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Habitat selection and use in winter by moose in sub-boreal forests of north-central British Columbia,.. 1977

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HABITAT SELECTION AND USE IN WINTER BY MOOSE IN SUB-BOREAL FORESTS OF NORTH-CENTRAL BRITISH COLUMBIA, AND RELATIONSHIPS TO FORESTRY by DONALD SIDNEY EASTMAN B. S c , U n i v e r s i t y of B r i t i s h Columbia 1962 M.Sc, U n i v e r s i t y of Aberdeen 1964 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY i n THE FACULTY OF GRADUATE STUDIES Department of P l a n t Science We accept t h i s t h e s i s -as conforming to the re q u i r e d standard THE UNIVERSITY OF BRITISH COLUMBIA December 1977 (c) Donald Sidney Eastman In p r e s e n t i n g t h i s t h e s i s i n p a r t i a l f u l f i l l m e n t of the requirements f o r an advanced degree at the U n i v e r s i t y of B r i t i s h Columbia, I agree th a t the L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r reference and study. I f u r t h e r agree that permission f o r extensive copying of t h i s t h e s i s f o r s c h o l a r l y purposes may be granted by the Head of my Department or by h i s r e p r e s e n t a t i v e s . I t i s understood that copying or p u b l i c a t i o n of t h i s t h e s i s f o r f i n a n c i a l gain s h a l l not be allowed without my w r i t t e n permission. Department of P l a n t Science The U n i v e r s i t y of B r i t i s h Columbia 2075 Wesbrook Place Vancouver, B.C., Canada V6T 1W5 Date Chairman: Dr. V. C. Brink ABSTRACT A study of wi n t e r h a b i t a t s e l e c t i o n and use by moose 2 was conducted i n a 11,300 km area of n o r t h - c e n t r a l B r i t i s h Columbia from May 1971 to August 19 73. The study area was lo c a t e d w i t h i n the f o r e s t e d sub-boreal spruce b i o g e o c l i m a t i c zone, a zone that i s r e c e i v i n g increased development, e s p e c i a l l y by f o r e s t r y . H a b i t a t s e l e c t i o n and use was examined mainly be p e l l e t group surveys and a e r i a l t r a n s e c t s . Wintering moose used p a r t i a l cutovers and burns more than coniferous f o r e s t s ; deciduous f o r e s t s and recent c l e a r c u t s were used l e a s t . L i m i t e d data suggested a s i m i l a r p a t t e r n i n summer. Winter use t y p i c a l l y increased from near zero a f t e r a recent disturbance such as c l e a r c u t t i n g , to a peak sometime between 10 and 25 years l a t e r , then d e c l i n e d to low l e v e l s during 25 and 9 0 years, and then apparently s t a b i l i z e d i n the mature f o r e s t stage at s l i g h t l y higher l e v e l s . On one i n t e n s i v e l y surveyed area, moose s e l e c t e d p a r t i a l cutovers and creek bottoms even though these h a b i t a t s comprised l e s s than 6 percent of the area. Moose began concentrating on winter ranges at l e a s t by mid- November, reached a peak i n November-January, and d e c l i n e d s t e a d i l y t h e r e a f t e r . Food h a b i t s and d i e t were examined by I l l rumen a n a l y s i s , t r a i l i n g and post-winter browse surveys. Moose had c a t h o l i c d i e t s but ate p r i m a r i l y deciduous browse f o r most of the year. Subalpine f i r becomes important i n l a t e w i n t e r . D i e t v a r i e d according to season and h a b i t a t . P r e f e r r e d species t y p i c a l l y were l e a s t common. Tagged twig t r a n s e c t s revealed that moose f r e q u e n t l y browsed p l a n t s more than once but r a r e l y re-browsed a twig. The time of browsing v a r i e d by species and by h a b i t a t w i t h most use recorded i n January and i n A p r i l . Levels of u t i l i z a t i o n were a l l l e s s than 100 percent of the previous y e a r 1 s production. U t i l i z a t i o n (weight-basis) ranged from 33 per- cent on r e d - o s i e r dogwood to 3 percent on subalpine f i r ; and from tr a c e amounts i n an upland burn h a b i t a t to more than 40 percent i n deciduous f o r e s t , p a r t i a l cutover and r i v e r bottom h a b i t a t s . Bedding .habits were examined i n an attempt to define cover f o r moose. Moose choose upper slopes t h a t faced south p a r t i c u l a r l y when snow depths became r e s t r i c t i v e (> 8 0 cm). Moose tended to s e l e c t l a r g e r than average trees and to bed on the southerly sides of them. S e l e c t i o n of bed s i t e s v a r i e d w i t h snow depth. As snow became deeper, moose bedded c l o s e r to l a r g e r t r e e s i n the denser canopied p a r t s of f o r e s t stands. Moose showed greater s e l e c t i o n f o r prot e c t e d s i t e s as w i n t e r c o n d i t i o n s became more severe. Secondary s e r a i succession was examined w i t h respect to s e v e r a l a t t r i b u t e s f o r mesic environments on the two commonest s u b s t r a t e s , g l a c i a l t i l l and l a c u s t r i n e deposits. i v F l o r i s t i c s of s e r a i stages from 1-200 years revealed t h a t on l a c u s t r i n e s o i l s , vegetation was more, diverse and the deciduous phase was prolonged. Species d i v e r s i t y d e c l i n e d around year 25 on t i l l but not on l a c u s t r i n e . S e v e r a l major changes occurred i n the tree l a y e r : f i r s t , a deciduous t r e e l a y e r developed e s p e c i a l l y on l a c u s t r i n e s o i l s ; second, a f t e r 25 years on t i l l (45 years or more on l a c u s t r i n e ) , lodgepole pine became most abundant; t h i r d , pine was g r a d u a l l y replaced by white spruce a f t e r 150-200 years or more; f o u r t h , subalpine f i r would probably become the domi- nant t r e e species i n the absence of f i r e . Understory phytomass, though c o n t r i b u t i n g l i t t l e to the mature f o r e s t mass, incre a s e d d r a m a t i c a l l y . t o peaks e a r l y i n succession and then remained low. Approximate net primary production of the understory on t i l l was greatest at age 11 w i t h 133 2 2 g/m / y r produced and l e a s t at age 39 w i t h 18 g/m /yr produced. Understory production i n the mature f o r e s t was an 2 estimated 27 g/m / y r . The shrubs c o n t r i b u t e d 70 percent, 26 percent, 44 percent, and 26 percent of annual production at ages 1, 11, 39, and 195 years, post-disturbance. Crude p r o t e i n and l i g n i n values were determined f o r 10 species (eight shrubs, one c o n i f e r , one l i c h e n ) f o r an annual c y c l e . Crude p r o t e i n averaged 7 percent.and l i g n i n , 9.8 percent. Crude p r o t e i n increased abruptly from steady winter values to peaks of 10-15 percent i n June-July and then returned to low l e v e l s by October. Leaf p r o t e i n was higher than, and V p r e d i c t a b l e from, stem l e v e l s . Crude p r o t e i n v a r i e d by- species, sometimes by substrate and r a r e l y by h a b i t a t - t y p e , at l e a s t f o r the species analyzed. The l i c h e n , lungwort, r e t a i n e d a high p r o t e i n value of approximately 11 percent throughout the year. L i g n i n l e v e l s v a r i e d s e a s o n a l l y , though l e s s d r a m a t i c a l l y than crude p r o t e i n . Levels were a f f e c t e d by species , substrate and age of s e r a i stage. P r o t e i n l e v e l s were s i m i l a r t o those reported i n the l i t e r a t u r e . Factors i n f l u e n c i n g crude p r o t e i n were d i f f i c u l t to disentangle due to confounding. Winter climate was s t u d i e d w i t h respect to d i f f e r e n c e s i n snow features between h a b i t a t s . Moose moved i n t o w i n t e r ranges before snow depths were l i m i t i n g . This i n d i c a t e s snow acts to t r i g g e r migration. On wi n t e r ranges, moose a l s o moved i n t o f o r e s t e d h a b i t a t s i n mid-winter (January) when snow depths approached 80 cm. Snow depths and d e n s i t i e s v a r i e d between h a b i t a t s . Snow cover was more v a r i a b l e i n p a r t i a l l y logged cutovers than i n the open or f o r e s t e d stands. The climate of f o r e s t , ecotone and adjacent open areas were documented. Compared t o adjacent open areas, the f o r e s t had higher r e l a t i v e humidity, l e s s wind, more moderate temperatures and approximately 50 percent of the snow depth. The t r a n s i t i o n zone from open to f o r e s t climates appeared to be r e l a t i v e l y narrow, l e s s than 50 m. The r e l a t i o n s h i p between c a r r y i n g c a p a c i t y , h a b i t a t s e l e c t i o n and home range are discussed w i t h reference to moose and management of t h e i r h a b i t a t . Management recommendations and suggestions f o r f u t u r e research are provided. TABLE OF CONTENTS Page ABSTRACT i i TABLE OF CONTENTS v i i LIST OF TABLES x i i i LIST OF FIGURES AND ILLUSTRATIONS xx LIST OF APPENDICES x x i v ACKNOWLEDGEMENTS x x i x 1. INTRODUCTION 1 1.1 The Study 1 1.2 The Approach to the Study 6 1.3 The Need f o r Integrated Management 8 1.4 A Land Use Pe r s p e c t i v e . 11 1.4.1 General I n t r o d u c t i o n 11 1.4.2 Mining 15 1.4.3 A g r i c u l t u r e . 18 1.4.4 F o r e s t r y 23 1.4.5 W i l d f i r e 29 2. THE STUDY AREAS 34 2.1 B i o p h y s i c a l S e t t i n g 34 2.2 The Primary Study Areas 52 2.3 The Secondary Study Areas 68 2.4 Moose D i s t r i b u t i o n and abundance 70 v i i v i i i 3. HABITAT USE AND SELECTION 73 3.1 I n t r o d u c t i o n 7 3 3.2 Methods • 77 3.2.1 The Synoptic Survey 77 3.2.2 P e l l e t Group Counting Methods f o r D e t a i l e d Survey. 