 Total 144 100.0 77 100.1 * Significant at p. < 0.05. Refers to simultaneous comparison. Simultaneous confidence interval for difference between availability and use. 5.1.11 Wetland Shrub Cover A v a i l a b i l i t y Sixty two percent of wetlands have over 10% shrub cover. The >.2 5% cover class i s represented more often than any other class at 35.6% (Table 17). Twenty four percent of the s i t e s provided <1% shrub cover. Use The amount of use of a wetland appears t o c o r r e l a t e p o s i t i v e l y with the percent cover of shrubs i n th a t wetland. While no observations were recorded i n wetlands with <5% shrub cover, 45.3% of wetland observations occurred i n wetlands with >25% shrub cover. 41-Selection The s e l e c t i o n r e s u l t s indicate that wetlands with <5% shrub cover were used less than they were available, whereas most other s h r u b c o v e r c l a s s e s were us e d i n p r o p o r t i o n t o t h e i r a v a i l a b i l i t y . Use of the 20% class exceeded a v a i l a b i l i t y . Table 17. Availability and moose use of wetland shrub cover classes in the West-Chilcotin during winter 1988. Shrub cover (%) Availability Use Chi square S .C. .I. 1 Selection plots % sightings % upper lower 1 25 24.0 0 0, .0 37.28 0. .158 0. ,323 -3 5 4.8 0 0, .0 0. .007 0. ,089 -5 7 6.7 2 2. 3 -0. 014 0. ,102 0 7 3 2.9 3 3, .4 -0. 056 0. ,044 0 10 8 7.7 9 10, ,5 -0. 110 0. ,055 0 15 9 8.7 9 10, ,5 -0. 102 0. ,066 0 20 10 9.6 24 27. ,9 -0. 293 • -0. 072 + >25 37 35.6 39 45. ,3 -0. 237 0. ,042 0 Total 104 100.0 86 99. ,9 * Significant at p_ < 0.05. Refers to simultaneous comparison. Simultaneous confidence interval for difference between availability and use. 5.1.12 Wetland Shrub Height A v a i l a b i l i t y T a l l shrubs are not abundant i n the study area. 90.4% of wetland shrubs are under 2 m (Table 18). Use Use of wetlands with low shrubs was high, e s p e c i a l l y wetlands with shrubs i n the 0.6-1.5 m range. These wetlands accounted for 63.9% of wetland observations. In wetlands with t a l l e r shrubs, -42-use declined as shrub height increased. Selection Wetlands with shrubs i n the 0.6-1.0 m c l a s s were selected, while wetlands with shrubs <0.5 m were avoided. Wetlands with s h r u b h e i g h t c l a s s e s >l m were u s e d i n p r o p o r t i o n t o a v a i l a b i l i t y . Table 18. Availability and moose use of wetland shrub height classes in the West-Chilcotin during winter 1988. Height (meters) Availability Use Chi square s.c. i . - Selection pi ots % si ghti ngs % upper lower 0.0-0.5 39 41.5 14 16. ,3 14.65 0. .134 0.370 -0.6-1.0 19 20.2 29 33. .7 -0. .256 -0.015 + 1.1-1.5 20 21.3 26 30. ,2 -0. ,209 0.030 0 1.6-2.0 7 7.4 9 10. .5 -0. .108 0.048 0 2.1-3.0 8 8.5 7 8. ,1 -0. ,072 0.079 0 >3.0 1 1.1 1 1. 2 -0. ,030 0.028 0 Total 94 100.0 86 100. .0 Significant at p_ < 0.05. Refers to simultaneous comparison. Simultaneous confidence interval for difference between availability and use. -43-5.2 MONTHLY ANALYSES OF USE 5.2.1 Cover Type Classes Forest Versus Wetland Use of f o r e s t s i n January of both y e a r s was s i m i l a r , accounting f o r two-thirds of a l l observations (Table 19). Use of wetlands i n b o t h y e a r s was a l s o s i m i l a r , a c c o u n t i n g f o r approximately one-third of a l l January observations. Moose used forests more and wetlands less i n February of 1987 than i n February 1988 (Table 20). Use of f o r e s t s i n March of both y e a r s was t h e same, accounting for s l i g h t l y less than 50% of observations (Table 21). Use of wetlands was also equal between the two years, accounting f o r s l i g h t l y greater than 50% of observations In both y e a r s , use of wetlands i n c r e a s e d as the w i n t e r progressed. In January, wetlands accounted for an average of 35% of o b s e r v a t i o n s , while i n March, wetlands accounted f o r an average of 56% of observations. -44-Table 19. Moose use of cover types i n the West-Chilcotin during January 1987 and 1988. Class # of observations % of observations 1987 1988 1987 1988 Forest Fine immature 2 1 4.3 2.8 intermediate 15 6 31.9 16.7 mature 6 10 12.8 27.8 over mature 0 0 0.0 0.0 Fine Total 23 17 49.0 47.3 Spruce immature 0 3 0.0 8.3 intermediate 0 0 0.0 0.0 mature 3 3 6.4 8.3 Spruce Total 3 6 6.4 16.6 Pine/spruce 4 0 8.5 0.0 Aspen 1 0 2.1 0.0 Forest Total 31 23 66.0 63.9 Wetland grass 0 0.0 sedge 0 0.0 spruce 6 16.7 low b i r c h 1 2.8 low willow 2 5.6 low shrub 4 11.1 t a l l willow 0 0.0 hay 0 0.0 sedge/shrub 0 0.0 Wetland Total 16 13 34.0 36.2 Total 47 36 100.0 100.2 1 Data was not coll e c t e d for s p e c i f i c wetland types i n 1987. -45-Table 20. Moose use of. cover types i n the West-Chilcotin during February 1987 and 1988. Class # of observations 1987 1988 % of monthly observations 1987 1988 Forest Pine immature 0 intermediate 11 mature 7 over mature 0 Pine Total 18 3 3 11 3 20 0.0 22.0 14.0 0.0 36.0 3.6 3.6 13.1 3.6 23.9 Spruce immature 0 intermediate 0 mature 2 Spruce Total 2 Pine/spruce 7 Aspen 1 Forest Total 28 Wetland Grass sedge spruce low b i r c h low willow low shrub t a l l willow hay sedge/shrub Wetland Total 22 Tot a l 50 2 1 14 17 3 0 40 0 0 24 3 2 11 2 0 2 44 84 0.0 0.0 4.0 4.0 14.0 2.0 56.0 44.0 100.0 2.4 1.2 16.7 20.3 3.6 0.0 47.8 0.0 0.0 28.6 3.6 2.4 13.1 2.4 0.0 2.4 52.5 100.3 Data was not coll e c t e d for s p e c i f i c wetland types i n 1987. -46-Table 21. Moose use. of cover types i n the West-Chilcotin during March 1987 and 1988. Class # of observations % of monthly observations 1987 1988 1987 1988 Forest Pine immature 2 0 5.4 0.0 intermediate 9 1 24.3 1.4 mature 3 3 8.1 4.1 over mature 0 3 0.0 4.1 Pine Total 14 7 37.8 9.6 Spruce immature 0 4 0.0 5.4 intermediate 0 0 0.0 0.0 mature 1 19 2.7 25.7 Spruce Total 1 23 2.7 31.1 Pine/spruce 1 1 2.7 1.4 Aspen 0 2 0.0 2.7 Forest Total 16 33 43.2 44.8 Wetland grass 0 0.0 sedge 3 4.1 spruce 14 18.9 low b i r c h 3 4.1 low willow 4 5.4 low shrub 8 10.8 t a l l willow 9 12.2 hay 0 0.0 sedge/shrub 0 0.0 Wetland Total 21 41 56.8 55.5 Total 37 74 100.0 100.3 Data was not co l l e c t e d for s p e c i f i c wetland types i n 1987. -47-Forest Although i t would be desirable to compare use of s p e c i f i c f o r e s t c l a s s e s between y e a r s , t h i s c a n n o t be done w i t h c o n f i d e n c e . Assessment of c l a s s e s used i n 1987 was done by a e r i a l o b s e r v a t i o n w i t h o u t s u b s e q u e n t g r o u n d v i s i t s . M i s c l a s s i f i c a t i o n between intermediate and mature c l a s s e s , was highly possible. Additionally, i n 1987 the overmature lodgepole pine c l a s s was not considered as a separate c l a s s from mature lodgepole pine c l a s s . For these reasons, the data w i l l be compared using 2 classes only, immature and non-immature. Use of p i n e accounted f o r n e a r l y o n e - h a l f o f J a n u a r y observations i n both 1987 and 1988 (Table 19). In f a c t , use of non-immature stands accounted for 43% of a l l observations i n both years. Immature stands accounted for few observations. Use of spruce stands d i f f e r e d between y e a r s , p r i m a r i l y due to an increase i n use of immature stands i n 1988. Use of mature spruce stands was s i m i l a r i n both years. Use of the pine/spruce cover type was greater i n 1987 than i n 1988. Use of aspen was low i n 1987 and nonexistent i n 1988. Use of non-immature stands i n February accounted f o r 36.0% and 20.3% of observations i n 1987 and 1988, r e s p e c t i v e l y (Table 2 0) . No moose were observed i n immature pine i n 1987 and only 3.6% of February 1988 observations were i n t h i s c l a s s . Use of spruce i n 1988 was 16.3 percentage points greater than i n 1987, with the mature c l a s s accounting f o r most of t h i s d i f f e r e n c e . Pine/spruce was used 14% of the time i n 1987, but only 3.6% of -48-the time 1988. Aspen was r a r e l y used, accounting f o r only 2.0% of 1987 observations and none of 1988 observations. While March use of f o r e s t s remained r e l a t i v e l y constant at approximately 44% i n both years, there was a s h i f t from pine to spruce from 1987 to 1988 (Table 21) . Use of non-immature pine stands equalled 32.4% i n 1987, but only 9.6% i n 1988. Immature pine accounted f o r 5.4% of 1987 observations, but none of 1988 observations. Use of mature spruce equalled 25.7% i n 1988, but only 2.7% i n 1987. Pine/spruce and aspen were r a r e l y used i n e i t h e r 1987 or 1988. In 1987, there was no apparent monthly trend i n use of pine and spruce. There was a 13 percentage point d e c l i n e i n use of pine between January and February, but l i t t l e change between February and March. There was a s l i g h t decline (3.7 percentage p o i n t s ) i n use of spruce throughout the winter. During 1987, there was a gradual s h i f t from forest to wetland cover types as winter progressed. In 1988, use of pine dropped from 47.3% to 9.6% as the winter progressed. Use of spruce, however, increased from 16.6% to 31.1% between January to March. Like 1987, i n 1988 moose also s h i f t e d from f o r e s t to wetland cover types as the w i n t e r progressed. -49-Wetlands Since a e r i a l observations were not mapped accurately i n 1987, i t was impossible to v i s i t l o c a t i o n s to determine c l a s s e s of wetlands used. Detailed information i s a v a i l a b l e only f o r the 1988 season. Spruce wetlands were used more often than any other wetland c l a s s i n a l l 3 months ( t a b l e s 19, 20, 21) . Use of spruce wetlands increased from 16.7% i n January to 28.6% i n February, and then d e c l i n e d to 18.9% i n March. The low shrub c l a s s received the second highest use of a l l the classes i n January and March and remained si m i l a r (10.8-13.1%) throughout the winter. Use of the t a l l willow class increased from 0% i n January to 2.4% i n February to 12.2% i n March. Use of low b i r c h and low willow c l a s s e s remained similar throughout the winter. Because of the s t r u c t u r a l s i m i l a r i t i e s of the low b i r c h , low willow, and low shrub classes, there i s j u s t i f i c a t i o n for lumping these three classes together. Use of these combined classes remained near 20% d u r i n g a l l 3 months. Use of g r a s s , sedge, hay and sedge/shrub was n e g l i g i b l e throughout the w i n t e r w i t h the exception of sedge which increased from no use i n January and February to 4.1% i n March. 