80 3.2.3 The A e r i a l Surveys 81 3.3 Results 84 3. 3.1 Habitat Use 84 3.3.2 Habitat S e l e c t i o n 9 7 3.3.3 The Timing of M i g r a t i o n and Occupany Periods 9 9 3.4 Discussion 102 3.4.1 The Importance of Habitat V a r i a b i l i t y . 102 4. FOOD HABITS 109 4.1 I n t r o d u c t i o n 109 4.2 Methods 110 4.2.1 Rumen A n a l y s i s 110 4.2.2 T r a i l i n g 117 4.2.3 The Browsed Stem Survey 118 4.3 Results 118 4.3.1 The Range of Species Taken 118 4.3.2 The Seasonal Trends 12 0 4.3.3 The E f f e c t of Habitat-Type on D i e t 125 4.4 D i s c u s s i o n 130 4.4.1 Methodology 130 i x Page 4.4.2 V a r i a t i o n s i n the Diet 131 4.4.3 Some Management I m p l i c a t i o n s of V a r i a t i o n s i n the Die t 133 4.4.4 Future Research. . . . 135 5. THE DYNAMICS OF WINTER BROWSING 137 5.1 I n t r o d u c t i o n 137 5.2 Methods 138 5.3 Results 143 5.3.1 The Incidence of Use 14 3 5.3.2 The Time of Use 149 5.3.3 The L e v e l of Use 152 6. BED SITE SELECTION BY MOOSE IN WINTER 15 7 6.1 I n t r o d u c t i o n 15 7 6.2 Methods 15 9 6.3 Results 162 6.4 Discussion 179 7. SECONDARY SUCCESSION IN SUB-BOREAL FORESTS 184 7.1 I n t r o d u c t i o n 184 7.2 Methods 18 8 7.2.1 S t r a t i f i c a t i o n 188 7.2.2 F i e l d Sampling Procedures 190 7.2.3 The P r e d i c t i o n of Mass and Height of Woody P l a n t s 198 7.2.4 Date-Analysis 207 7.3 Results f o r Mesic Upland S i t e s 208 7.3.1 The Data Base 20 8 X Page 7.3.2 F l o r i s t i c Changes i n S e r a i Succession 210 7.3.3 Temporal Dynamics of the Tree Layer 237 7.3.4 Phytomass, Height and Basal Area of the Shrub Layer i n S e r a i P l a n t Communities 246 7.3.4.1 For the combination of species 246 7.3.4.2 Trends i n phytomass and height of food species 251 7.3.5 Phytomass of the Herb Layer i n S e r a i P l a n t Communities 254 7.3.6 Net Primary P r o d u c t i v i t y of the Understory 258 7.4 Results f o r R i p a r i a n S i t e s 264 7.5 Discussion 269 7.5.1 P r e d i c t i n g Successional Development. 269 7.5.2 Trends i n Production of Cover and Food 2 76 8. NUTRITIVE ASPECTS OF MOOSE FORAGES 2 85 8.1 I n t r o d u c t i o n 2 85 8.2 Methods 288 8.3 Crude P r o t e i n Levels 291 8.4 L i g n i n Levels 30 8 8.5 Discussion 314 8.5.1 Crude P r o t e i n Levels i n Moose Forages 314 x i Page 8.5.2 Assessing the. N u t r i t i v e Values of Forages 318 8.5.3 Factors A f f e c t i n g N u t r i e n t Levels 322 9. EFFECTS OF FORESTS ON. WINTER CLIMATE 333 9.1 I n t r o d u c t i o n 333 9.2 Methods 335 9.2.1 The Estima t i o n . o f M i g r a t i o n and Winter Range Occupancy . 335 9.2.2 Snow C h a r a c t e r i s t i c s of Habitat-Types 336 9.2.3 Climate of the Forest Edge 340 9. 3 Results 341 9.3.1 Mi g r a t i o n and Snow Accumulation of Habitat-Types 341 9.3.2 Snow C h a r a c t e r i s t i c s of Habitat-Types 345 9.3.3 Climate of the Forest Edge 355 9.4 Discussion 365 9.4.1 The Role of Snow Pack i n I n i t i a t i n g M i g r a t i o n 365 9.4.2 The Role of Climate i n D i f f e r e n t i a l Use Between H a b i t a t s . . . . 367 10. DISCUSSION 379 10.1 Habitat R e l a t i o n s h i p s i n Moose Management 379 10.2 The E f f e c t s of Timber Management on Moose Hab i t a t . 39 4 10.2.1 F e l l i n g 397 10.2.2 S i t e P r e p a r a t i o n 402 x i i Page 10.2.3 Stand Establishment 405 10.2.4 Stand Tending . 408 10.2.5 Stand P r o t e c t i o n 412 10.2.6 General Management Considerations. 414 10.3 Overview and Recommendations 416 11. LITERATURE CITED. . 42 7 12. APPENDICES 460 VITA LIST OF TABLES TABLE Page 1.1 Population Growth and Future P r o j e c t i o n s f o r P r i n c e George and the Surrounding D i s t r i c t 14 1.2 Major Events i n the. Settlement and Growth of P r i n c e George and the Surrounding Region 16 1.3 Number and Area of. Farms, and Numbers of C a t t l e f o r the Province and f o r the Pr i n c e George Region, 1881-1971 20 1.4 Trends i n Logging Methods and Area Cut i n the P r i n c e George Forest D i s t r i c t , 1950-1973 . . . 27 1.5 Estimated Areas of Broad Vegetation Classes i n F i v e Major Drainages i n North-Central B r i t i s h Columbia (from Whitford and C r a i g 1918) 31 2.1 C l i m a t i c Parameters f o r the Study Area 40 2.2 Major S o i l A s s o c i a t i o n s f o r the Study Area and Their R e l a t i o n s h i p to Parent M a t e r i a l s and Moisture Regimes 44 2.3 Estimated R e l a t i v e Abundance and Herd S t r u c t u r e f o r Wintering Moose on the Int e n s i v e Study Areas,, 1964-65 to 1975-76 58 2.4 Types of Analyses Conducted on the Study Area.. . . 69 3.1 Results from T r i a l P e l l e t Group Survey: Time/Plot and Number of Group/Plot 7 8 3.2 R e l a t i v e Winter Use of A v a i l a b l e H abitats on S e l e c t e d Study Areas, Based on P e l l e t Group Surveys 8 7 3.3 R e l a t i v e Winter Use of Major Habitat-Types i n the Sub-Boreal F o r e s t , Based on P e l l e t Group Surveys 89 x i i i x i v TABLE Page 3.4 Winter Use of Ecotones Between Forests and V a r i o u s l y Aged S e r a i Stages, Based on P e l l e t Group Surveys 9 2 3.5 Winter U t i l i z a t i o n of Roads and Habitats i n which they were Located, Based on Accumulated P e l l e t Groups i n 1973 at McKenzie. . . 95 3.6 R e l a t i v e Summer Use of.Habitat-Types on Accumulated Summer Feces Recorded i n the 1973 Synoptic Survey. . . 9 6 3.7 D i s t r i b u t i o n of P e l l e t Group P l o t s According to Habitat-Type and the Number of Groups they Contained on the Intensive Salmon Area 98 3.8 S e l e c t i o n of Habitat-Types by Moose i n Winter as I n d i c a t e d by Accumulated P e l l e t Groups on the Intensive Salmon Area. . 98 4.1 Components of Rumen O l i g e s t a A f t e r Sample Pre p a r a t i o n 112 4.2 The E f f e c t of A n a l y t i c a l Method on Frequency of Occurrence of P l a n t Taxa Recorded i n Moose Rumen Samples 114 4.3 The E f f e c t of A n a l y t i c a l Method on Amounts of P l a n t Taxa I d e n t i f i e d i n Moose Rumen Samples . . . 116 4.4 V a r i e t y of P l a n t Species Eaten by Moose, by Forage C l a s s , i n Various P a r t s of Th e i r North American Range . . . 119 4.5 Food Habits of Moose, i n North-Central B r i t i s h Columbia, Based on T r a i l i n g and Rumen A n a l y s i s , 1971^-74 (%-basis) 122 4.6 Comparisons of Food Habits of Moose Between D i f f e r e n t H a b i t a t s i n E a r l y and Late Winter. . . . 126 4.7 Winter Food Preferences of Moose i n North- i Ce n t r a l B r i t i s h Columbia, by ̂ Habitat-Type 127 5.1 Proportions of Twigs that were Browsed Once and Twice i n Major Habitats on the Eagle, Grove and Salmon Winter Ranges During the 1972-73 Winter 144 X V TABLE Page 5.2 Number of Times P l a n t s of Subalpine F i r , Paper B i r c h , Red-osier Dogwood and Willow were Browsed on the Eagle, Grove and Salmon Study Areas during the 1972-73 Winter. 146 5.3 P r o p o r t i o n of P l a n t s Browsed.by Species and H a b i t a t , i n the 1972-73 Winter 148 5.4 Time of Browsing and L e v e l of U t i l i z a t i o n (Weight-Basis) f o r A l l Species, Habitat Types, Study Areas and Months 15 0 6.1 Major H a b i t a t s , Snow Depth Classes and Study Areas Sampled f o r Bed S i t e Examinations 160 6.2 Example of Data Sheets-Used to Study Bed S i t e s . . 161 6.3 Time Spent by Moose i n Beds as I n d i c a t e d by Feces and Urine, According to H a b i t a t and Month. 165 6.4 Locations of Moose.Beds w i t h Respect to P o s i t i o n on Slope, and Aspect 166 6.5 Comparison of C o n i f e r Species A v a i l a b l e as S h e l t e r Trees, With Those -Used by Moose 169 6.6 O r i e n t a t i o n of Moose i n Their Beds, and i n R e l a t i o n to the S h e l t e r Tree 171 6.7 Location of Beds i n Quamaniqs, as A f f e c t e d by Habitat and Snow Depth Class 178 6.8 Comparison of Snow Depths between Moose Bedding S i t e s and Adjacent Areas 179 7.1 Scale Used to Assess Canopy-Coverage of Understory Vegetation (Clayer) ( a f t e r Daubenmire 19 59), plus Domin Scale E q u i v a l e n t s 195 7.2 Summary of Features Sampled i n the Synoptic Study of Succession 197 7.3 The E f f e c t of S i t e on P r e d i c t i n g Mass f o r S e l e c t e d Shrub Species . . • 201 x v i TABLE Page 2 7.4 C o e f f i c i e n t s of Determination (r value) f o r S i x Independent V a r i a b l e s Used to P r e d i c t Phytomass of 19 Sub-Boreal Shrubs . . . . 203 7.5 Regression C o e f f i c i e n t s f o r P r e d i c t i n g Height from Diameter Measurements 2 05 7.6 Regression C o e f f i c i e n t s f o r P r e d i c i n g Oven-Dried, Above-Ground Phytomass of 19 Sub-Boreal Shrubs from Diameter, and from Diameter Squared by Length Measure- ments. A l l V a r i a b l e s Based.on Logarithmic Transformed Data 206 7.7 D i s t r i b u t i o n of Sampling S i t e s f o r the P l a n t Succession Study. . 209 7.8 P l a n t Community Names f o r Successional Stages on T i l l and L a c u s t r i n e Substrates 211 7.9 Percent Canopy-Coverage/Frequency of Occurrence Values f o r Major P l a n t Species of the Herb (C) Layer i n a Sub-Boreal Forest Sere i n a Mesic Environment on the T i l l Substrate. . . 