5.2.2 Distance-to-Wetland In both years, use i n a l l months was concentrated near the wetland edge. Approximately one-half of the observations i n each -50-month o c c u r r e d i n the 0-20 m c l a s s , w h i l e over 78% of the observations i n each month occurred within 100 m of a wetland edge. Table 22. Monthly moose use of distance-to-wetland classes in the West-Chilcotin during winter 1987 and 1988. Distance to # of observations % of monthly observations wtlnd (m) Jan Feb Mar Jan Feb Mar 1987 1988 1987 1988 1987 1988 1987 1988 1987 1988 1987 1988 0-20 9 11 13 19 17 21 56. .3 40. .7 46. ,4 47. ,5 56. .7 63. .6 21-40 0 6 2 7 3 5 0. 0 22. .2 7. .1 17. .5 10. .0 15. ,2 41-60 4 4 0 3 0 1 25. .0 14. .8 0. .0 7, .5 0. .0 3. .0 61-80 0 1 7 4 6 3 0. .0 3. .7 25. .0 10, .0 20. .0 9. .1 81-100 3 0 0 2 0 0 18. .8 0. .0 0. .0 5, .0 0. .0 0. .0 100-149 0 0 0 2 0 0 0. .0 0. .0 0. .0 5, .0 0. .0 0. .0 150-200 0 5 1 3 0 1 0. 0 18. ,5 3. .6 7. ,5 0. .0 3, .0 >200 0 0 5 0 4 2 0. 0 0. ,0 17. .9 0. ,0 13. .3 6. .1 Total 16 27 28 40 30 33 100. 1 99. 9 100. 0 100. ,0 100. 0 100. .0 5.2.3 Distance-to-Forest S i m i l a r to use of d i s t a n c e - t o - w e t l a n d c l a s s e s , use of distance-to-forest classes was highest near the edge. Use of the 0-20 m c l a s s ranged from 57.1% to 93.8% during the 1987 and 1988 winter seasons. Use of wetland classes beyond 100 m was <5% i n a l l months. A g a i n , as w i n t e r p r o g r e s s e d , the number of o b s e r v a t i o n s i n each c l a s s d i d not change i n any n o t i c e a b l e pattern. -51-Table 23. Monthly moose use of d i s t a n c e - t o - f o r e s t c l a s s e s i n the W e s t - C h i l c o t i n d u r i n g w i n t e r 1987 and 1988. Distance t o # of obse r v a t i o n s  f o r e s t (m) Jan Feb Mar 1987 1988 1987 1988 1987 1988 0-20 6 8 15 28 15 25 66. .7 57. .1 88. .2 54. .9 93, ,8 59. .5 21-40 3 4 0 6 0 7 33. ,3 28. .6 0. ,0 11. .8 0, .0 16. .7 41-60 0 0 0 9 0 6 0, .0 0. .0 0. .0 17 .6 0, .0 14. .3 61-80 0 2 2 4 1 1 0. .0 14. .3 11. ,8 7 .8 6, .2 2 .4 81-100 0 0 0 1 0 2 0. .0 0. .0 0. .0 2 .0 0, .0 4 .8 100-149 0 0 0 0 0 1 0. ,0 0. 0 0. 0 0, .0 0. ,0 2. ,4 150-200 0 0 0 2 0 0 0. ,0 0. 0 0. 0 3, .9 0. .0 0, .0 >200 0 0 0 1 0 0 0. ,0 0. .0 0. .0 2. .0 0. .0 0. .0 Total 9 14 17 51 16 42 100. ,0 100. 0 100. ,0 100 .0 100. .0 100. .1 5.2.4 Forest Age As the winter progressed the percentage of observations i n younger forests decreased (Table 24). For example, i n January 71.5% of observations occurred i n forests younger than 60 years, but i n February and March the percentage dropped to <28%. Use of i n d i v i d u a l classes was highly variable between months. % of monthly o b s e r v a t i o n s  Jan Feb Mar 1987 1988 1987 1988 1987 1988 -52-Table 24. Monthly moose use of f o r e s t age c l a s s e s i n the W e s t - C h i l c o t i n d u r i n g w i n t e r 1988. Age (years) Jan Feb Mar # % # % # % 0-20 2 14.3 1 3 4 3 10 0 21-40 6 42.9 4 13 8 2 6 7 41-60 2 14.3 3 10 3 2 6 7 61-80 1 7.1 7 24 1 6 20 0 81-100 0 0.0 7 24 1 5 16 7 101-120 1 7.1 2 6 9 4 13 3 121-140 2 14.3 1 3 4 4 13 3 >141 0 0.0 4 13 8 4 13 3 To t a l 14 100.0 29 99 8 30 100 0 5.2.5 Forest Height Use of fo r e s t height classes did not follow any obvious trend (Table 25) . i n a l l 3 months fo r e s t s <2 m t a l l were not used. Use of other classes ranged from 3.3% to 28.6%. Table 25. Monthly moose use of f o r e s t height c l a s s e s i n the W e s t - C h i l c o t i n d u r i n g w i n t e r 1988. Height Jan Feb Mar (m) # % # % # % 0.0-2.0 0 0 0 0 0 0 0 0 0 2.1-5.0 3 21 4 3 10 0 3 9 3 5.1-10.0 4 28 6 6 20 0 4 12 5 10.1-12.0 3 21 4 6 20 0 7 21 9 12.1-14.0 0 0 0 5 16 7 5 15 6 14.1-16.0 2 14 3 6 20 0 7 21 9 16.1-18.0 2 14 3 1 3 3 2 6 3 >18.0 0 0 0 3 10 0 4 12 6 To t a l 14 100 0 30 100 0 30 100 1 -53-5.2.6 Forest Density In a l l 3 months forests of most f r e q u e n t l y (Table 26). between 9 and 19 trees per 100 of f o r e s t observations. intermediate density were used the Forests with d e n s i t i e s ranging sq. m. received between 43 and 74% Table 26. Monthly moose use of f o r e s t d e n s i t y c l a s s e s i n the W e s t - C h i l c o t i n d u r i n g w i n t e r 1988. De n s i t y (stems/ 100 m ) Jan Feb Mar # 0 1 # 0 i # 0 i 0-4 1 7 .1 2 6, ,5 1 3. .1 5-8 1 7 .1 5 16, .1 1 3. .1 9-11 0 0 .0 10 32, .3 12 37 .5 12-14 2 14 .3 5 16, .1 8 25. .0 15-19 4 28 .6 5 16, .1 4 .4 20-24 2 14. .3 0 0. ,0 3 9, .4 25-29 1 7 .1 1 3, ,2 2 6, .3 >30 3 21. .4 3 9, ,7 1 3, .1 Total 14 99. .9 31 100. .0 32 99, .9 5.2.7 Forest Canopy Closure The percentage of observations i n forest with canopy closure >10% decreased as the winter progressed (Table 27). In January, 71.4% of f o r e s t observations were i n forests with canopy closure values >10%. F i f t y percent of moose observations i n January were i n the 2 0% c l a s s . In February and March, use of f o r e s t s with c a n o p y c l o s u r e v a l u e s >10% d r o p p e d t o 48.4 and 47.0% r e s p e c t i v e l y . -54-Table 27. Monthly moose use of f o r e s t canopy c l o s u r e c l a s s e s i n the W e s t - C h i l c o t i n d u r i n g w i n t e r 1988. Canopy Jan Feb Mar c l o s u r e (%) # % # % # % <1 1 7, .1 0 0. .0 2 6, .3 3 1 7. ,1 4 12. .9 1 3, .1 5 2 14. .3 10 32. .3 7 21, ,9 7 0 0. ,0 2 6. .5 7 21. .9 10 1 7. .1 3 9, ,7 7 21, .9 15 2 14, .3 4 12. ,9 3 9, .4 20 7 50. .0 3 9. ,7 2 6, ,3 >25 0 0, .0 5 16. .1 3 9, .4 Total 14 99 .9 31 100. ,1 32 100, .2 5.2.8 Forest DBH As the winter progressed, the number of observations i n f o r e s t s with tree diameters-at-breast height >17 cm increased ( T a b l e 2 8 ) . F o r example, i n J a n u a r y 50% o f t h e moose observations were i n stands with trees >17 cm DBH. In February and March, t h i s p e r c e n t a g e i n c r e a s e d t o 69.9 and 83.9% re s p e c t i v e l y . 5.2.9 Forest Shrub Cover In a l l 3 months, forests with shrub cover <5% were used the most extensively (Table 29) . Use of f o r e s t s with shrub cover >10% increased from 21.3% i n January to 32.2% i n February and to 34.4% i n March. -55-Table 28. Monthly moose use of forest DBH classes in the West-Chilcotin during winter 1988. DBH Jan Feb Mar (cm) # % # % # % 0-5 2 14. .3 1 3. 3 2 6. .5 5-7 1 7. ,1 1 3. 3 0 0. 0 7-10 3 21 .4 2 6, .7 2 6. 5 10-12 0 0. 0 1 3 .3 0 0. ,0 13-16 1 7. 1 4 13, .3 1 3. .2 17-20 3 21. .4 10 33, .3 7 22. ,6 21-24 2 14. .3 4 13, .3 8 25. 8 >25 2 14 .3 7 23 .3 11 35. ,5 Total 14 99, .9 30 99 .8 31 100. ,1 Table 29. Monthly moose use of forest shrub cover classes in the West-Chilcotin during winter 1988. Shrub cover Jan Feb Mar (%) # % # % # % <1 7 50. ,0 12 38, ,7 10 31. .3 3 0 0. .0 3 9, .7 3 9. .4 5 4 28, ,6 2 6, .5 8 25, ,0 7 0 0, ,0 4 12, .9 0 0. .0 10 1 7. ,1 1 3, ,2 6 18. ,8 15 1 7. ,1 4 12, .9 2 6. .3 20 1 7. 1 4 12, .9 1 3. ,1 >25 0 0, ,0 1 3, .2 2 6. .2 Total 14 99. ,9 31 100. ,0 32 100. ,1 - 5 6 -5.2.10 Wetland shrub Cover Use of wetland shrub cover classes was consistent throughout the winter (Table 30). In a l l three months use was concentrated i n wetlands with shrub cover values >15%. No use was recorded i n wetlands with shrub cover values <5%. Table 30. Monthly moose use of wetland shrub cover c l a s s e s i n the W e s t - C h i l c o t i n d u r i n g w i n t e r 1988. Shrub cover Jan Feb Mar (%) # % # % # % <1 0 0. .0 0 0, .0 0 0. ,0 3 0 0. .0 0 0. ,0 0 0. .0 5 0 0. .0 2 4. .9 0 0. .0 7 0 0. .0 2 4, ,9 1 2. .8 10 0 0. .0 6 14. ,6 3 8. 3 15 1 11. .1 6 14. .6 2 5. .6 20 4 44. .4 11 26. ,8 9 25. .0 >25 4 44. 4 14 34. .0 21 58. 3 Total 9 99. .9 41 99. ,8 36 100. 0 5.2.11 Wetland Shrub Height In a l l three months, use was concentrated i n wetlands with shrub heights between 0.6 and 1.5 m (Table 31). In each month, over 61% of wetland observations were i n wetlands with shrubs of t h i s height. -57-Table 31. Monthly moose use of wetland shrub height c l a s s e s i n the W e s t - C h i l c o t i n d u r i n g w i n t e r 1988. Shrub height Jan Feb Mar ( m ) # % i % # % 0.0-0.5 0 0 .0 11 26. .8 3 8 .3 0.6-1.0 5 55, .5 13 31. ,7 11 30, .6 1.1-1.5 2 22 .2 13 31, ,7 11 30, .6 1.6-2.0 2 22 .2 2 4, .9 5 13 .9 2.1-3.0 0 0, .0 2 4, ,9 5 13, .9 >3.0 0 0, .0 0 0. ,0 1 2, .7 Total 9 99, .9 41 100, ,0 36 100, .0 5.3 WINTER HOME RANGE SIZE The winter home ranges of r a d i o - c o l l a r e d moose v a r i e d from 2.7 to 45.2 km2 i n 1987 and from 3.6 to 155.5 km2 i n 1988 (Table 32). Average winter home range s i z e equalled 20.7 km i n 1987 2 and 45.0 km i n 1988. The l a r g e ranges of moose A and B c o n t r i b u t e s i g n i f i c a n t l y t o the l a r g e r a v e r a g e i n 1988. 2 Excluding these home ranges, the 1988 average s i z e i s 23.4 km . The home ranges of moose A and B are large because they include 6 observations before these moose a r r i v e d i n the wintering area. These moose were one month l a t e r i n returning to the wintering area than the previous year and were l a t e r than a l l the other c o l l a r e d moose to r e t u r n , not a r r i v i n g u n t i l Feb. 1 and 11 resp e c t i v e l y . -58-Table 32. Winter home range s i z e s of r a d i o - c o l l a r e d moose i n the W e s t - C h i l c o t i n . Moose 1987 1988 S i z e # of obs. S i z e # of obs. (km ) (km ) A 19.6 16 155.5 27 B 41.3 15 63.9 27 C 45.2 9 17.2 30 D 13.5 16 22.7 29 E 6.2 15 3.6 30 F 2.7 16 16.2 29 G 12.2 10 I 42.9 20 J 37.6 23 Average 20.7 45.2 (23.4) Average 1988 home-range s i z e e x c l u d i n g moose A and B. 5.4 WINTER DIET 5.4.1 Composition The winter d i e t of moose i n both years consisted mainly of shrubs (Table 33), which contributed >60% of the d i e t i n each month. C o n i f e r s were the second most consumed forage group, comprising 10-30% of the d i e t . Bog b i r c h was the s i n g l e most consumed species, making up 28-35% of the d i e t . Lodgepole pine was the second most consumed s p e c i e s , comprising 10-30% of monthly d i e t s . Two other plants that contributed s i g n i f i c a n t l y to winter d i e t were willow (7-19%) and Service berry (Amelanchier  a l n i f o l i a ) (5-13%). In February and March of 1988, sedges were a s i g n i f i c a n t component of the d i e t making up 22 and 14% of the t o t a l r espectively. -59-5.4.2 Monthly Analysis In 1987, the d i e t composition remained r e l a t i v e l y constant throughout the winter. Bog bi r c h and lodgepole pine were the two most often consumed species. In February and March of 1988, there appears to be a switch from pine to sedge. While pine made up 22.6% of the January d i e t , i t decreased to 11.8 and 10.3% of February and March d i e t s r e s p e c t i v e l y . During the same time period, sedges increased from 6.6% to 21.7% and then decreased to 14.4 %. Table 33. Winter d i e t composition o£ r a d i o - c o l l a r e d moose i n the W e s t - C h i l c o t i n d u r i n g 1987 and 1988. Ve g e t a t i o n Jan Feb Mar Type 1987 1988 1987 1988 1987 1988 % % % % % % C o n i f e r s Lodgepole pine 21 .0 22 .6 30 .5 11 .8 26 .5 10 .3 Other c o n i f e r 0 .4 0 .0 1 .2 0 .0 0 .1 0 .0 C o n i f e r T o t a l 21 .4 22 .6 31. .7 11 .8 26 .6 10 .3 Grasses 2 .7 3. .3 0. .4 5. .6 3. .4 2. .9 Sedges/Rushes 5. .6 6. .6 0, ,0 21, .7 3. .4 14. .4 Ferns 0. .1 0, .0 0. ,0 0. .0 0. .7 0, .0 Mosses 0, .5 0. .0 0. .0 0, ,0 0, .0 0. .0 Forbs - 1. .7 0. ,0 0. ,3 0. .0 0. .7 1. ,8 Lichen 0. .0 1. .3 0. ,0 0. .0 0. .5 0. .0 Shrubs Bog b i r c h 34. ,2 28. .1 29. .3 32. .8 31. .3 34. .5 Wi11ow 13. .2 9. .8 9. .7 7. .4 18. .9 11. .3 S e r v i c e b e r r y 5. .9 8. 