213 7.10 Percent Canopy-Coverage/Frequency of Occurrence Values f o r Major P l a n t Species of the Herb (C) Layer i n a Sub-Boreal Forest Sere i n a Mesic Environment on the L a c u s t r i n e Substrate. . 215 7.11 Percent Canopy-Coverage/Frequency of Occurrence Values f o r Major P l a n t Species of the Herb (C) Layer i n P a r t i a l l y Logged Sub-Boreal Forest Stands i n a Mesic Environment on T i l l and L a c u s t r i n e Substrates . .217 7.12 Percent Species Composition (Stem-Basis) of the Shrub (B) Layer i n a Sub-Boreal Forest Sere i n a Mesic Environment Over T i l l Substrates 218 7.13 Percent Species Composition (Stem-Basis) of the Shrub (B) Layer i n a Sub-Boreal Forest Sere i n a Mesic Environment Over L a c u s t r i n e Substrates 219 x v i i TABLE x Page 7.14 Percent Species Composition (Stem-Basis) of the Shrub (B) Layer i n P a r t i a l l y Logged Stands of Sub-Boreal Forests i n a Mesic Environment 220 7.15 Temporal Changes i n Tree Species Composition f o r Mesic Sub-Boreal Forests on T i l l and L a c u s t r i n e Substrates 238 7.16 Temporal Trends f o r Coniferous Regeneration i n a Mesic Environment Over T i l l and L a c u s t r i n e Substrates 240 7.17 Temporal Changes i n Composition and Proportions of Dead Trees i n Mesic Sub- Bor e a l Forest Stands on T i l l and L a c u s t r i n e Substrates 242 7.18 Temporal Changes i n Basal Area, Canopy Closure, and Height of Dominant Trees i n Mesic Sub-Boreal Forests on T i l l and L a c u s t r i n e Substrates 244 7.19 S t a t i s t i c s (mean = sd) f o r the Shrub (B) Layer i n Mesic Sub-Boreal Seres Over T i l l and L a c u s t r i n e Substrates 247 7.2 0 Trends i n Phytomass of the Herb (C) Layer f o r Seres i n Mesic Sub-Boreal Forests on T i l l and L a c u s t r i n e Substrates 256 7.21 Approximate Net Primary P r o d u c t i v i t y of Understory Vegetation (Layers B and C) at Four Successional Stages on T i l l Substrate. . . . 260 7.22 Changes i n Proportions of P l a n t Components of S e lected Shrub Species a t Four Successional Stages on T i l l Substrates 264 7.2 3 Major Features of S e r a i Stages i n Forest Succession on R i p a r i a n ( A l l u v i a l ) H a b itats (Adapted from Sumanik (1968) and Waring (1970) ) 267 8.1 L o c a t i o n , H a b i t a t , Substrate, and Species C o l l e c t e d f o r Crude P r o t e i n and L i g n i n Analyses 2 89 x v i i i TABLE Page 8.2 Estimates of Experimental E r r o r i n P r o t e i n Analyses f o r Selected Species.. 291 8.3 Crude P r o t e i n and L i g n i n Levels i n Major Moose Forages Averaged Over an Annual Cycle (May 1972 - A p r i l 1973) 292 8.4 Crude P r o t e i n and L i g n i n Levels i n the Current Year's Stems and Leaves of Se l e c t e d Browse Species 295 8.5 Proportions of Stem and Leaf Tissue i n Current Annual Growth of S e l e c t e d Browse Species C o l l e c t e d i n September 19 72 297 8.6 Comparison of P r o t e i n and L i g n i n Contents of Willow and Paper B i r c h C o l l e c t e d from T i l l and L a c u s t r i n e Substrates of the Burn H a b i t a t at the Grove Study Area 299 8.7 E f f e c t of Habitat on Crude P r o t e i n Levels i n S e l ected Browse Species at the Eagle and Salmon Study Areas . . . . . 303 8.8 Year-to-Year V a r i a t i o n s i n the Content of Crude P r o t e i n and L i g n i n 307 8.9 D i f f e r e n c e i n Percent L i g n i n Content as A f f e c t e d by Substrate, H a b i t a t and Stand Age 312 8.10 Comparison of Crude P r o t e i n Values f o r Current Annual Growth of Common Winter Foods of Moose (November-March) 316 9.1 L o c a t i o n , S i t e Number, E l e v a t i o n , and Ha b i t a t of Snow Courses. . 337 9.2 Annual V a r i a t i o n s i n Snow Depths i n Open or Deciduous Forest Habitats at Eagle, Grove and Salmon Winter Ranges f o r March 1972, 1973, and 1974 346 9.3 Monthly Snow Depths and D e n s i t i e s f o r Three H a b i t a t s on the Eagle Winter Range 34 8 9.4 Monthly Snow Depths f o r Four Habitats on the Grove Winter Range 349 x i x TABLE Page 9.5 Monthly Snow D e n s i t i e s f o r Four Habitats on the Grove Winter Range. 350 9.6 Monthly Snow Depths, f o r F i v e Habitats at the Salmon Winter Range. 351 9.7 Monthly Snow D e n s i t i e s . f o r F i v e Habitats at the Salmon Winter Range 352 9.8 Comparison of Sel e c t e d C l i m a t i c Parameters Between the South-Facing Ecotone at the Grove Study Area ( S t a t i o n at 76 m i n the Open) and Pr i n c e George. A i r p o r t . 357 9.9 Mean Monthly Temperature, R e l a t i v e Humidity, Snow Pack, and Wind f o r C l e a r c u t and Adjacent Forest S i t e s at the Bowron Study Area, 1972-73 Data 359 9.10 Mean Monthly Depth, Density, and Penetrance of Snow Across the South-Facing, Fo r e s t - Burn Ecotone at the Grove Study Area, 1972-73 Winter 363 10.1 General Features of B a s i c Resources Required by Moose 395 10.2 S o i l Disturbance and Slash Accumulations R e s u l t i n g from D i f f e r e n t Types of Logging i n Western North America (Derived from Bockheim e t a l . 1975) 400 LIST OF FIGURES FIGURE Page 1.1 The general r e l a t i o n s h i p of sub-models tha t comprise a moose-forest model. Derived from Haagenrud and H j e l j o r d (1976) and Houston (1968) 4 1.2 Development of f o r e s t r y i n the P r i n c e George Forest D i s t r i c t as i n d i c a t e d by the annual cut and the number of operating sawmills, 1914-1974 25 2 1.3 Annual area (km ) burned by w i l d f i r e i n the P r i n c e George Forest D i s t r i c t , 1910-1975 32 2.1 Locations of the study areas, and of place names mentioned i n the t e x t 35 2.2 Longterm monthly averages of some temperature and p r e c i p i t a t i o n para- meters f o r the P r i n c e George weather s t a t i o n 38 2.3 Oblique a e r i a l photographs i l l u s t r a t i n g the general t e r r a i n and vegetation of the P r i n c e George study area 43 2.4 A schematic i l l u s t r a t i o n of the major s o i l a s s o c i a t i o n s i n the study area, and t h e i r topographic r e l a t i o n s h i p to each other 45 2.5 Photographs of the Eagle, Grove and Salmon w i n t e r ranges 5 3 2.6 A s o i l a s s o c i a t i o n map of the Eagle study area 55 2.7 A f o r e s t cover map of the Eagle study area 56 2.8 A s o i l a s s o c i a t i o n map of the Grove study area 61 xx xx i FIGURE Page 2.9 A f o r e s t cover map of the Grove study area 6 3 2.10 A s o i l a s s o c i a t i o n map of the Salmon study area 65 2.11 A f o r e s t cover map of the Salmon study area 6 7 3.1 Photographs i l l u s t r a t i n g logged h a b i t a t s i n sub-boreal f o r e s t s : a) s e l e c t i v e , b) cut and leave, and c) c l e a r c u t 76- 3.2 The r e l a t i o n s h i p between d a i l y s n o w f a l l and the timing of the a e r i a l t r a n s e c t surveys, January 19 72 to May 19 7 3 8 3 3.3 A map showing f l i g h t l i n e s used f o r the a e r i a l t r a n s e c t surveys on the Grove study area 85 3.4 R e l a t i v e use by moose of ecotones and adjacent h a b i t a t s , based on p e l l e t group t r a n s e c t s 9 3 3.5 Number of moose seen/minute of f l y i n g on the Eagle, Grove and Salmon study areas during a e r i a l t r a n s e c t surveys i n the 1972-73 winter 101 4.1 The seasonal changes i n forage c l a s s e s eaten by moose i n n o r t h - c e n t r a l B r i t i s h Columbia 124 5.1 Photographs i l l u s t r a t i n g the methods of tagging twigs and measuring diameter at p o i n t of browsing 141 6.1 The r e l a t i o n s h i p between snow depth and the length of time moose spent i n beds, as i n d i c a t e d by r e l a t i v e amounts of feces and urine 163 6.2 The r e l a t i o n s h i p between snow depth and the distance between bed s i t e s and t h e i r a s s o c i a t e d s h e l t e r t r e e s 174 x x i i FIGURE Page 6.3 The r e l a t i o n s h i p between snow depth and the d i f f e r e n c e i n crown cl o s u r e between a bed s i t e and the f o r e s t stand i n which i t was l o c a t e d . . . . 176 6.4 An i l l u s t r a t i o n of e f f e c t i v e snow i n t e r c e p t i o n by the f o r e s t canopy 177 7.1 The s i t e and s t a t i o n l a y o u t used to study secondary p l a n t succession 192 7.2 Photographs i l l u s t r a t i n g s e l e c t e d s u c c e s s i o n a l stages on mesic t i l l and l a c u s t r i n e substrates 221 7.3 Some trends i n the f o r e s t stand features of b a s a l area, dominant tree height, and canopy c l o s u r e i n sub-boreal f o r e s t seres on t i l l and l a c u s t r i n e substrates 245 7.4 Trends i n height and mass of browse and non-browse species i n sub-boreal f o r e s t succession on t i l l and l a c u s t r i n e substrates 252 7.5 Percentage composition, by forage c l a s s , of phytomass i n the "C" or Herb l a y e r at your s u c c e s s i o n a l stages of the sub- bo r e a l f o r e s t on t i l l and l a c u s t r i n e substrates 255 7.6 Percentage composition, by forage c l a s s , of the net primary production of the understory vegetation (l a y e r s B and C) at four s u c c e s s i o n a l stages of the sub- bo r e a l f o r e s t on the t i l l substrate 259 7.7 Percentage composition, by species, of the net primary production of the shrubs ( l a y e r s B and C) at four s u c c e s s i o n a l stages of the sub-boreal f o r e s t on the t i l l s ubstrate 262 7.8 Photographs i l l u s t r a t i n g p l a n t s u c c e s s i o n a l stages on r i p a r i a n ( a l l u v i a l ) substrates 268 8.1 Crude p r o t e i n l e v e l s i n major p l a n t species eaten by moose i n sub-boreal f o r e s t s 293 x x i i i FIGURE Page 8.2 Comparisons of crude p r o t e i n l e v e l s i n w i l l o w and subalpine f i r growing on s i m i l a r substrates but i n stands of d i f f e r e n t ages 301 8.3 Comparisons of crude p r o t e i n l e v e l s i n d i f f e r e n t species growing at the same s i t e s 304 8.4 Consistency i n crude p r o t e i n l e v e l s between aspen and w i l l o w f o r three d i f f e r e n t s i t e s 306 8.5 L i g n i n l e v e l s i n major shrub species eaten by moose i n sub-boreal f o r e s t s 310 8.6 E f f e c t of s i t e on l i g n i n l e v e l s i n s e l e c t e d sub-boreal shrubs 313 9.1 Photographs showing the use of the western snow sampler and the penetrometer. . . . 338 9.2 Patterns of snow accumulation and snow melt f o r the Eagle, Grove and Salmon study areas, and f o r weather s t a t i o n s at P r i n c e George and A l e z a Lake 342 9.3 Some temperature and r e l a t i v e humidity gradients across the forest-open ecotone at the Grove study area during the 1972-73 winter 358 9.4 Wind run and snow depths across the forest-open ecotone at the Grove study area during the 1972-73 winter 361 10.1 Major f a c t o r s and how they i n t e r - r e l a t e moose population l e v e l s (modified from Houston 1968) 384 LIST OF APPENDICES Page APPENDIX A. SCIENTIFIC AND COMMON NAMES OF PLANT SPECIES RECORDED IN THE STUDY AREA 4 60 APPENDIX B. SCIENTIFIC AND COMMON -NAMES OF BIRD AND MAMMAL SPECIES MENTIONED FOR THE STUDY AREA 467 APPENDIX C. STATISTICAL DATA USED FOR THE INTRODUCTION (SECTION 1) 469 , - Table C - l . Estimated Annual Economic Value of Minerals Produced.in the Omineca Mining D i s t r i c t , 1926-1974. . . . . 470 • . •• Table C-2. Number and T o t a l Area of Farms, and Number of C a t t l e f o r the Province and f o r the Pr i n c e George Region, 1881-1971 471 Table C-3. Annual Cut of Timber ( A l l Species) and the Sawmills Operating i n the P r i n c e George Forest D i s t r i c t , 1909-1975 472 Table C-4. Area of Forest Land Disturbed by W i l d f i r e s and by Logging i n the P r i n c e George Forest D i s t r i c t , 1912-1975 473 APPENDIX D. - HABITAT USE AND SELECTION DATA (SECTION 3) 474 Table D-l. I n d i v i d u a l P l o t Data on Time and Number of Accumulated P e l l e t Groups Counted in- the A p r i l , 19 72 T r i a l Used to Determine the P e l l e t Group Survey Method 4 75 Table D-2. Example of the Recording Format Used f o r the A e r i a l Transects of the Intensive Study Areas, and the Type of Data Recorded 477 x x i v X X V Page Table D-3. Example of the Summary Derived from A e r i a l Transect Data . 479 APPENDIX E. Table D-4. Data from P e l l e t Group Transects f o r Synoptic Surveys i n 19 72. Table D-5. Data from P e l l e t Group Transects f o r Synoptic Surveys i n 1973. CHARACTERISTICS OF SAMPLES COLLECTED FOR THE FOOD HABITS STUDY (SECTION 4) . Table E - l . Rumen Samples: Date of K i l l , Sex, Age,- and Location of K i l l . . APPENDIX F. SUCCESSION DATA (FOR SECTION 7) Table F - l . Percentage Canopy-Coverage/ Frequency of Occurrence Values of Major* P l a n t Species Recorded i n the Herb Layer at the Succession Study S i t e s , MF2-MF7. . 491 Table F-2. Percentage Canopy-Coverage/ Frequency of Occurrence Values of Major* P l a n t Species Recorded i n the Herb Layer at the Succession Study S i t e s , MF8-MF13 . . 494 Table F-3. Percentage Canopy-Coverage/ Frequency of Occurrence Values of Major* P l a n t Species Recorded i n the Herb Layer at the Succession Study S i t e s , MF14-MF19. . 497 Table F-4. Percentage Canopy-Coverage/ Frequency of Occurrence Values of Major* P l a n t Species Recorded i n the Herb Layer at the Succession Study S i t e s , MF20-MF22. . 500 Table F-5. Percentage Canopy-Coverage/ Frequency of Occurrence Values of Major* P l a n t Species Recorded i n the Herb Layer at the Succession Study S i t e s , SR1-SR6. . . 503 Table F-6. Percentage Canopy-Coverage/ Frequency of Occurrence Values of Major* P l a n t Species Recorded i n the Herb Layer at the Succession Study S i t e s , SR7-SR12 . . 506 x x v i Page Table F-7. Percentage Canopy-Coverage/ Frequency of Occurrence Values of Major* P l a n t Species Recorded i n the Herb Layer at the Succession Study S i t e s , SR13-SR18. . 509 Table F-8. Percentage Canopy-Coverage/ Frequency of Occurrence Values of Major* P l a n t Species Recorded i n the Herb Layer at the Succession Study S i t e s , SR19-SR23. . 512 2 Table F-9. Phytomass (g/m , Oven-Dried Basis) of the Shrub Layer, by Species, i n a Sub-Boreal Forest Sere i n a Mesic Environment on T i l l Substrates 515 2 Table F-10. Phytomass (g/m , Oven-Dried Basis) of the Shrub Layer, by Species, i n a Sub-Boreal Forest Sere i n a Mesic Environment on L a c u s t r i n e Substrates. . . . 516 2 Table F - l l . Phytomass (g/m , Oven-Dried Basis) of the Shrub Layer, by Species, i n P a r t i a l l y Logged Sub-Boreal Forests i n a Mesic Environment on T i l l and L a c u s t r i n e Substrates 517 2 2 Table F-12. Basal Area (cm /m ) of the Shrub Layer, by Species, i n a Sub-Boreal Forest Sere i n a Mesic Environment on T i l l Substrates 518 2 2 Table F-13. Basal Area (cm /m ) of the Shrub Layer, by Species, i n a Sub- Boreal Forest Sere i n a Mesic Environment on L a c u s t r i n e Substrates 520 2 2 Table F-14. Basal Area (cm /m ) of the Shrub Layer, by Species, i n P a r t i a l l y Logged Sub-Boreal Forests i n a Mesic Environment on T i l l and L a c u s t r i n e Substrates 521 Table F-15. Height (cm) of the Shrub Layer, by Species, i n a Sub-Boreal Forest Sere i n a Mesic Environment on T i l l Substrates 522 x x v i i Page Table F-16. Height (cm) of the Shrub Layer, by Species, i n a Sub-Boreal Forest Sere i n a Mesic Environment on L a c u s t r i n e Substrates 524 Table F-17. Height (cm) of the Shrub Layer, by Species, i n P a r t i a l l y Logged Sub-Boreal Forests i n a Mesic Environment on T i l l and L a c u s t r i n e Substrates 525 Table F-18. Number of Stems Sampled i n the Shrub Layer, by Species, i n a Mesic Sub-Boreal Forest Sere on T i l l Substrates 526 Table F-19. Number of Stems Sampled i n the Shrub Layer, by Species, i n a Mesic Sub-Boreal Forest Sere on L a c u s t r i n e Substrates 527 Table F-20. Number of Stems Sampled i n the Shrub Layer, by Species, i n P a r t i a l l y Logged, Mesic Sub-Boreal Forests on T i l l and L a c u s t r i n e Substrates 528 Table F-21. Oven-Dried Weights of Components of Major Shrub Species i n Sub-Boreal Forests 529 APPENDIX G. DATA FOR NUTRIENT CONTENTS OF SAMPLED PLANT SPECIES (SECTION 8) . . . 534 Table G-l. Crude P r o t e i n Levels (%) i n P l a n t Samples C o l l e c t e d from the P r i n c e George Study Area, A p r i l 1972 to A p r i l 1973 . . . ' 535 Table G-2. L i g n i n Values (%) i n P l a n t Samples C o l l e c t e d from the Pr i n c e George Study Area, A p r i l 1972 to A p r i l 1973 541 APPENDIX H. CLIMATIC DATA USED FOR SECTION 9 545 Table H - l . Penetrance Values (1-11 Scale) f o r Snow Hardness Estimates Across the South-Facing Ecotone at the Grove Area, 1973 546 x x v i i i Page Table H-2. Monthly Means f o r Temperature and R e l a t i v e Humidity Across the South- Facing Forest-Burn Ecotone at the Grove Study Area, 1972-73 Winter 547 Table H-3. Wind Run (km/day) Across the South-Facing Forest-Burn Ecotone at Grove Study Area and the Exposed Burn S i t e at Buckhorn 548 Table H-4. Snow Depths (cm) Across the West-Facing Ecotone at the Grove Study Area, 19,72-73 Winter 548 APPENDIX I. GLOSSARY OF TERMS AND ABBREVIATIONS USED IN THE TEXT 549 C ACKNOWLE DGEMENTS I t i s a pleasure to acknowledge the many persons who co n t r i b u t e d to t h i s p r o j e c t . Thanks go to my committee, Drs. P. J . Bandy, F. B u n n e l l , I . M. Cowan, V. C. Runeckles, J. H. G. Smith and K. Sumanik, f o r t h e i r advice and c r i t i c a l review of my work. To my su p e r v i s o r , Dr. V. C. "Bert" B r i n k , I express my g r a t i t u d e f o r h i s i n t e l l e c t , p a t i e n c e , p e r s p e c t i v e , diplomacy and honesty. I t was an e n r i c h i n g experience to be one of h i s graduate students. My study was financed, encouraged and f a c i l i t a t e d l a r g e l y by the B.C. F i s h and W i l d l i f e Branch. F i s h and W i l d l i f e personnel i n the Pr i n c e George region supported and a s s i s t e d me i n many ways. Ken Sumanik encouraged me to study the problem, and provided me w i t h the b e n e f i t of h i s experience and the w i t of h i s i n s i g h t . M i l t Warren was a mine of information. Conservation O f f i c e r s D. Adolph, B. Clapp, L. Cox, W. Richmond, D. Turner, G. Vincent introduced me to t h e i r d i s t r i c t s , c o l l e c t e d rumen samples, and acted as e x c e l l e n t guides and sources of info r m a t i o n . Thanks a l s o go to P. Brade, K. C h i l d , K. F u j i n o and R. Goodlad of the Prin c e George o f f i c e , and to many i n the V i c t o r i a o f f i c e f o r t h e i r v a rious c o n t r i b u t i o n s to my study. Members of the B.C. Forest Service provided much x x i x X X X u s e f u l i n f o r m a t i o n on f o r e s t r y r e l a t e d a f f a i r s . I wish to acknowledge C. P. Axhorn, J. B u l l e n , R. C l i f f o r d , D. G