9 13. 0 8. 5 0. .0 5. 2 Other shrubs 14. ,7 19. 4 15. 6 12. .2 14. .5 19. .6 Shrub Total 68. .0 66. 2 67. 6 60. 9 64. .7 70. .6 Determined by f e c a l fragment analyses. -60-5.4.3 Quality Crude p r o t e i n values of winter browse ranged from 4.0% to 7.8% (Table 34). Percent crude p r o t e i n values between January and March were highest i n bog b i r c h ranging from 7.2% to 7.8%. Lodgepole pine also had high values, ranging from 6.5% to 7.9%. Sedges had the lowest crude protein l e v e l s , ranging from 4.0% to 5.3% Gross energy l e v e l s ranged from 4188 calories/gram to 5872 c a l o r i e s / g r a m . Bog b i r c h and lodgepole pine a g a i n had the highest v a l u e s , ranging from 5580 to 5872 and 5444 to 5655 calories/gram respectively. Sedges had the lowest energy content of a l l measured forages, ranging from 4188 to 4222 calories/gram. Neutral detergent f i b e r (NDF) leve l s ranged from 47.6-79.3%. Sedges had the highest percentage NDF, ranging from 77.2-79.3%. Levels were second highest i n bog birch, ranging from 52.7-58.1%. -61-Table 34. Monthly n u t r i t i o n a l parameters of major forage sp e c i e s consumed by w i n t e r i n g moose i n the W e s t - C h i l c o t i n , 1988. % Crude P r o t e i n Gross Energy cal/gm % Neutral Detergent F i b e r Lodgepole pine January February March A p r i 1 6.5 7.0 7.9 6.8 Trembli ng Aspen January February 5.5 6.6 Wi 1 low January February March A p r i l 6.2 6.2 7.1 6.9 Sedges March A p r i 1 5.3 4.0 Bog B i r c h January February March A p r i 1 7.2 7.8 7.2 5.2 5567 5498 5655 5444 54.4 51.5 47.6 51.1 5247 5327 55.0 50.9 5390 5321 5427 5309 56.9 52.4 47.2 46.5 4222 4188 79.3 77.2 5580 5799 5865 5872 58.1 56.4 52.7 57.4 5.5 Snow Depth Snow depth i n 1987 was greater than i n 1988 (Table 35). For example, i n early January the 1987 and 1988 snow depths were 49 and 11 cm r e s p e c t i v e l y . The mean snow depths f o r January, February and March 1987 were 48, 48, and 42 cm respe c t i v e l y . The mean monthly values for 1988 for the same period were 21, 35, and 24 cm respectively. -62-Table 35. Winter 1987 and 1988 d a i l y snow depths (cm) recorded at 5 M i l e Ranch i n the W e s t - C h i l c o t i n . Day Dec/86 Dec/87 Jan/87 Jan/88 Feb/87 Feb/88 Mar/87 Mar/88 Apr/87 Apr/88 1 36 0 49 11 48 37 47 33 30 13 2 36 0 49 11 48 42 47 33 28 13 3 36 3 49 11 48 36 47 33 28 12 4 36 5 49 11 48 38 47 30 28 12 5 36 6 49 11 48 38 47 30 27 11 6 36 6 49 11 50 36 48 31 27 11 7 39 6 49 11 52 36 48 31 27 9 8 39 6 49 11 49 35 47 28 26 9 9 39 6 48 11 49 35 47 28 23 9 10 39 5 48 13 49 38 47 27 21 8 11 39 5 48 17 49 35 48 27 21 8 12 47 5 48 17 48 35 47 26 19 6 13 50 5 48 17 48 35 46 26 17 6 14 50 5 48 17 48 35 46 26 13 4 15 49 5 47 19 48 37 45 26 13 2 16 49 6 47 19 48 37 44 26 11 2 17 48 6 47 19 47 37 44 23 10 0 18 48 6 47 19 47 36 44 22 8 0 19 46 6 47 19 47 36 43 21 8 0 20 46 9 47 25 47 36 42 21 5 0 21 46 9 47 29 47 35 42 21 4 0 22 47 9 47 32 47 35 42 21 2 0 23 47 9 47 32 48 34 41 19 0 0 24 47 9 47 31 48 33 37 19 0 0 25 47 9 47 31 48 33 36 18 0 0 26 49 9 47 31 47 32 36 18 0 0 27 52 9 47 31 47 32 33 16 0 0 28 51 9 47 31 47 32 33 16 0 0 29 51 11 47 32 32 31 16 0 0 30 51 11 47 37 31 16 0 0 31 49 11 48 37 30 16 Average 45 7 48 21 48 35 42 24 13 5 Range 36-52 0-11 47-49 11-37 47-52 32-42 30-48 16-33 0-30 o-i; Atmospheric Environment S e r v i c e (1986, 1987, 1988). - 6 3 -6. DISCUSSION 6.1 SELECTION INDICES Selection indices involve many assumptions and have several c h a r a c t e r i s t i c s that are not immediately obvious; t h e r e f o r e , t h e i r r e s u l t s can be e a s i l y m i s i n t e r p r e t e d and management strat e g i e s derived from these misinterpretations can have severe consequences. This discussion b r i e f l y o u t l i n e s the concept of s e l e c t i o n and then addresses important aspects of h a b i t a t s e l e c t i o n indices that are not r e a d i l y perceived, but need to be understood i f the index i s going to be i n t e r p r e t e d c o r r e c t l y . S p e c i f i c a l l y , the following w i l l be discussed: 1. Errors i n interpreting habitat s e l e c t i o n indices 2. Assumptions underlying habitat s e l e c t i o n indices 3. The character of habitat selection indices i n r e l a t i o n to habitat a v a i l a b i l i t y . 6.1.1 Selection Hess and Swartz (1940) were among the f i r s t to develop the ideas of s e l e c t i o n using food preference r a t i n g s . A preferred food species i s one which i s proportionately more frequent i n the d i e t of an animal than i n the available environment. From t h i s idea, s e l e c t i o n theory has expanded to d i f f e r e n t l e v e l s , such as s e l e c t i o n f o r p h y s i c a l or geographical range by a p a r t i c u l a r species, s e l e c t i o n of a home range by an i n d i v i d u a l or s o c i a l group, and s e l e c t i o n of habitat components within the home range of an i n d i v i d u a l (Johnson 1980). The concept i n v o l v e d with a s e l e c t i o n index (SI) i s that i n the absence of s e l e c t i o n , use (u) -64-of a v a i l a b l e components (a) w i l l be random. That i s , i f an animal does not purposefully select the components i t uses, the use of a component w i l l be proportional to the a v a i l a b i l i t y of that component. This i s quantified i n the following r a t i o : SI = u/a In theory, a selection index of 1 ( i . e . use = a v a i l a b i l i t y ) indicates that the component i s neither selected nor avoided, but i s used i n proportion to i t s a v a i l a b i l i t y . Values >1 i n d i c a t e s e l e c t i o n , values <1 indicate avoidance. Often, r e l a t i v e degrees of s e l e c t i o n are defined. For example, Ault and Stormer (1983) rated s e l e c t i v i t y as high (SI >1.5), medium (0.75< SI <1.50), and low (SI <0.75). P i t t (pers. comm.), i n a bighorn sheep study, a r b i t r a r i l y defined selection for a p a r t i c u l a r plant species as SI >2.5 and avoidance as SI <0.7. The assumption of a s e l e c t i o n index i s that i t allows the r e l a t i v e p r e f e r e n c e f o r a component to be i d e n t i f i e d . For example, i f three plant species comprise equal percentages of an animal's d i e t , but one of these equally-consumed species i s less a v a i l a b l e than the others, then the animal feeding on i t must have sought i t out. The animal, then, may be said to prefer t h i s species over the other two (Petrides 1975). The s e l e c t i o n index p r o v i d e s a r a t i o between h a b i t a t a v a i l a b i l i t y and h a b i t a t use, but i t does not i n c o r p o r a t e a s t a t i s t i c a l t e s t . Neu et. a l . (1974) provided a s t a t i s t i c a l t e c h n i q u e f o r e v a l u a t i n g p r e f e r e n c e or avoidance u s i n g the Bonferroni z s t a t i s t i c i n conjunction with the Chi-square. Byers -65-and S t e i n h o r s t (1984) demonstrated how the s i m u l t a n e o u s confidence l i m i t s for the Bonferroni approach can be calculated. Johnson (1980) recognized that preference of components depended, to a large degree, on the array of components that the researcher deemed a v a i l a b l e . He proposed a method based on ranks of components by usage and by a v a i l a b i l i t y that provides comparable r e s u l t s whether a questionable component i s included or excluded from consideration. Chesson (1983) developed a method where the measure of p r e f e r e n c e does not change with changes i n food d e n s i t i e s unless consumer behaviour also changes. A l l r e d g e and R a t t i (1986) d i s c u s s e d two other s t a t i s t i c a l techniques f o r analysis of resource selection, the Friedman t e s t and the Quade t e s t . Despite the s o p h i s t i c a t i o n of these various techniques, they are a l l d e r i v e d from the b a s i c s e l e c t i o n i n d e x and i n c o r p o r a t e some of the l o g i c f a u l t s a ssociated with i t . An understanding of these f a u l t s i s important before management decisions based on selection indices are made. 6.1.2 Errors i n Interpretation The Selection Index i s not a Direct Measure of Importance When assessing u s e - a v a i l a b i l i t y i n d i c e s , land managers may misconceive what the s e l e c t i o n index measures. For example, a s e l e c t e d habitat i s often misinterpreted to mean an important habitat. This misinterpretation may be c a r r i e d further such that habitats with large selection index values are considered to be more important than habitats with small s e l e c t i o n values. I t i s i m p o r t a n t t o r e a l i z e t h a t the u n i t s f o r s e l e c t i o n a r e u s e / a v a i l a b i l i t y . The index alone does not provide a d i r e c t -66-measure of habitat importance; importance can only be infe r r e d from these kind of data. An example of t h i s kind of m i s i n t e r p r e t a t i o n e x i s t s i n a recently published paper i n the Journal of W i l d l i f e Management. Fa i r b a n k s e t . a l . (1987) found that open grasslands, i n a l l seasons, were used by Rocky Mountain b i g h o r n sheep (Ovis c a n a d e n s i s canadensis) to a l e s s e r degree than they were av a i l a b l e . They state that grasslands are avoided i n a l l seasons and imply that grasslands are not important, yet grasslands were used more than any other component (41-63%). Researchers often i n t e r p r e t s e l e c t i o n to mean preference. C l e a r l y , s e l e c t i o n i s a f u n c t i o n of many f a c t o r s o ther than simple p r e f e r e n c e . S e l e c t i o n i n v o l v e s a s e r i e s of f a c t o r s i n c l u d i n g i n t e r n a l a n i m a l f a c t o r s , l e a r n e d and e v o l v e d b e h a v i o u r s , and environmental i n f l u e n c e s (Heady 1964). A s e l e c t i o n index may imply preference but i s not a d i r e c t measure of preference. 