i l b e r t , E. Lemon, J . Revel, W. Young and the ranger s t a f f s at P r i n c e George, Hixon, Summit Lake and A l e z a Lake. I r e c eived valuable help from the Resource A n a l y s i s Branch of the Environment M i n i s t r y (formerly the B.C. Land Inventory). In p a r t i c u l a r , I thank Greg Cheeseman, A l Dawson, the l a t e A l Luckurst, Gary Runka and Jim van Barneveld. Forest companies i n the area generously provided maps, informat i o n and a s s i s t a n c e . I wish to acknowledge Rustad Brothers Lumber Company and t h e i r f o r e s t e r , Don Frood; Holger Thomsen; Northwood Pulp and Timber Company Ltd.; P r i n c e George Pulp and Paper Company L t d . ; and Weldwood of Canada Company L t d . The f o l l o w i n g people a s s i s t e d me i n d a t a - c o l l e c t i o n , both i n tedious l a b o r a t o r y analyses and i n mosquito-plagued, patience-demanding f i e l d work: Rick Bonar, Dave Dunbar, Chris Easthope, O l l i e F r i c k e , John K e l l y , Ben Koop, Margaret L a r k i n , Mike Masson, W i l l a Noble, Sharon R u s s e l l , E r i c Rutt, Rankin Smith, Don Stevenson and C h r i s Whyte. John K e l l y deserves s p e c i a l thanks both f o r h i s i n v a l u a b l e help and h i s f r i e n d s h i p . Other o r g a n i z a t i o n s who provided help: The College of New Caledonia and the Canada A g r i c u l t u r e Experimental Farm f o r l a b o r a t o r y space and f a c i l i t i e s ; the Canadian x x x i W i l d l i f e Service,, f o r s c h o l a r s h i p s . Other i n d i v i d u a l s I wish to acknowledge are: Les Bower, master of the Cessna 185; Richard Revel, who f i r s t d escribed the sub-boreal spruce zone to me; Ed T e l f e r , whose experience, common sense and ideas o f f e r e d example and s t i m u l a t i o n ; John Powell and Douglas Golding of the Canada Fo r e s t r y S e r v i c e , f o r t h e i r data and advice on f o r e s t c l i m a t o l o g y ; Jim Peek, f o r advice on snow measurements and s e v e r a l aspects of moose ecology; Rod S i l v e r , f o r u s e f u l d i s c u s s i o n s of what makes moose " t i c k " ; Rick E l l i s , f o r inform a t i o n and d i s c u s s i o n on p l a n t succession; and Ralph R i t c e y , f o r advice and comment on my ideas and w r i t i n g . For making the f i n a l copy of the t h e s i s so readable thanks go to Barbara Smith, who d i d an e x c e l l e n t job of ty p i n g ; and to Laura F r i i s , who competently prepared the graphs and f i g u r e s . As i n most f i e l d p r o j e c t s , many i n d i v i d u a l s and f a m i l i e s provided a human environment which complemented and enhanced my experiences while we l i v e d i n P r i n c e George. I wish to acknowledge i n p a r t i c u l a r the f o l l o w i n g f a m i l i e s and i n d i v i d u a l s : .the Clapps, Froods, Gagnons, Jaroschs, Manns, Pagets, John Sawitsky, Spurrs and Sumaniks. L a s t , but c e r t a i n l y not l e a s t , I wish to thank my f a m i l y : the support and forebearance of my w i f e , E l a i n e , and my two c h i l d r e n , Jenny and S t u a r t ; i n many respects t h i s t h e s i s i s as much t h e i r s as i t i s mine. To my parents, Ben and B e r n i c e , I owe an unpayable debt. My in-laws, Len and Mabel Weston, gave f r e e l y t h e i r understanding and support. 1. INTRODUCTION 1.1 The Study Moose are e l u s i v e , s o l i t a r y ungulates of the sub- b o r e a l f o r e s t s i n n o r t h - c e n t r a l B r i t i s h Columbia. They have been h i g h l y s u c c e s s f u l i n t h i s comparatively harsh environment. Their success i s probably due to three main c h a r a c t e r i s t i c s . F i r s t l y , moose are adapted to winters that are long, c o l d and snowy ( K e l s a l l 1969, K e l s a l l and T e l f e r 1971). Secondly, they are browsers i n an area where shrubs and trees form the major food resource, e s p e c i a l l y i n winter. F i n a l l y , they are a f i r e - or successionally-adapted species (Geist 1971) and so can c a p i t a l i z e e f f e c t i v e l y on the superabundance of forage produced i n the e a r l y stages of f o r e s t succession ( T e l f e r 1974). Since frequent f i r e s leave large p o r t i o n s of b o r e a l and sub-boreal f o r e s t s i n these e a r l y stages (Heinselman 1973 and o t h e r s ) , t h i s response i s a d i s t i n c t advantage to the species. Their response i s manifested by i n c r e a s i n g p r o d u c t i v i t y when n u t r i t i o u s and abundant forage becomes a v a i l a b l e (Geist 1974, Markgren 1969) . Moose are an important n a t u r a l resource. H i s t o r i c a l l y , i n the e a r l y 1900's, meat and hides were used f o r food and c l o t h i n g by indigenous peoples and European 1 2 s e t t l e r s . Since then, the r e c r e a t i o n a l value of moose has increased d r a m a t i c a l l y , e s p e c i a l l y i n the n o r t h - c e n t r a l region. During the 1970-74 p e r i o d , the Pri n c e George area (old management Areas 20-22) provided approximately an average of 16,000 man-days of r e c r e a t i o n and a harvest of 5,400 annually ( B r i t i s h Columbia F i s h and W i l d l i f e Branch 1970-1974). Most r e c e n t l y , the non-consumptive use of moose has been recognized as a growing and important value. A d d i t i o n a l l y , moose may w e l l provide a p r o t e i n source. The t e c h n i c a l f e a s i b i l i t y of "game ranching" moose has been demonstrated by Knorre (197 4) and others. Hence, moose w i l l become an i n c r e a s i n g l y v a l u a b l e resource. To deal adequately w i t h t h i s important resource, moose management must i n t e n s i f y . The f o r e s t s that moose c o l o n i z e d so s u c c e s s f u l l y have a l t e r e d since moose f i r s t appeared. This change i s an i n e v i t a b l e , n a t u r a l consequence of f o r e s t succession. The s i g n i f i c a n t d i f f e r e n c e w i t h f u t u r e changes w i l l be the impact of man and h i s a c t i v i t i e s . Of these p u r s u i t s , h a r v e s t i n g timber and c o n t r o l l i n g f o r e s t f i r e s w i l l be the most important. Since these sub-boreal f o r e s t s are e a s i l y a c c e s s i b l e and h i g h l y productive, f o r e s t - r e l a t e d development w i l l i n e v i t a b l y be widely d i s t r i b u t e d . Since moose are a l s o widely spread i n these f o r e s t s , the question a r i s e s , "What w i l l be the e f f e c t s of human a c t i v i t i e s upon moose?" The purpose of t h i s t h e s i s i s to examine moose 3 h a b i t a t w i t h p a r t i c u l a r reference to the e f f e c t s of f o r e s t - r e l a t e d a c t i v i t i e s . Much i s known g e n e r a l l y about moose and t h e i r h a b i t a t s (Bedard et a l . 1974). However, l i t t l e i s known about 1) moose h a b i t a t i n the n o r t h - c e n t r a l region of B r i t i s h Columbia, or 2) the impact of f o r e s t p r a c t i c e s upon t h i s h a b i t a t . These gaps are c r i t i c a l l i n k s i n achieving i n t e g r a t e d management of f o r e s t s and moose. C l e a r l y , a h a b i t a t - o r i e n t e d study does not deal w i t h a l l those components that a f f e c t moose populations. I t i s therefore u s e f u l to place such a study i n context. V a r i a t i o n s i n moose populations are determined by two broad types of mechanisms (Houston 1968) - environmental and popul a t i o n . The i n t e r f a c e or li n k a g e between them i s the energy and n u t r i e n t supply ( i n c l u d i n g water) a v a i l a b l e to moose. The a v a i l a b l e supply i s determined l a r g e l y by environmental mechanisms, although d e n s i t i e s and behaviour of moose can obviously modify a v a i l a b i l i t y as w e l l . The focus of the present study i s on some of the major environmental mechanisms that determine the a v a i l a b l e supply of energy and n u t r i e n t s to moose. This i s shown sch e m a t i c a l l y i n Figure 1.1. The method by which I studied these mechanisms i s r e f l e c t e d i n the o r g a n i z a t i o n of t h i s t h e s i s . Four t o p i c s make up the f i r s t p a r t of the t h e s i s , v i z . , h a b i t a t use and s e l e c t i o n , d i e t , l e v e l of use and bedding behaviour. In the u l t i m a t e sense, the a v a i l a b i l i t y of energy and n u t r i e n t s to 3a Figure 1.1 The general r e l a t i o n s h i p of sub-models th a t comprise a moose-forest model. Derived from Haagenrud and H j e l j o r d (1976) and Houston (1968). 4 WILDLIFE MANAGEMENT DECISIONS POPULATION MODEL CHANGES IN NATALITY, MORTALITY AND BODY WEIGHT. YEARLY HARVEST HERD STRUCTURE, DENSITY AND NUMBER OF ANIMALS BEHAVIOUR MODEL hACTIVITY ENERGY MODEL 7 SATISFAC- TION COVER GRAZ- ING HABIT FOOD REQUIREMENT FOREST/PATTERN/ PLANT SUCCES- SION MODEL 5T • FOOD ["AVAILABLE .GRAZING QUANTITY GRAZING/ BROWSING MODEL FORES MANAGEMENT DECISIONS TIMBER HARVEST 5 moose i s set by the conversion of s o l a r energy i n t o the chemical energy stored i n p l a n t s . However, not a l l of the vege t a t i o n i s forage f o r moose, and not a l l the forage i s a v a i l a b l e to them. Moreover, vegetation a l s o provides escape cover and s h e l t e r from the elements. Thus the f i r s t task was an attempt to perceive the sub-boreal f o r e s t environment i n terms meaningful to moose. Therefore, h a b i t a t s were defined by examining the use and s e l e c t i o n of vegetation types. Moose forage was assessed by food h a b i t s t u d i e s . Browse surveys were used to estimate the pr o p o r t i o n of the forage t h a t was eaten. F i n a l l y , since the a v a i l a b i l i t y of s h e l t e r a f f e c t s the a v a i l a b i l i t y of food (Bunnell 1974), I a l s o examined the s e l e c t i o n of bed s i t e s . These four components make up the f i r s t p a r t of the t h e s i s . Once these b i o l o g i c a l features were de f i n e d , I examined three major f a c t o r s that modified them. These three t o p i c s comprise the second p a r t of the t h e s i s , v i z . , f o r e s t succession, forage n u t r i t i v e v a l u e s , and c l i m a t e . Succession was studied since i t determines long term trends i n forage production and the p r o v i s i o n of cover. N u t r i t i v e aspects of moose forages were q u a n t i f i e d to determine the r e l a t i o n s between species eaten and n u t r i e n t content, and to assess what f a c t o r s i n f l u e n c e d n u t r i e n t l e v e l s . Climate was examined, e s p e c i a l l y snow, as i t i n f l u e n c e s the occupancy of winter ranges, the a v a i l a b i l i t y of forage, and the s e l e c t i o n and use of h a b i t a t s . These three t o p i c s t h a t comprise the 6 second p a r t of the t h e s i s are a l s o modified by f o r e s t p r a c t i c e s . The t h i r d p a r t of the t h e s i s , the D i s c u s s i o n , attempts to combine the f i r s t two p a r t s , to assess the e f f e c t s of f o r e s t r y a c t i v i t i e s on moose, and to provide an overview and recommendations f o r w i l d l i f e managers. 1.2 The Approach to the Study The present management of harvestable w i l d l i f e i n B r i t i s h Columbia has emphasized population dynamics r a t h e r than h a b i t a t r e l a t i o n s h i p s . U n t i l q u i t e r e c e n t l y , w i l d l i f e managers regulated harvest through manipulation of hunting seasons and bag l i m i t s . Data were c o l l e c t e d from game and hunter checks, harvest and hunter que s t i o n n a i r e surveys, and carry-over and post-season counts. L i t t l e attempt was made to modify the production of w i l d l i f e through h a b i t a t manipulation, although the importance of h a b i t a t was w e l l recognized, e.g., Smith (1955). In the 1960's, the t a c i t r e c o g n i t i o n of h a b i t a t ' s r o l e evolved to an a c t i v e p u r s u i t of h a b i t a t management. E f f i c i e n t f o r e s t f i r e c o n t r o l and expanded logging were the two prime f a c t o r s inducing the development of moose h a b i t a t management programs i n n o r t h - c e n t r a l B r i t i s h Columbia. The i n c r e a s i n g e f f i c i e n c y of p r o t e c t i n g sub-boreal f o r e s t s from w i l d f i r e v i t i a t e d the main agent f o r c r e a t i n g e a r l y s u c c e s s i o n a l ranges. R e a l i z i n g the r e l a t i o n s h i p between 7 high moose d e n s i t i e s and e a r l y s u c c e s s i o n a l stages, the adverse impact of f i r e c o n t r o l on moose h a b i t a t s became r e a d i l y apparent. The r a p i d l y i n c r e a s i n g acreage of f o r e s t s t h a t were logged was a l s o apparent. Thus, the p r i s t i n e , u n c o n t r o l l e d agent of range c r e a t i o n was p r o g r e s s i v e l y being suppressed and replaced by an agent whose impact was c o n t r o l l e d by human design. Although c r e a t i o n or m o d i f i c a t i o n of moose h a b i t a t was a n c i l l a r y to f o r e s t r y , b i o l o g i s t s r e a l i z e d that h a b i t a t management f o r moose was r e a d i l y f e a s i b l e . A passive regard f o r v e g e t a t i o n a l changes was transformed to an a c t i v e i n t e r e s t i n modifying f o r e s t s by logging f o r moose production. The broadened pe r s p e c t i v e of w i l d l i f e management from p r i m a r i l y animals-only to animal s - a n d - t h e i r - h a b i t a t was a major development. The growing awareness of o p p o r t u n i t i e s f o r h a b i t a t management were accompanied by the r e a l i z a t i o n that moose h a b i t a t r e l a t i o n s h i p s i n sub-boreal f o r e s t s were l a r g e l y unknown, except i n the most general f a s h i o n . E x i s t i n g i n f o r m a t i o n w i t h respect to logged h a b i t a t s was inadequate or poorly understood. Previous moose studie s i n B r i t i s h Columbia were n e i t h e r w i t h i n the sub-boreal f o r e s t s nor d e a l t w i t h other aspects of moose ecology (Hatter 1950, Cowan et a l . 1950, Baynes 1956, R i t c e y and Verbeek 1969, Finnegan 1973). Although they provided u s e f u l background i n f o r m a t i o n , the major problem of h a b i t a t r e l a t i o n s h i p s i n n o r t h - c e n t r a l 8 f o r e s t s was not addressed. Previous s t u d i e s of moose h a b i t a t s elsewhere i n circumboreal ranges were a l s o u s e f u l as background informat i o n (e.g., Bergerud and Manuel 1968, Houston 1968, Lykke 1964, Peek 1971, Peterson 1955,,Pimlott 1961, Stevens 1970, T e l f e r 1967). However, despite r a t h e r extensive s t u d i e s of fo r e s t e d h a b i t a t s , logged areas have received s u r p r i s i n g l y l i t t l e a t t e n t i o n . Most st u d i e s have d e a l t w i t h n a t u r a l areas, or, where logging had occurred, i t received almost i n c i d e n t a l a t t e n t i o n (e.g., Stevens 1970). Even i n studie s c a r r i e d out p r i m a r i l y on cutovers, e.g., Bergerud and Manuel (1968), l i t t l e i n formation on r e l a t i v e use of d i f f e r e n t types and ages of cutovers has been provided. Studies such as those r e c e n t l y published by Peek et a l . (1976)are exceptions to the r u l e . 1.3 The Need f o r Integrated Management U n t i l q u i t e r e c e n t l y , s u p p l i e s of the common property resources exceeded demands. Low p r i c e s f o r land, g r a z i n g a l l o t m e n t s , c u t t i n g r i g h t s , and hunting l i c e n c e s , a l l r e f l e c t e d the apparent superabundance of resources. In f a c t , considerable e f f o r t was expended by government to encourage settlement and development. Two concomitant pressures r a d i c a l l y a l t e r e d t h i s s i t u a t i o n . Human population increased at an exponential r a t e , i n c r e a s i n g demands f o r goods and s e r v i c e s which i n turn 9 a c c e l e r a t e d demands f o r n a t u r a l resources. Many c i t i z e n s experienced an i n c r e a s i n g amount of l e i s u r e time r e s u l t i n g from wealth accrued through n a t u r a l resource developments i n the r e g i o n . These growing demands f o r a v a r i e t y of goods and s e r v i c e s , confronted one b a s i c l i m i t : a f i n i t e land base. As demands approached the l i m i t s imposed by b i o p h y s i c a l parameters, they i n e v i t a b l y l e d t o c o n f l i c t and competition. The i n t e r e s t i n m u l t i p l e and i n t e g r a t e d use was an attempt to r e s o l v e these c o n f l i c t s i n a r a t i o n a l manner. Whether t h i s attempt w i l l be s u c c e s s f u l or not i s s t i l l unknown. I t can be assumed th a t an i n t e g r a t e d system of n a t u r a l resource a l l o c a t i o n and management i s a prime option a v a i l a b l e to the p u b l i c . However, i n the present system, important demands are u s u a l l y made by the producers and users of these resources. In s i m p l i s t i c terms, these can be stated as: d e f i n i n g o b j e c t i v e s , d e v i s i n g a planning process, i n v e n t o r y i n g of resources, implementation of plans followed by e v a l u a t i o n and re-adjustment. The o b j e c t i v e s of i d e n t i f i e d user groups must a l s o be defined i n o p e r a t i o n a l terms so t h a t they can be evaluated, i n t e g r a t e d and i n i t i a t e d w i t h i n the l i m i t s of a v a i l a b l e resources, time and technology. E f f e c t i v e i n t e g r a t e d f o r e s t and w i l d l i f e management must be dynamic r a t h e r than s t a t i c . The need f o r c o n t i n u a l re-assessment and e v a l u a t i o n of programs i s obvious. 