6.1.3 Assumptions Selection Assumes That Animal Has Choice A ha b i t a t s e l e c t i o n index i s often used to t e s t the n u l l hypothesis that use of habitats i s random. If use i s not random, the assumption i s then made that the animal consciously s e l e c t s the h a b i t a t s i t uses. However, t h i s l o g i c f a i l s to consider a t h i r d p o s s i b i l i t y . The animal may be forced to use the habitats i t does because i t does not have a choice. For example, i t may be f o r c e d to l i v e there because human encroachment through settlement or industry has rendered more desirable habitats -67-u n a v a i l a b l e . A l t e r n a t i v e l y , p r e f e r r e d h a b i t a t s may be made unavailable because of the presence of predators or because of density dependent factors. In these cases, the preferred habitat w i l l be u n d e r - u t i l i z e d and subsequently underestimated by the r e s e a r c h e r . The s e l e c t i o n index cannot d i s t i n g u i s h between conscious s e l e c t i o n and forced "no choice." 6.1.4 Selection and A v a i l a b i l i t y The s e l e c t i o n index value depends, to a large degree, on the a v a i l a b i l i t y component. Various problems are associated with the a v a i l a b i l i t y component and subsequently r e s u l t i n problems with the index. Defining A v a i l a b i l i t y i s Subjective While determining h a b i t a t use i s r e l a t i v e l y o b j e c t i v e , d e f i n i n g h a b i t a t a v a i l a b i l i t y u s u a l l y r e q u i r e s p e r s o n a l , subjective judgement. The researcher cannot know absolutely what i s a v a i l a b l e t o t h e a n i m a l and must, t h e r e f o r e , i n f e r a v a i l a b i l i t y from the animal's behaviour. Herein l i e s a major problem of s e l e c t i o n i n d i c e s . What the researcher d e f i n e s as a v a i l a b l e may g r e a t l y i n f l u e n c e the s e l e c t i o n index and, consequently, what i s defined as selected and avoided. What the r e s e a r c h e r deems a v a i l a b l e to the animal may be e n t i r e l y d i f f e r e n t from what the animal deems a v a i l a b l e to i t s e l f . The r e s u l t i n g s e l e c t i o n v a l u e s may be more a f u n c t i o n of the researcher's perception of the animal than of the biology of the animal. Johnson (1980) provided an hypothetical example of how the researcher's notion of which components are a v a i l a b l e can - 6 8 -i n f l u e n c e whether an i n d i v i d u a l component i s used above, i n p r o p o r t i o n t o , or below i t s a v a i l a b i l i t y . He d e s c r i b e d a s i t u a t i o n where two investigators have d i f f e r e n t opinions as to which food items are a v a i l a b l e to a f i s h . The d i f f e r e n c e i n t h e i r opinion i s r e f l e c t e d i n the conclusions regarding s e l e c t i o n and avoidance (Table 36). When item A i s considered ava i l a b l e , item B, i s defined as a preferred component. However, when item A i s considered unavailable, item B then becomes defined as an avoided component. C l e a r l y , the change i n "preference" i s not r e l a t e d to a change i n f i s h behaviour, but rather i s dictated by the i n v e s t i g a t o r ' s n o t i o n of the array of components deemed av a i l a b l e . Table 36. Example i l l u s t r a t i n g r e s u l t s of comparing usage and a v a i l a b i l i t y when a common but seldom-used item i s included (A) and when excluded (B) from consideration . Item Usage A v a i l a b i l i t y Conclusion (%) (%) A A 2 60 Avoided B 43 30 Preferred C 55 10 Preferred B B 44 75 Avoided C 56 25 Preferred Johnson (1980). -69-Selection Order The r e s e a r c h e r ' s d e f i n i t i o n of which components a r e a v a i l a b l e to the animal can influence s e l e c t i o n indices i n a less obvious manner. Often, i n v e s t i g a t o r s define components within the animal's home range as being available to the animal. Use of the various components within the home-range can then be compared to the a v a i l a b i l i t y of each component and the degree of s e l e c t i o n assessed. A major, but rather pervasive flaw with t h i s technique i s that i t f a i l s to consider that the animal may have selected i t s home range because of the abundance of a p a r t i c u l a r component. The magnitude of the "a" value f o r t h i s component w i l l l i k e l y prevent selection from being indicated even i f use i s high. The following example i l l u s t r a t e s t h i s i d e a . During the w i n t e r months, moose i n the W e s t - C h i l c o t i n Region of B.C. congregate i n an area containing abundant wetlands ( l e t ' s assume moose congregate there because of the abundance of wetlands) . T h i s area i s surrounded by hundreds of square k i l o m e t e r s of lodgepole pine f o r e s t . For sake of argument, l e t ' s say that wetlands make up 90% of the w i n t e r i n g area and 10% of the surrounding forested area. A u s e - a v a i l a b i l i t y study might f i n d that moose use wetlands 50% of the time and f o r e s t s 50% of the time. The conclusion, then, would be that wetlands are avoided (use i s less than a v a i l a b i l i t y ) and forests are selected, when i n f a c t , the opposite i s true. Selection for the winter range had been based on the abundance of wetlands, yet w i t h i n the winter range, i t appears as i f wetlands are avoided. -70-The above argument i l l u s t r a t e s that s e l e c t i o n can occur at d i f f e r e n t l e v e l s . Johnson (1980) defined 4 orders of se l e c t i o n . From lowest to highest these are: 1. F i r s t order selection - selection of physical or geographical range of a species 2. Second order selection - selection of the home range of an in d i v i d u a l within the physical or geographic range 3. Third order selection - selection of habitat components within a home range 4. Fourth order selection - selection of av a i l a b l e food items within a habitat component. Recognizing that lower orders of se l e c t i o n (e.g. s e l e c t i o n of home range) determine what i s a v a i l a b l e a t hig h e r o r d e r s of s e l e c t i o n (e.g. s e l e c t i o n of h a b i t a t s w i t h i n home range) can a s s i s t managers i n interpreting selection r e s u l t s . In the above moose example, the value of wetlands may have been underestimated i f home range se l e c t i o n had not been considered. Defining a v a i l a b i l i t y by an animal's home range incorporates a l o g i c f a u l t related to selection order. To measure sel e c t i o n , the use must be measured and then compared to the a v a i l a b i l i t y . The a v a i l a b i l i t y i s determined by the home range which i s determined by what was used! This r e s u l t s i n use being compared to use, not a v a i l a b i l i t y . 6.1.5 Is Use Equivalent to Selection? As discussed e a r l i e r , s e l e c t i o n i n d i c e s r e l y on what i s av a i l a b l e . This i s where an inherent problem of these indice s l i e s . For a habitat unit to be selected, i t must be used to a greater extent than i t i s available; therefore, a common habitat w i l l r a r e l y be defined as selected f o r . For example, a habitat that covers 50% of an animal's home range may have to be used over 75% of the time to s t a t i s t i c a l l y be c l a s s i f i e d as selected f o r (McLellan 1985). Furthermore, i t would be impossible to c l a s s i f y a h a b i t a t as h i g h l y s e l e c t e d (SI >1.5 by A u l t and Stormer's (1983) d e f i n i t i o n ) , i f that habitat exceeded 67% of the a n i m a l ' s home range ( l 0 0 % / 6 7 % = 1.5). T h i s i s a common occurrence i n g r i z z l y bear studies where timber often makes up a la r g e component of g r i z z l y bear home ranges. Because of the abundance of timber i n the home range, timber i s r a r e l y defined as s e l e c t e d f o r , despite being used as much as 40% of the time (McLellan 1985). In these cases, high use of hi g h l y a v a i l a b l e habitats may be purposeful " s e l e c t i o n " on the part of the animal, despite a negative selection value. 6.1.6 Is The Logic Flawed? The h a b i t a t s e l e c t i o n index has several f a u l t s associated with i t as has been discussed. I f these f a u l t s are considered when the index values are being interpreted, then, t h e o r e t i c a l l y , u s e f u l information can be obtained. However, the f o l l o w i n g argument suggests that there i s a major f a u l t i n the l o g i c of the index which negates i t s v a l i d i t y . The reasons f o r t h i s argument are that habitat s e l e c t i o n indices are predicated on two f a l s e presumptions and measured with an i l l o g i c a l u n i t . 72-Presumptions 1. Presumes that animals select habitat. I t i s l i k e l y that animals select a t t r i b u t e s associated with habitats rather than habitats themselves. Habitats are units of land around which we a r b i t r a r i l y draw boundaries based on our own perceptions. Thus, for example, we may define spruce forests and pine f o r e s t s as separate habitats. A moose, however, i s not a taxonomist and may not care whether a forest i s pine or spruce, but rather i s only concerned that the forest provides cover. I f animals are r e a l l y s e l e c t i n g f o r a t t r i b u t e s , then s e v e r a l h a b i t a t s may provide the a t t r i b u t e that the animal seeks. The habitats that we define are therefore probably d i f f e r e n t from the habitats a moose "sees." 2. Presumes that we need to know animal's habitat preferences. I f one accepts the argument that animals s e l e c t s p e c i f i c p h y s i c a l a t t r i b u t e s , then i t f o l l o w s t h a t d e t e r m i n a t i o n of h a b i t a t s e l e c t i o n may be in a p p r o p r i a t e . What we are r e a l l y i n t e r e s t e d i n i s a t t r i b u t e s e l e c t i o n . These a t t r i b u t e s may or may not be associated with the habitats that we have de f i n e d ; therefore, habitat selection indices w i l l only be representative i f there i s a high c o r r e l a t i o n between the h a b i t a t and the att r i b u t e ( s ) f o r which the animal selects. -73-6.1.7 The Unit i s I l l o g i c a l The concept of selection was i n i t i a l l y derived i n reference to food preference. The r a t i o of percent of each food item i n the d i e t versus percent of each food item a v a i l a b l e as a measure of preference seems l o g i c a l . The units are consistent. However, the u n i t s f o r h a b i t a t s e l e c t i o n are not c o n s i s t e n t . The proportion of time spent i n a habitat i s compared to the area of the habitat. I t appears i l l o g i c a l to expect any r e l a t i o n s h i p to ex i s t between these variables. I t i s l i k e l y that the s i z e of the area encompassed