10 Competing demands f o r l i m i t e d resources and t h e i r i n t e g r a t i o n has important e c o l o g i c a l i m p l i c a t i o n s . F i r s t , the o p t i o n of " l e t t i n g nature take i t s course" may appear l e s s tenable because n a t u r a l catastrophes such as w i l d f i r e and i n s e c t outbreaks may s e r i o u s l y d i s r u p t the production of de s i r e d and needed f o r e s t products. The apparent impact of n a t u r a l catastrophes on the economy may be d i r e c t l y r e l a t e d to the degree of resource commitment. That s o c i e t y has accepted t h i s r e l a t i o n s h i p i s c l e a r l y demonstrated by commitments to f o r e s t f i r e suppression and i n s e c t c o n t r o l programs. However, i t i s worth n o t i n g that economic f a c t o r s may be as c a t a s t r o p h i c as n a t u r a l ones (K. Sumanik, pers. comm.). A second e c o l o g i c a l i m p l i c a t i o n i s that human beings impose novel means of r e - d i s t r i b u t i n g and using resources by a l t e r i n g n a t u r a l patterns and processes. In f o r e s t r y , wood products w i t h t h e i r n u t r i e n t and energy content are mechanically removed and i n j e c t e d i n t o eco- systems of t e n f a r removed from t h e i r source. In w i l d l i f e and f i s h e r i e s management, v e r t e b r a t e species are ex t r a c t e d by unnatural methods and transported to other systems. None of these i m p l i c a t i o n s i s n e c e s s a r i l y d e l e t e r i o u s to the producing systems, although some logging methods may cause n u t r i e n t d e p l e t i o n on poor s i t e s . Since we have great expectations from n a t u r a l systems, and since we l i m i t or at l e a s t modify t h e i r f u n c t i o n i n g , i t i s 11 obvious that we should understand how they work i n order to manage them b e t t e r . We should r e a l i z e what e f f e c t s our a c t i v i t i e s have upon t h e i r continued a b i l i t y to produce what we r e q u i r e . 1.4 A Land Use P e r s p e c t i v e 1.4.1 General i n t r o d u c t i o n The purpose of t h i s s e c t i o n i s to r e l a t e human settlement and i n d u s t r i a l development i n the general study area, to moose and t h e i r h a b i t a t . Although the main subject of t h i s t h e s i s i s w i t h f o r e s t r y - r e l a t e d impacts, t h i s land use cannot be considered i n i s o l a t i o n from others. A b r i e f h i s t o r y of land use a l s o c l e a r l y i d e n t i f i e s changes i n r a t e s of land use a c t i v i t i e s : that the l a r g e s c a l e of modifying n a t u r a l systems i s c l e a r l y a recent phenomenon. Morice (1905) r e l a t e d that from time immemorial, w i l d l i f e of n o r t h - c e n t r a l B r i t i s h Columbia "have been trapped or chased by the American r e p r e s e n t a t i v e s of the human species who c a l l themselves Bene (men). . . ." (Morice 1905:4). From Morice's d e s c r i p t i o n , C a r r i e r s u b s i s t e d by hunting and f i s h i n g and depended e s p e c i a l l y upon salmon (Oneorkynehus spp. ) . They were semi-nomadic, s h i f t i n g winter quarters to meet f u e l needs, moving to lakes i n s p r i n g f o r f i s h i n g , and camping at s u i t a b l e salmon f i s h i n g s i t e s i n l a t e summer and e a r l y f a l l (Morice 1905). Morice made no mention of the C a r r i e r s s t a r t i n g f o r e s t f i r e s although he 12 s t a t e d t h a t "black pine i s f a i r l y common a l l over the country. . ." (Morice 1905:2). Based on these accounts, which date back to approximately 1600, the indigenous people probably had only a minor i n f l u e n c e upon the land and r e s i d e n t w i l d l i f e , . a t l e a s t u n t i l the beginnings of the f u r trade i n the e a r l y 1800's. Furbearers were the i n i t i a l reason f o r e x p l o r a t i o n and settlement i n the study area by Europeans. The f i r s t European e x p l o r e r was Alexander MacKenzie i n 1793. Simon Fraser followed i n 18 0 6 when he and h i s companions e s t a b l i s h e d F o r t St. James, and i n 18 07 when he founded F o r t George (now Pr i n c e George). Fur t r a d i n g continued to be the economic mainstay of the area u n t i l development of the Grand Trunk P a c i f i c Railway i n 1915. P l a c e r mining was an e a r l y i n d u s t r y , but i t i n v o l v e d p r i m a r i l y i n d i v i d u a l prospectors. The impact of the small number of f u r - t r a d e r s and pl a c e r miners on moose and moose h a b i t a t was probably in c o n s e q u e n t i a l when compared wi t h l a t e r a c t i v i t i e s . Moose were uncommon, and the need f o r l a n d - c l e a r i n g was minimal. The r a t e of tre e c u t t i n g must have been slow s i n c e , f o r example, lumber f o r the Hudson Bay f o r t s was a l l hand sawn. The most l i k e l y impact was an increased incidence of f o r e s t f i r e s , although evidence on t h i s p o i n t i s scanty. The f i r s t major burst of a c t i v i t y r e s u l t e d from the survey and c o n s t r u c t i o n of the Grand Trunk P a c i f i c Railway. The era surrounding t h i s development (1910-1916) witnessed a land boom spurred by a n t i c i p a t i o n of great expansion once the r a i l w a y was completed ( K e l l y and Farstad 1946). By about 1915, approximately 84,000 ha of land were a l i e n a t e d i n the general area, although more than 50 percent subsequently r e v e r t e d to the Crown. This high r e v e r s i o n r a t e was a t t r i b u t e d to economic c o n d i t i o n s a f t e r the completion of the Grand Trunk P a c i f i c r a i l l i n e , f a i l u r e of the P a c i f i c Great Eastern to reach Pr i n c e George, and the departure of many s e t t l e r s to World War I ( K e l l y and Farstad 1946). An i n t e r e s t i n g personal r e c o l l e c t i o n of t h i s c o l o r f u l p e r i o d can be found i n Walker (1972). A l s o , i t was during t h i s time t h a t a g r i c u l t u r e and f o r e s t r y began, p r i m a r i l y i n response to demands associated w i t h r a i l w a y c o n s t r u c t i o n and by the l o c a l populace. Towns and v i l l a g e s were slow to develop. Most of them were s i t e d at h i s t o r i c a l f o r t s , or c l o s e to f o r e s t r y and a g r i c u l t u r a l a c t i v i t i e s . C o n f l i c t s between settlement and moose were probably minimal at that time since both s e t t l e r s and moose were uncommon and sp a r s e l y d i s t r i b u t e d . The post-World War I I era saw tremendous expansion i n population and i n d u s t r y . A u s e f u l index of t h i s development i s the growth of Prin c e George (Table 1.1). The expansion was accompanied by r a p i d development of t r a n s p o r t a t i o n and u t i l i t y c o r r i d o r s . These changes have had a profound impact on moose not only through l o s s and disturbance of h a b i t a t , but a l s o by i n c r e a s i n g a c c e s s i b i l i t y of moose herds to 14 Table .1.1 Population Growth and Future P r o j e c t i o n s f o r Pr i n c e George and the Surrounding D i s t r i c t Recorded Population Census Pr i n c e George C i t y * * Population Year D i s t r i c t 8* d i d bdy new bdy P r o j e c t i o n * * * pre-1793 >12,000+ >350+ 1807 >12,000+ >350+ 1911 ca. 2,000++ c700 1915 >5,500 c3,500 1921 17,631 2,053 1931 21,534 2,479 1941 25,276 2,027 1951 40,276 4,703 1956 60,067 10,563 1961 74,240 13,877 32,268 1966 103,767 24,471 51,671 1971 128,205 33,101 49,365 64,365 1976 58,292 86,677 1981 111,279 1986 135,874 1991 160,030 1996 184,854 *Dominion Bureau of 186,440 km.2 S t a t i s t i c s 1 Census D i s t r i c t 8 covers * * C i t y 1968, boundaries were extended i n 1961, 1964, 1965, 1967, 1970 and most r e c e n t l y i n 1975. ***From B.C. Research (1974). Study area boundaries included P r i n c e George and the immediate l o c a l i t y . +Hudson Bay Company made these estimates i n 1856. ++Area population based on Land Recording D i s t r i c t . 15 hunters from the l a r g e population centres i n southern and c e n t r a l B r i t i s h Columbia. C u r r e n t l y , the study area i s tra v e r s e d by two major p r o v i n c i a l highways, a myriad of secondary and logging roads, two major r a i l w a y l i n e s , one e l e c t r i c a l t r a n s m i s s i o n l i n e , and one n a t u r a l gas l i n e . Prime and c r i t i c a l moose h a b i t a t i n the F i n l a y and Parsnip Rivers has been flooded and destroyed by the W. A. C. Bennett Dam and W i l l i s t o n Reservoir. Flooding of v i r t u a l l y the e n t i r e McGregor River and many of i t s t r i b u t a r i e s i s under a c t i v e c o n s i d e r a t i o n by government. Dams proposed f o r the upper Fraser River would be d e v a s t a t i n g f o r moose. The Kenney Dam at the headwaters of the Nechako River has so c o n t r o l l e d and modified the n a t u r a l h y d r o l o g i c a l regime of the r i v e r t h a t i t s c a p a b i l i t y as a "moose r i v e r " i s l i k e l y reduced. I t has al s o pre-empted h i g h l y productive moose h a b i t a t . Major development events are l i s t e d b r i e f l y i n Table 1.2. 1.4.2 Mining As i n most other regions of B r i t i s h Columbia, mining i n the Pr i n c e George d i s t r i c t has had an e r r a t i c h i s t o r y . Major developments began i n 1861, when p l a c e r miners came northwards from the Cariboo p l a c e r gold f i e l d s to explore the F i n l a y and Parsnip R i v e r s . Eight years l a t e r , gold was discovered i n the Manson, Germansen and Omineca Rivers and t h e i r t r i b u t a r i e s . The f o l l o w i n g gold rush was h e c t i c , and 16 Table 1.2 Major Events i n the Settlement and Growth of Pri n c e George and the Surrounding Region Year Event pre'^17 93 Indian v i l l a g e ( " L h e i t l i " ) at confluence of Nechako and Fraser Rivers 1793 F i r s t recorded e x p l o r a t i o n by Europeans (A. MacKenzie) 1807 F o r t George e s t a b l i s h e d 18 21 Merger of HBC and North West Fur companies 1861 P l a c e r gold mining on F i n l a y and Parsnip Rivers 18 69 P l a c e r gold mining on Manson, Germansen and Omineca Rivers and t h e i r t r i b u t a r i e s 19 09 F i r s t sawmill opened i n Pr i n c e George 1914 Completion of Grand Trunk P a c i f i c Railway, W.W.I began. 