by a p a r t i c u l a r habitat has nothing to do with the length of time that an animal spends i n that h a b i t a t , and t h a t the d u r a t i o n of occupation i s more a f u n c t i o n of the a c t i v i t y associated with that habitat. For example, the s i z e of a feeding s i t e has nothing to do with how long i t takes an animal to eat. The area encompassed by protective cover has nothing to do w i t h how lon g an animal beds. The time spent i n each "habitat" i s l i k e l y determined by the a c t i v i t y performed i n that h a b i t a t . I f a moose spends 3 0% of i t s day feeding, you would expect i t to be i n a feeding s i t e 30% of the day. I t does not seem l o g i c a l to expect the moose to be i n the feeding s i t e longer because the feeding s i t e i s larger than 30% of i t s home range or sh o r t e r because the feeding s i t e i s l e s s than 30% of i t s home range. To t h i s end, the argument that use of a habi t a t by an animal i s a c t u a l l y selection of that habitat appears to be more appropriate. The habitat i s chosen because of the a c t i v i t y the animal chooses to pursue, and the length of time the animal 74-spends i n that habitat depends upon the a c t i v i t y chosen. This argument addresses two other problems associated with habitat s e l e c t i o n indices: 1) Resource density 2) Contagion/Dispersion. Resource Density Consider the following scenario: . = resource (food) a b 'a' and 'b' are equal areas i d e n t i c a l i n every r e s p e c t except that the density of food i n 'b' i s much greater than i n 'a'. An animal eating i n 'a' w i l l be required to eat longer to consume the same amount of food as an animal eating i n 'b'. The l i k e l i h o o d of fin d i n g an animal i n 'a' i s therefore greater than the l i k e l i h o o d of finding an animal i n 'b'. A habitat s e l e c t i o n index would show that 'a' has a higher s e l e c t i o n value than 'b', yet 'b' provides a greater resource density. As an a l t e r n a t i v e , the researcher could t r y to determine which a c t i v i t i e s the animal performs i n each habitat. I f these a c t i v i t i e s are known, a r e l a t i v e ranking of the importance of a h a b i t a t , compared to other h a b i t a t s which p r o v i d e the same at t r i b u t e s , can be made. -75-Contagion and Dispersion The e f f e c t s of contagion and dispersion on habitat s e l e c t i o n indices can also be demonstrated by example. • • • • • A . B . A . A . A . B x y . = moose location Assume that A i s a feeding habitat and B i s a habitat that moose normally don't use. 'x' and 'y' are i d e n t i c a l i n every way except i n the p o s i t i o n i n g of the u n i t s . In 'x' h a b i t a t B separates two units of habitat A. In 'y' both units of habitat A are adjacent. In 'x' the dispersion of the A units r e s u l t s i n equal use of a l l three units (B i s used only as a t r a v e l corridor between A and B), whereas i n 'y', B i s never used. Contagion and dispersion c l e a r l y can a f f e c t the habitat s e l e c t i o n index. Once again, determining the a c t i v i t y associated with each habitat would i d e n t i f y why each habitat was used and would allow the importance of each habitat to be assessed. 6.1.8 When Do Habitat Selection Indices Work? I believe that there are only 2 s i t u a t i o n s where a habitat s e l e c t i o n index can provide an accurate assessment of habitat importance: (1) when a habitat of r e l a t i v e l y low a v a i l a b i l i t y i s used extensively, and (2) when a habitat of r e l a t i v e l y high -76 a v a i l a b i l i l t y i s r a r e l y used. Provided the problems which have been discu s s e d regarding a v a i l a b i l i t y are addressed, u s e f u l information concerning these habitats can be elucidated. 6.1.9 Recommendations Discussion i n t h i s paper has revealed that there are many a s p e c t s o f h a b i t a t s e l e c t i o n i n d i c e s t h a t make t h e i r i n t e r p r e t a t i o n l e s s than s t r a i g h t - f o r w a r d . The f o l l o w i n g recommendations are suggested. F i r s t , t o e n s u r e t h a t v a l u a b l e h a b i t a t s a r e n o t u n i n t e n t i o n a l l y l o s t t o development or i n d u s t r y , i t i s recommended that land use managers use other t e c h n i q u e s , i n c o n j u n c t i o n w i t h s e l e c t i o n i n d i c e s , to e v a l u a t e w i l d l i f e habitat. For example, " p r i n c i p a l habitats" should be i d e n t i f i e d . Petrides (1975) describes a p r i n c i p a l food as one which i s eaten i n greatest quantity. A p r i n c i p a l habitat could be defined as one which i s used the greatest amount of time. Whether or not the animal prefers t h i s habitat i s not important. What i s important i s that despite the fact i t may not be selected f o r , i t i s used the majority of the time. Use of t h i s technique w i l l safeguard against discounting heavily-used, "avoided" habitats as important components. A p o t e n t i a l flaw with t h i s technique i s a habitat may be used extensively due to "forced" use, yet w i l l s t i l l be c l a s s i f i e d as a p r i n c i p a l habitat. The second recommendation concerns r e s e a r c h . H a b i t a t s e l e c t i o n i n d i c e s have been extensively used i n the past as a means of i n d i r e c t l y determining habitat importance. I t has been -77-considered too d i f f i c u l t to c o l l e c t t h i s kind of information v i a other techniques. Discussion i n t h i s paper suggests that the v a l i d i t y of habitat s e l e c t i o n indices i s questionable and that the information gained i s therefore l i k e l y u n r e l i a b l e . I t i s , t h e r e f o r e , recommended t h a t i f a measure of importance of habitats i s desired, data which more d i r e c t l y measure importance should be c o l l e c t e d . Determining the a c t i v i t i e s associated with s p e c i f i c habitat a t t r i b u t e s , f o r example, w i l l determine more e f f e c t i v e l y why animals use the areas they do and w i l l allow the importance of habitats to be assessed. With t h i s background of selection indices, s e l e c t i o n r e s u l t s from t h i s study w i l l be discussed. 6.2 SELECTION RESULTS 6.2.1 Cover Type Classes The n u l l hypothesis that moose are randomly d i s t r i b u t e d with respect to cover types was rejected. Three cover types were used p r o p o r t i o n a t e l y greater than they were av a i l a b l e , while 5 cover types were used proportionately less than they were a v a i l a b l e . Of a l l f o r e s t cover types, only immature spruce and mature spruce were used i n proportions greater than t h e i r a v a i l a b i l i t y . This suggests that there i s a c h a r a c t e r i s t i c of spruce forests that moose seek. A l i k e l y c h a r a c t e r i s t i c i s the r e l a t i v e l y high abundance of browse associated with spruce f o r e s t s compared to pine f o r e s t s . The average shrub cover i n spruce f o r e s t i s 7%, compared to 1% i n lodgepole pine forests. Spruce forests then, can provide both a source of food and a source of cover. -78-Mature spruce forests also provide a greater canopy closure than pine f o r e s t s . For example, the average canopy closure of both intermediate and mature pine was 5%, whereas the average canopy closure f o r mature spruce was 7%. The average canopy closure value for spruce forests that moose used was 10%. Spruce wetlands were the only wetlands that were used i n greater proportion than t h e i r a v a i l a b i l i t y . These wetlands also received the greatest amount of use of any f o r e s t or wetland c l a s s . The main distinguishing c h a r a c t e r i s t i c of spruce wetlands from other cover types (with the exception of spruce forests) i s the presence of both food and cover (in the form of spruce trees) within one cover type. Ty p i c a l l y , forests provide cover, but not food, while wetlands provide food, but not cover. I t may be then, that moose use spruce wetlands because they provide not only an abundant food source, but also protective cover. Use of these wetlands would reduce the amount of t r a v e l moose undertake between wetland feeding areas and forested bedding areas, and may also provide a decreased r i s k of predator attack while foraging. Mature lodgepole pine forests were used i n lower proportion than a v a i l a b i l i t y ; however, they were the second most often used f o r e s t cover type and t h i r d most often used o v e r a l l cover type (12.4%). I t i s u n l i k e l y that use was l e s s than a v a i l a b i l i t y because moose do not choose mature lodgepole pine, but rather because the a v a i l a b i l i t y of mature lodgepole pine was so high (29.2%). -79-Use of both grass and sedge wetlands was low, despite the f a c t that each of these classes makes up a s i g n i f i c a n t component of the a v a i l a b l e cover types. I t i s l i k e l y , then, that moose choose n o t t o use t h e s e t y p e s o f w e t l a n d s . The main di s t i n g u i s h i n g c h a r a c t e r i s t i c between these 2 classes of wetlands and other wetland classes i s that they contain few shrubs. While sedges and g r a s s e s are eaten by moose, the snow cover may preclude t h e i r a v a i l a b i l i t y . I t may be that moose do not use grass and sedge wetlands because, u n l i k e wetlands c o n t a i n i n g shrubs, they do not contain available forage during the winter. 6.2.2 Distance To Edge The n u l l hypothesis that moose are randomly d i s t r i b u t e d with respect to distance from forest/wetland border was rejected. In both winters, use of the 0-20 m class was proportionately greater than i t s a v a i l a b i l i t y . In general, areas greater than 100 m from the edge were used proportionately less than they were a v a i l a b l e . Of p a r t i c u l a r importance i s that despite the f a c t that the 0-20 m clas s was abundant (31.1%), use s t i l l exceeded a v a i l a b i l i t y . The high l e v e l of use of t h i s class l i k e l y r e f l e c t s i t s importance. The suggestion that moose seek areas that provide both food and cover can again be used to explain t h i s pattern of use. By concentrating use close to the edge, moose take advantage of the hig h shrub p r o d u c t i o n of wetlands and the cover provided by fo r e s t s . Regardless of whether moose were observed i n the forest or i n the wetland, use was concentrated near the edge; however, moose i n wetlands did not move away from the edge as frequently -80-as moose i n the forests. This supports the idea that moose use f o r e s t s f o r cover. Venturing away from the f o r e s t c o u l d be dangerous, but venturing away from the wetland would increase protection from predators or thermal stress. 6.2.3 Forest Age Moose seem to select older age classes. Forests older than 12 0 years were used p r o p o r t i o n a t e l y g r e a t e r than they were a v a i l a b l e . T h i s c o u l d be a r e s u l t of moose s e e k i n g these f o r e s t s , or i t could be an arithmetic r e s u l t caused by the low abundance of these f o r e s t s . Due to the low a v a i l a b i l i t y , few o b s e r v a t i o n s i n t h e s e f o r e s t s would be r e q u i r e d t o show se l e c t i o n . This v a r i a b l e may not be a useful c r i t e r i o n f o r explaining patterns of use of cover types by moose. Large v a r i a t i o n s i n p h y s i c a l structure can e x i s t i n f o r e s t s within each age c l a s s . For example, i n age class 41-60 years old, height varied between 5 and 16.5 m, density varied between 3 and 33 stems/100 square meters, and DBH varied between 8 and 22 cm. I f moose choose cover types because of physical structure of the cover type, then age c l a s s i s l i k e l y not a useful variable to measure. 6.2.4 Forest Density and DBH The pattern of use of density classes suggests that moose prefer forests of intermediate density. The reason f o r t h i s may be that dense forests r e s t r i c t locomotion, while open forests do not provide adequate cover, either for hiding from predators or reducing thermal s t r e s s . Forests of intermediate d e n s i t y may provide the optimum balance of locomotion ease and cover. -81-While the v a r i a b i l i t y of ph y s i c a l s t r u c t u r e i n f o r e s t s of the same age c l a s s may be l a r g e , there i s , n e v e r t h e l e s s , a r e l a t i o n s h i p among forest age, density, and DBH. As forest age increases, density decreases and DBH increases. The s e l e c t i o n data indicates that moose use forests older than 120 years, with diameters greater than 25 cm, and densities between 9 and 11 cm i n greater proportion than they are av a i l a b l e . The average DBH and density of f o r e s t s >120 years i s 25 cm and 9-11 stems/100 square meters respectively. The fa c t that older f o r e s t s provide the DBH and density that moose prefer may be the reason why they were used i n proportion greater than a v a i l a b i l i t y . 6.2.5 Forest Height Tree height does not appear to influence which f o r e s t cover types moose choose to use. I f moose use f o r e s t s p r i m a r i l y f o r hiding, t h i s observation i s l o g i c a l . Providing trees are t a l l e r than a c e r t a i n minimum height (2 m) they w i l l provide s u f f i c i e n t cover to hide a moose. Forests less than 2 m i n height may not provide s u f f i c i e n t cover and therefore may not used. Use of f o r e s t s w i t h t r e e s l e s s than 2 m t a l l was not s i g n i f i c a n t l y d i f f e r e n t from a v a i l a b i l i t y ; however, t h i s may be a r e s u l t of the low a v a i l a b i l i t y of these f o r e s t s . Because the a v a i l a b i l i t y of these forests was so low, even no observations of moose r e s u l t e d i n no s i g n i f i c a n t d i f f e r e n c e between use and a v a i l a b i l i t y . -82-6.2.6 Forest Canopy Closure The s e l e c t i o n r e s u l t s indicate that moose use f o r e s t s with canopy c l o s u r e s >25% p r o p o r t i o n a t e l y greater than they were a v a i l a b l e , while f o r e s t s with canopy closures of 3% were used l e s s than they were available. This pattern of use may r e f l e c t the magnitude of the a v a i l a b i l i t y of these c l a s s e s rather than moose biology. The >25% class was only 2.4% a v a i l a b l e , and thus r e l a t i v e l y few observations were needed to show use greater than a v a i l a b i l i t y . In contrast, the 3% c l a s s was l a r g e l y a v a i l a b l e (21.1%) and thus many observations would be needed i n order to show use equal to or greater than a v a i l a b i l i t y . I f t h i s i s the case, then moose do not choose forests because of canopy-closure. Some other variable/s is/are more important. Canopy closure values were compared to f o r e s t age, density and DBH. Independent comparisons between each of these variables and canopy-closure did not demonstrate any r e l a t i o n s h i p s . This i s l i k e l y because canopy closure i s related to a combination of these v a r i a b l e s ; therefore, forests of a va r i e t y of d i f f e r e n t age classes l i k e l y provide the canopy closure that moose prefer. 6.2.7 Forest Shrub Cover The s e l e c t i o n pattern demonstrated f o r f o r e s t shrub cover may be explained by arguments si m i l a r to that f o r f o r e s t canopy closure. Use d i f f e r e d from a v a i l a b i l i t y i n only 2 classes. This i s l i k e l y a f a l l a c y r e s u l t i n g from the extreme a v a i l a b i l i t y values and not a r e s u l t of moose selection or avoidance. Use of the 1% cover class was less than a v a i l a b i l i t y , yet was greater -83-t h a n i n any o t h e r c l a s s . The h i g h a v a i l a b i l i t y (63.9%) outweighed the high use (37.7%). Conversely, use of the 20% c l a s s was g r e a t e r than a v a i l a b i l i t y , l i k e l y because of the extremely low a v a i l a b i l i t y (0.7%). Few observations i n t h i s c l a s s would r e s u l t i n selection being indicated. The extensive use of forests with l i t t l e shrub cover implies that moose l i k e l y use forests for shelter and not for feeding. 6.2.8 Wetland Shrub Cover The s e l e c t i o n index i n d i c a t e s that moose d i d not s e l e c t wetlands with shrub cover greater than 25%. This r e s u l t e d from the high a v a i l a b i l i t y of t h i s c l a s s (35.6%) rather than moose s e l e c t i o n against t h i s c l a s s . In fact, t h i s class received 45.3% of use. This pattern of use i s l i k e l y an example of s e l e c t i o n at the home-range l e v e l as discussed by Johnson (1980) . Moose l i k e l y winter i n the study area because of the high abundance of browse-producing wetlands r e l a t i v e to surrounding areas. The high a v a i l a b i l i t y of these wetlands prevents s e l e c t i o n from being demonstrated despite high use. If sel e c t i o n were being studied at the home range l e v e l rather than the cover type within home range l e v e l , the importance of these high browse p r o d u c t i o n wetlands would l i k e l y be demonstrated. The s e l e c t i o n index worked well for wetlands with <5% shrub cover. In t h i s case, the negative s e l e c t i o n index r e s u l t i s l i k e l y i n d i c a t i v e of the l i t t l e importance of these wetlands for moose. Despite comprising a l a r g e p r o p o r t i o n of a v a i l a b l e wetlands (28.8%), there were no observations of moose i n these wetlands. The negative selection r e s u l t for the 1 and 3% shrub -84-cover c l a s s e s demonstrates that these wetlands are of l i t t l e value to moose. The use p a t t e r n was one of increasing use as shrub cover increased. The fac t that the a v a i l a b i l i t y of each shrub cover cl a s s also increased as shrub cover increased makes i t d i f f i c u l t to discern i f moose seek out high shrub cover classes or i f they are randomly using wetlands. Considering that wetlands with l i t t l e shrub cover were l a r g e l y a v a i l a b l e but never used, i t appears that moose do seek areas with high shrub cover. 6.2.9 Wetland Shrub Height Selection occurred for the 0.6-1.0 m-class only. Use of the 1.1-1.5 m c l a s s was h i g h (30.2%), but s e l e c t i o n was not demonstrated because a v a i l a b i l i t y was also high (21.3%). I t i s l i k e l y that these two classes are preferred by moose. The n e g a t i v e s e l e c t i o n of the 0.0-0.5 m-class l i k e l y r e f l e c t s the low value of t h i s c l a s s f o r w i n t e r i n g moose. Despite comprising 41.5% of the a v a i l a b l e wetlands, t h i s class was only used 16.3% of the time. Two possible explanations f o r such low moose use of t h i s class are related to shrub cover and snow depth. The percent shrub cover i s related to shrub height. For example, the average shrub cover class for shrubs under 0.5 m i s 7%, while the average shrub cover class f o r shrubs over 3.0 m i s 40%. Moose may use the <0.5 m c l a s s i n f r e q u e n t l y because they do not have enough forage. A l t e r n a t i v e l y , snow i n wetlands was deep enough to bury shrubs <0.5 m. This r e s u l t s i n shrubs les s than <0.5 m t a l l being d i f f i c u l t to f i n d . The <0.5 m shrub height c l a s s may have been used proportionately less than -85-i t s a v a i l a b i l i t y then, because browse a v a i l a b i l i t y was low. 6.3 Importance of Wetlands S e l e c t i o n r e s u l t s from the wetland shrub cover classes and the wetland shrub height c l a s s e s s u b s t a n t i a t e s the idea that moose use wetlands f o r feeding. During the winter, the only p h y s i c a l d i f f e r e n c e among wetlands i s the d i f f e r e n c e i n shrub abundance. The fa c t that wetlands with low shrub cover values or with short shrubs were r a r e l y used and that wetlands with high shrub cover values were often used suggests that the a v a i l a b i l i t y of food plays a r o l e i n determining why moose use wetlands. 6.4 Monthly Analyses of Use This research project was designed to determine patterns of h a b i t a t s e l e c t i o n of a l l r a d i o - c o l l a r e d moose f o r the e n t i r e winter season, not for in d i v i d u a l moose or for i n d i v i d u a l months. Th i s was done i n order to o b t a i n adequate sample s i z e s f o r s t a t i s t i c a l t e s t s . The sample s i z e s on a monthly basis or for i n d i v i d u a l animals are not adequate to s a t i s f y s t a t i s t i c a l requirements and therefore test s t a t i s t i c s were not applied. For example c h i - s q u a r e t e s t s can be u s e d t o d e t e r m i n e i f d i s t r i b u t i o n s of moose use of measured variables d i f f e r e d between months or between the same month of d i f f e r e n t years. However, i n order to use the chi-square approximation, no more than 20 percent of a l l categories may contain l e s s than f i v e expected observations (Roscoe and Byars 1971). Since t h i s requirement was not met, the chi-square approximation could not be used. For -86-t h i s r e a s o n , the monthly use r e s u l t s must be i n t e r p r e t e d s u b j e c t i v e l y . They are presented not as a b s o l u t e f a c t , but rather to stimulate development of other hypotheses which could be tested i n the future. 6.4.1 Forest Versus Wetland The increasing use of wetlands as winter progressed d i f f e r s from p a t t e r n s of moose use documented i n other winter habitat s t u d i e s . Eastman (1979), P i e r c e and Peek (1984), and Schwab (1985) a l l documented use of heavy f o r e s t cover during l a t e winter. The explanation proposed by Pierce and Peek as well as by Eastman was that moose were taking advantage of decreased snow depths i n forests as compared to open areas. The decreased snow depth f a c i l i t a t e s e a s i e r locomotion and thus reduces energy expenditure. Schwab d i d not r e l a t e t h i s p a t t e r n of use to snow-depth, but hypothesized that avoiding heat s t r e s s i n l a t e winter may play an important r o l e . His b e l i e f i s substantiated by Renecker and Hudson (1986), who demonstrated t h a t moose metabolic rates were unaffected by low temperatures (-25 - -30 C) but temperatures >-5 C i n the winter caused stress as measured by increased r e s p i r a t i o n r a t es. Data from my study suggests that neither of these factors influences patterns of winter cover type use by moose. I t i s u n l i k e l y that seeking areas of reduced snow-depth provided by forested areas i s a s i g n i f i c a n t f a c t o r i n causing moose to change cover type preference, at l e a s t over the range of snow depths encountered i n t h i s study. For example, despite 87-average snow-depths remaining nearly constant i n January (48 cm), February (48 cm) and March (42 cm) of 1987, use of wetlands increased from 34% to 57%. In 1988, the average snow depths i n January and March were nearly equal (21 and 24 cm r e s p e c t i v e l y ) , while the average snow depth i n February was much greater (35 cm) . Moose use of wetlands d i d not r e f l e c t these changes, but increased p r o g r e s s i v e l y throughout the winter. To substantiate t h i s argument furt h e r , snow depths i n 1987 were s u b s t a n t i a l l y greater than i n 1988. In fact, the average snow-depth i n January 1987 was 27 cm greater than i n January 1988. I f snow depth played a factor i n determining which cover types moose use, then use of forested cover types i n January 1987 should have been greater than i n January 1988. This was not the case. Avoiding heat s t r e s s a l s o does not e x p l a i n the observed p a t t e r n of use. In 1988, the average monthly temperature, c a l c u l a t e d from data a s s o c i a t e d with each moose r e l o c a t i o n f l i g h t , dropped between January and February, while the average March temperature exceeded that of January. The average monthly temperature did not change among a l l 3 months i n 1987. I f moose were avoiding heat stress, then i n March 1988 there should have been more observations i n forests than i n wetlands. In 1987 the p a t t e r n of cover type use s h o u l d have remained the same throughout the winter. This argument does not completely ru l e out heat stress as a factor i n determining cover type use because absolute temperature i s not the only variable influencing body temperature of an -88-animal. Two other important variables, which were not measured, are d i r e c t s u n l i g h t and wind speed. C o n s i d e r a t i o n of these variables i s necessary before heat stress can be eliminated as a contributing factor i n moose habitat s e l e c t i o n . 6.4.2 Forests The only dramatic change i n patterns of f o r e s t cover use occurred i n 1988 when use of pine dropped 38 percentage points between January and March. This decline i n use of pine coincided with a 15 percentage point increase i n use of spruce as well as a 19 percentage point increase i n wetland use. This s h i f t i n cover type use i s d i f f i c u l t to explain using arguments of snow-depth and heat stres s because spruce f o r e s t s and w e t l a n d s a r e extreme o p p o s i t e s i n terms of p h y s i c a l s t r u c t u r e . In other s t u d i e s , moose move from open areas to f o r e s t s . In t h i s study, there was an increase i n use of open areas and spruce forests as winter progressed. One explanation may be that low forage i n West-Chilcotin forests means that moose must use wetlands to f o r a g e . R e g a r d l e s s of e n v i r o n m e n t a l conditions, moose may be forced to feed i n open areas. However, they w i l l be f r e e to choose t h e i r bedding s i t e s and perhaps because of c e r t a i n environmental c o n d i t i o n s (e.g. high s o l a r r a d i a t i o n ) they choose spruce f o r e s t s (which provide greater canopy cover). As discussed previously, spruce f o r e s t s c o n t a i n abundant browse r e l a t i v e to p i n e f o r e s t s . T h i s s h i f t i n use from lodgepole pine forests to wetlands and spruce f o r e s t s may be a r e s u l t of moose seeking areas of abundant browse. -89-6.5 DIET COMPOSITION Moose are p r i m a r i l y a browsing species, e s p e c i a l l y during the winter. In south-central Alaska, willows and Kenai b i r c h (Betula kenaica) are the most preferred species, with cottonwoods (Populus balsamifera), high bush cranberry (Viburnum edule), red elder (Sambucus racemosa), rose (Rosa spp) and raspberry (Rubus idaeus) being l e s s important (Peek, 1974). Spencer and Hakala (19 64) recorded various willows as the most important d i e t a r y component, w i t h bog b i r c h , dwarf b i r c h ( B e t u l a n a n a ) , s e r v i c e b e r r y , mountain ash (Sorbus e d u l e ) , and h i g h - b u s h cranberry being of minor importance. Moose food habit studies from the Kenai peninsula i n Alaska show that sedges were used when snow depths were less than 30 cm. When snow depths exceeded 30 cm, b i r c h stems comprised 72% of d i e t s from February to May (LeResche and Davis 1973). Studies i n B r i t i s h Columbia have shown important wintering moose f o r a g e s p e c i e s t o be r e d - o s i e r dogwood (C o r n u s s t o l o n i f e r a ) , paper b i r c h ( B e t u l a p a p y r i f e r a ) , w i l l o w s , s e r v i c e b e r r y , quaking aspen, h a z e l ( C o r y l u s c a l i f o r n i c a ) . high-bush cranberry (Viburnum pauciflorum) a l p i n e f i r (Abies  lasiocarpa and mountain ash (Peek 1974). This study demonstrates that lodgepole pine and bog b i r c h are two important browse species for moose i n the C h i l c o t i n . I am not aware of any other North American moose study that shows lodgepole pine being a component of moose d i e t s . One Swedish paper documents damage to North American lodepole pine -90-provenances by moose (Hansson 1985). S i m i l a r l y , bog-birch, which i s considered to be of only minor importance i n other studies, was an important component of moose d i e t s i n the C h i l c o t i n . S u r p r i s i n g l y , bog bi r c h was preferred over willow. The high use of sedges i n 1988 versus 1987 may be due to snow depths i n 1988 being less than 30 cm f o r a large part of the winter. This i s consistent with the fi n d i n g s of LeResche and Davis (1973). 6.6 DIET QUALITY Crude Protein Protein i s considered by many to be the most important plant n u t r i e n t because i t p r o v i d e s n i t r o g e n r e q u i r e d by rumen microorganisms for growth and i s es s e n t i a l f o r body maintenance, growth, reproduction and l a c t a t i o n (Oldemeyer et a_l. 1977) . Plant protein l e v e l s cycle throughout the year with lowest l e v e l s o c c u r r i n g during the winter months. In t h i s study, with the exception of sedges, a l l the major forage species consumed during the period of study had crude p r o t e i n l e v e l s between 6 and 7%. These l e v e l s are s i m i l a r to values obtained from moose browse i n Alaska (Gasaway and Coady 1974). These l e v e l s are l i k e l y minimum r e q u i r e d p r o t e i n l e v e l s . Schwartz et a l . (1987) demonstrated that Alaskan moose need a minimum dietary crude protein l e v e l of 6.8% to meet maintenance requirements. Corbett (1969) stated that the minimum l e v e l f o r ruminants i s about 7%. Murphy and Coates (19 66) found that white-tailed deer fed 7% protein diets throughout the winter were p h y s i c a l l y stunted and t h a t does produced fewer fawns compared to does fed higher protein l e v e l -91-0 d i e t s . Dietz et a l . (1962) c l a s s i f i e d mule deer winter range as good i f browse contained a minimum of 7% crude protein. Gross Energy and Neutral Detergent Fiber These values were presented as a matter of i n t e r e s t . Gross energy values f o r domestic animal feeds range between 4300 and 4700 cal / g (Maynard and L o o s l i 1969). Values f o r several species of moose browse range from 4250 c a l / g to 4870 c a l / g . An outstanding exception i s common birc h (5440 cal/g) ( H j e l j o r d et a l . 1982). Gross energy values are not uniform throughout the plant. The values reported i n t h i s study, with the exception of sedges appear unusually high. This may be a r e s u l t of moose feeding p r e f e r e n t i a l l y on plant components which c o n t a i n high gross energy. The n e u t r a l detergent f i b e r percentages reported here are a l s o s u s p i c i o u s l y high. N e u t r a l detergent f i b e r v a l u e s of composite moose d i e t s reported by Renecker and Hudson (1985) ranged from 48.9 to 52.6% during the winter period. 6.7 Summary This study demonstrates the importance of forest-wetland edges to wintering moose. The edge ecotone i s l i k e l y important b e c a u s e i t s i m u l t a n e o u s l y p r o v i d e s f o o d and c o v e r . Spruce-wetlands and spruce f o r e s t s , a l s o important h a b i t a t s , provide the same combination of e s s e n t i a l s . The high use of f o r e s t s with low shrub a v a i l a b i l i t y i s consistent with the idea that moose use f o r e s t s p r i m a r i l y as a source of cover and not forage. Conversely, the p o s i t i v e c o r r e l a t i o n between wetland -92-shrub cover and moose use i s c o n s i s t e n t with the idea that wetlands are used primarily for food. The i n c r e a s e d use o f wetlands as w i n t e r p r o g r e s s e d contradicts other North American moose studies, but snow depth i n t h i s study area did not exceed the c r i t i c a l threshold f o r moose. The importance of snow depth i n determining cover type s e l e c t i o n i n other studies was l i k e l y not s i g n i f i c a n t i n t h i s study. Selection r e s u l t s of forest attributes were as follows: 1. Age - only forest age classes 121-140 and >140 years were s e l e c t e d i n proportions g r e a t e r than random (use = 9.6 and 11.0% re s p e c t i v e l y ) . Use was high i n f o r e s t s aged 21-40 year s (16.4%), 41-80 y e a r s (19.2%), and 81-100 years (16.4%), but d i d not exceed that expected by random use.. 2 2. D e n s i t y - f o r e s t s w i t h 9-11 stems/100 m were selected i n proportions greater than random (use = 28.6%). 3. Height - s e l e c t i o n g r e a t e r than random was not demonstrated for any of the forest height classes. 4. DBH - only forests with DBH >24 cm were selected i n proportions greater than random (use = 26.7%). Use was high i n the 16.1-20.0 cm class (26.7%) and the 2 0.1-24.0 cm class (18.7%), but did not exceed that expected by random use. 5. Canopy closure - only forests with canopy closure > 25% were selected i n proportions greater than random (use = 10.4%), but use was s i g n i f i c a n t (>10%) i n a l l forests with canopy closure >5%. 6. Forest shrub cover - f o r e s t s with 2 0% shrub cover were s e l e c t e d i n proportions g r e a t e r than random (use = 7.8%). Use was high i n the 1, 5, and 10% s h r u b c o v e r c l a s s e s (37.7, 18.2, and 10.4% respe c t i v e l y ) , but did not exceed that expected by random use. -93-7. MANAGEMENT RECOMMENDATIONS T h i s r e s e a r c h h as d o c u m e n t e d s e v e r a l i m p o r t a n t c h a r a c t e r i s t i c s of moose use of cover types that have important i m p l i c a t i o n s to f o r e s t h a r v e s t i n g p r a c t i c e s . Of c r i t i c a l importance to moose i s the ecotone between upland f o r e s t s and wetlands. A d d i t i o n a l l y , the importance of spruce f o r e s t s and spruce wetlands i s highlighted. Management recommendations are concerned with these cover types. 1. Use of the edge between f o r e s t s and wetlands i s high very close to wetlands, e s p e c i a l l y wetlands with > 15% shrub cover. Use drops o f f dramat i c a l l y as distance from the edge increases. Beyond 2 00 m from the edge, use was low i n 1988 (2%), but was n o t a b l e i n 1987 (12.1%). I t i s recommended that a border of conifers no le s s than 2 00 m be l e f t around wetlands, e s p e c i a l l y i f they contain >15% shrub cover. In order f o r a moose to remain 200 m from the open, there must also be a 200 m s t r i p of conifer l e f t around the cut-block; therefore, logging should be r e s t r i c t e d within 400 m of wetlands. In a r eas where moose d e n s i t i e s a r e h i g h , and the dispersion of wetlands i s such that l i t t l e c o nifer cover e x i s t s beyond 400 m from wetlands, logging should be avoided. Spruce wetlands and mature spruce f o r e s t s , e s p e c i a l l y those within 200 m from to wetlands, appear to be very important to moose. Disturbances near these cover types should be minimized. While sedge meadows were not used e x t e n s i v e l y by wintering moose, they may be very important cover types, e s p e c i a l l y i n the spring, as sedges are among the f i r s t vegetation to break dormancy. In t h i s study, use of sedge meadows was commonly observed i n A p r i l . Sedges were also a s i g n i f i c a n t component of some p e l l e t groups c o l l e c t e d i n A p r i l ; therefore, logging should also be r e s t r i c t e d within 400 m of sedge meadows. Use of immature l o d g e p o l e p i n e stands was low i n t h i s study, but may have resulted from low a v a i l a b i l i t y of t h i s forest c l a s s . Lodgepole pine was an important component of moose winter d i e t s , and heavy browsing of immature lodgepole pine branches was often noted. The p o t e n t i a l e x i s t s to make a v a i l a b l e abundant c o n i f e r browse by logging lodgepole pine f o r e s t s and allowing them to regenerate. Moose may take advantage of the food and cover that occur simultaneously i n t h i s newly c r e a t e d type of f o r e s t cover. Heavy moose use of regenerating stands could, however, create s i l v i c u l t u r e problems. 5. I t must be noted t h a t t h i s study was s e l e c t i v e i n terms of the animals used and the period i n which data were co l l e c t e d . These data are from adult female moose i n the winter p e r i o d . Use of cover t y p e s d u r i n g d i f f e r e n t parts of the year and by other sex and age c l a s s e s of moose may d i f f e r ; t h e r e f o r e , data from d i f f e r e n t times of the year as well as from bull-moose and immature animals are desirable. -96-8. LITERATURE CITED Allredge, J.R., and J.T. R a t t i . 1986. Comparison of some s t a t i s t i c a l techniques for analysis of resource s e l e c t i o n . J . W i l d l . Manage. 50:157-165. Annas, R.M., and R. Coupe, (eds.). 1979. Biogeoclimatic zones and subzones of the Cariboo Forest Region. B.C. M i n i s t r y of Forests Publication. Atmospheric Environment Service. 1981. Canadian Climate Normals 1951 - 1980. Environment Canada. Atmospheric Environment Service. 1986, 1987, 1988. Anahim Lake 5-Mile Ranch Climate Summary. Ault, S.C., and F.A. Stormer. 1983. Seasonal food s e l e c t i o n by scaled q u a i l i n northwest Texas. J . W i l d l . Manage. 47: 222-228. B r i t i s h Columbia Ministry of Forests. 1989. Sub-boreal pine spruce zone. Cariboo Forest Region. Forest Science Section. Draft. Brusnyk, L.M., and F.F. G i l b e r t . 1983. Use of shoreline timber reserves by moose. J. Wild l . Manage. 47:673-685. Byers, C.R., and R.K. Steinhorst. 1984. C l a r i f i c a t i o n of a technique f o r analysis of u t i l i z a t i o n - a v a i l a b i l i t y data. J. W i l d l . Manage. 48:1050-1053. Chesson, J . 1983. The estimation and analysis of preference and i t s r e l a t i o n s h i p to foraging models. Ecology 64:1297-1304. Coady, J.W. 1974. Influence of snow on behaviour of moose. Naturaliste Can. 101:417-436. Corbett, J.L. 1969. The n u t r i t i o n a l value of grassland herbage. In Gasaway and Coady (1974). Des Meules, P. 1964. The influence of snow on the behaviour of moose. Trans. Northeastern Sec. W i l d l . Conf. 21. Dietz, D.R., R.H. Udall, and L.E. Yeager. 1962. Chemical composition and d i g e s t i b i l i t y by mule deer of selected forage species, Cache l a Poudre Range, Colorado, i n Oldemeyer (1974) . Eastman, D.S. 1978. Habitat selection and use i n winter by moose i n sub-boreal spruce forests of northcentral B r i t i s h Columbia, and relationships to forestry. Ph.D. t h e s i s , University of B r i t i s h Columbia. -97-Fairbanks, W.S., J.A. Bailey, and R.S. Cook. 1987. Habitat use by a low-elevation, semicaptive bighorn sheep population. J . Wildl. Manage. 51:912-915. Gasaway, W.C., and J.W. Coady. 1974. Review of energy requirements and rumen fermentation i n moose and other ruminants. Nat. Can. 101:227-262. Hansson, L. 1985. Damage by w i l d l i f e , e s p e c i a l l y small rodents, to North American Pinus contorta provenances introduced into Sweden. Can. J . For. Res. 15:1167-1171. Hatter, J . 1950. The moose of central B r i t i s h Columbia. Ph.D th e s i s , Washington State University, Pullman. Heady, H.F. 1964. P a l a t a b i l i t y of herbage and animal preference. J . Range Manage. 17:76-82. Hess, A.D., and A. Swartz. 1940. The forage r a t i o and i t s use i n determining the food grade of streams. In Petrides (1975). H j e l j o r d , O., F. Sundstol, and H. Haagenrud. 1982. The n u t r i t i o n a l value of browse to moose. J . W i l d l . Manage. 46:333-343. Johnson, H.D. 1980. The comparison of usage and a v a i l a b i l i t y measurements for evaluating resource preference. Ecology 61:65-71. K e l s a l l , J.P., and W. Prescott. 1971. Moose and deer behaviour i n snow i n Fundy National Park, New Brunswick. Can. W i l d l . Serv. Rep. Ser. 15. Knowlton, F.F. 1960. Food habits, movements, and populations of moose i n the Gravelly Mountains, Montana. J . W i l d l . Manage. 24:162-170. LeResche, R.E., and J.L. Davis. 1973. Importance of non-browse foods to moose on the Kenai Peninsula, Alaska. J . W i l d l . Manage. 37:279-287. Lyon, L.J., and C.E. Jensen. 1980. Management implications of elk and deer use of clear-cuts i n Montana. J . W i l d l . Manage. 44:353-362. Marcum, C.L., and D.O. Loftsgaarden. 1980. A nonmapping technique for studying habitat preferences. J . W i l d l . Manage. 44:963-968. Maynard, L.A., and J.K. L o o s l i . 1969. Animal N u t r i t i o n . 6th ed. McGraw-Hill Inc. pp 30-33. -98-McLellan, B.N. 1985. U s e - a v a i l a b i l i t y analysis and timber s e l e c t i o n by g r i z z l y bears. G r i z z l y Bear Habitat Symposium, Missoula, MT. McNicol, J.G., and F.F. G i l b e r t . 1980. Late winter use of upland cutovers by moose. J . Wildl. Manage. 44:363-371 Mendenhall, W. 1971. Introduction to p r o b a b i l i t y and s t a t i s t i c s , 3rd ed. Duxbury Press, Belmont, C a l i f . Murphy, D.A., and J . A. Coates. 1966. E f f e c t s of dietary protein on deer. 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The value and c h a r a c t e r i s t i c s of resident hunting. B.C. Ministry of Environment p u b l i c a t i o n . Renecker, L.A., and R.J. Hudson. 1985. Estimation of dry matter intake of free-ranging moose. J . W i l d l . Manage. 49:785-792. Renecker, L.A., and R.J. Hudson. 1986. Seasonal energy expenditures and themoregulatory responses of moose. Can. J . Zool. 64:322-327. Roscoe, J.T., and J.A. Byars. 1971. An inspection of the r e s t r a i n t s with respect to sample siz e commonly imposed on the use of the chi-square s t a t i s t i c . In Byers and Steinhorst (1984) . -99-Schwab, F.E. 1985. Moose habitat selection i n r e l a t i o n to for e s t cutting practices i n Northcentral B r i t i s h Columbia. Ph.D. th e s i s , University of B r i t i s h Columbia. Schwartz, C.C., W.L. Regelin, and W.W. Franzmann. 1987. Protein digestion i n moose. J . Wildl. Manage. 51:352-357. Spencer, D.L., and J.B. Hakala. 1964. Moose and f i r e on the Kenai. Proc. 3rd Ann. T a l l Timbers F i r e Ecology Conf. T e l f e r , E.S. 1970. Winter habitat s e l e c t i o n by moose and white-tailed deer. J. Wildl. Manage. 34:553-559. T e l f e r , E.S. 1978. Habitat requirements of moose - the p r i n c i p a l taiga range animal. Proc. 1st Int. Rangeland Congr. 462-265. Van Ballenberghe, V., and J.M. Peek. 1971. Radiotelemetry studies of moose i n northeastern Minnesota. J . W i l d l . Manage. 35:63-71. Walmsley, M., 6. Utzig, T. Void, D. Moon, and J . van Barneveld. (eds.). 1980. Describing ecosystems i n the f i e l d . Res. Anal. Br. Tech. Paper 2, Land Manage. Rep. No. 7, B.C. Ministry of Environment, B.C.Ministry of Forests, V i c t o r i a , B.C. -100-

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