1915 P r i n c e George incorporated 1918 W.W.I, ended 1952 Kenney Dam completed. John Hart highway completed between P r i n c e George and Dawson Creek. B.C.R. l i n e completed from Quesnel to Pri n c e George. 1956 Vancouver to Squamish l i n k of B.C.R. completed 1958 Pri n c e George to Dawson Creek, and to F t . St. John se c t i o n s of B.C.R. completed 1965 Yellowhead highway between P r i n c e George and McBride completed 1967 B.C.R. extension to F t . St. James 1968 Three pulp m i l l s opened i n Pr i n c e George. W. A. C. Bennett power p r o j e c t on Peace River completed. 1975 A g r i c u l t u r a l land reserve e s t a b l i s h e d around Pr i n c e George o 1.7 i t d e c l i n e d q u i c k l y a f t e r 1875. Mining a c t i v i t y was q u i e t during the next 50 years u n t i l i n the 1930's, when the pl a c e r deposits northwest of P r i n c e George were developed. Again, the f l u r r y of a c t i v i t y was b r i e f . Today, an o l d mining road can s t i l l be seen on the Salmon River study area, and u n t i l 1974, a l a r g e s l u i c i n g device was at the o l d E.M.K. m i l l s i t e , approximately 60 km NE of P r i n c e George. The outbreak of World War I I strengthened mineral p r i c e s and sti m u l a t e d e x p l o r a t i o n and development of many lode metals. A f t e r the war, a c t i v i t y subsided once more, but the recent, dramatic increases i n the i n t e r n a t i o n a l gold p r i c e s again spurred renewed i n t e r e s t i n p l a c e r mining on many streams and r i v e r s i n the P r i n c e George area, e s p e c i a l l y to the southeast. In general, these a c t i v i t i e s are undertaken by small operators. These operations can adversely a f f e c t f i s h and w i l d l i f e through s i l t a t i o n and h a b i t a t d e s t r u c t i o n . C u r r e n t l y , the only mine operating w i t h i n the P r i n c e George area (Fraser-Ft. George Regional D i s t r i c t ) i s a small limestone quarry west of P r i n c e George. The c i t y , however, plays an important r o l e as a supply and s e r v i c e center f o r much of the a c t i v e e x p l o r a t i o n to the north and northwest. The economic value of mines i n the Omineca Mining D i s t r i c t from 1926 to 1974 i s tabulated i n Appendix Table C-2. Future development p o s s i b i l i t i e s appear l i m i t e d , except p o s s i b l y f o r c o a l (I.P.A. 1976). A non-coking c o a l 18 d e p o s i t , estimated at 73 x 10 6 t , l i e s on the Bowron R i v e r , approximately 60 km southeast of P r i n c e George. Development of t h i s deposit would see increased access, p o s s i b l y a r a i l spur connecting to the BCR or the CNR, and d i r e c t employment of 1,000 people (I.P.A. 1976). Con s t r u c t i o n of a proposed s t e e l m i l l at P r i n c e George would a l s o increase P r i n c e George's population as w e l l as increase l i k e l i h o o d of c o a l e x t r a c t i o n from the Bowron V a l l e y . The most important impact of mining and associated e n t e r p r i s e s on moose r e l a t e s to increased access and increased hunting. Past and p o t e n t i a l mining a c t i v i t i e s have d i s t u r b e d r e l a t i v e l y l i t t l e h a b i t a t , except some r i p a r i a n h a b i t a t s along p l a c e r streams. The roads l e a d i n g to mines, and e s p e c i a l l y the i n c r e a s i n g number of people at P r i n c e George associated w i t h supplying and s e r v i c i n g , have a much greater impact on moose. 1.4.3 A g r i c u l t u r e A g r i c u l t u r a l development has a f f e c t e d moose p r i m a r i l y through conversion of v a l u a b l e h a b i t a t s i n lowland areas i n t o farmland. The f i r s t record of farming was a garden planted by D. W. Harmon i n 1811 at F o r t St. James (Runnalls 1946). Most i n i t i a l attempts at land c l e a r i n g were unsuccessful and t h e i r r e v e r s i o n was b e n e f i c i a l to moose. K e l l e y and Farstad (1946) noted that most e a r l y attempts to farm were small (2 to 8 ha) and s c a t t e r e d , w i t h a high r a t e of abandonment. This p a t t e r n l i k e l y b e n e f i t e d moose by c r e a t i n g small patches of e a r l y s u c c e s s i o n a l vegetation i n a p r i m a r i l y f o r e s t e d area. From a f t e r the r a i l w a y boom u n t i l the depression, farming was i n e c l i p s e , being unable to compete w i t h the more southern producers who had b e t t e r s o i l s and c l i m a t e . At t h i s time, many farmers turned to logging and road b u i l d i n g ( K e l l e y and Farstad 1946). With the r e d u c t i o n of these l a t t e r occupations during the depression, i n t e r e s t i n farming and land c l e a r i n g returned, and K e l l e y and Farstad (1946) b e l i e v e d that depression farming played an important r o l e i n developing a g r i c u l t u r e . S i m i l a r to growth i n settlement, a g r i c u l t u r e expanded i n the post-war p e r i o d . I n i t i a t i o n of t h i s phase of a g r i c u l t u r e a l s o represented the beginnings of i n c o m p a t a b i l i t y between moose and farming and ranching. Large land areas were logged or burned or both, and put to the plow. Most of t h i s land was at low e l e v a t i o n , e.g., along r i v e r v a l l e y s and i n the g l a c i o - l a c u s t r i n e sediments below 760 m i n e l e v a t i o n . While these areas represented the highest c a p a b i l i t y a g r i c u l t u r e lands i n the r e g i o n , they a l s o represented high c a p a b i l i t y winter moose h a b i t a t s . The r a t e of expansion i s presented i n Table 1.3. By the l a t e 1960's, a g r i c u l t u r e was s t i l l l a r g e l y i n the development stage and was c o n t i n u i n g to increase i t s p r o d u c t i v i t y annually (Oswell 1969). 20 Table 1.3 Number and Area of Farms, C a t t l e f o r the Province P r i n c e George Region, and Numbers of and f o r the 1881-1971 No. of farms/km 2* Area of farms/km 2* No. of ca t t i e / k m 2 * Year p r o v i n c i a l r e g i o n a l p r o v i n c i a l r e g i o n a l p r o v i n c i a l r e g i o n a l 1881 .003 .002 .086 1891 .007 .136 1901 .007 .007 .134 1911 .018 .011 .001( 1%) ' .150 1921 .024 .012 .031( 1%) .234 (46%) 1931 .028 .011(8%) .015 .012(16%) .251 (12%) 1941 .028 .012(9%) .018 .013 .359 .228(13%) 1951 .028 .020 1956 .027 .020 1961 .021 .007(7%) .020 .014(15%) 1966 .021 .008(7%) .020 1971 .020 .012 .616 .223( 2%) *Data expressed on a per km 2 b a s i s s i n c e the sample area changed during sample time. The r e g i o n a l census from 1881-1931 was the Cariboo e l e c t o r a l d i s t r i c t (81,417 km 2); from 1941-1966, Census D i s t r i c t 8 (186,440 km 2); and i n 1971, the F r a s e r - F o r t George Regional D i s t r i c t (51,196 km 2). Area of the province i s 930,528 km2. O r i g i n a l data i n Appendix Table C-2. **Regional t o t a l s as p r o p o r t i o n ( i n percent) of p r o v i n c i a l t o t a l s i n paren t h e s i s . 21 Now, as i n the past, the mainstay of r e g i o n a l a g r i c u l t u r e i s l i v e s t o c k and a s s o c i a t e d forage production. The major areas around P r i n c e George are: a) Reid, Chief and Nukko Lakes b) the lower Salmon River V a l l e y , east of Highway 97 c) south of P r i n c e George to Woodpecker, p r i m a r i l y on the east side of the Fraser R i v e r d) s c a t t e r e d areas i n the Willow River - Aleza Lake v i c i n i t y e) west of P r i n c e George along Highway 97. The Pineview c l a y s o i l s and a l l u v i a l m a t e r i a l s are used most commonly f o r a g r i c u l t u r a l purposes. Despite o p t i m i s t i c expectations f o r a g r i c u l t u r e , economic and b i o p h y s i c a l c h a r a c t e r i s t i c s f o r e t e l l an u n c e r t a i n f u t u r e . The Pineview s o i l s r e q u i r e expensive c l e a r i n g and are d i f f i c u l t to manage: they are slow to heat, puddle when wet and pack when dry. The a l l u v i a l t e r r a c e s i n bottomlands are fragmented, l e a d i n g to a "pocket" a g r i c u l t u r e where access and c l e a r i n g costs are high. Except f o r m i c r o c l i m a t i c v a r i a t i o n s along the major drainages, the growing season i s c o o l , w i t h frequent r a i n f a l l ; crop maturity i s o f t e n delayed by these f a c t o r s . Crop a l t e r n a t i v e s are l i m i t e d . The long and c o l d winters n e c e s s i t a t e extended periods of winter feeding. Distance to markets, unstable p r i c e s f o r products and more productive lands elsewhere a l s o add to the d i f f i c u l t i e s f a c i n g 22 a g r i c u l t u r a l development i n the re g i o n . Other a g r i c u l t u r a l e n t e r p r i s e s i n c l u d e sheep ranching, and vegetable, egg, and l i m i t e d h o r t i c u l t u r a l production. However, these make a minor c o n t r i b u t i o n to the o v e r a l l r e g i o n a l a g r i c u l t u r e compared with' the beef i n d u s t r y . Steps by government to increase and promote a g r i c u l t u r e i n the region are many and v a r i e d . Community pastures are sponsored and p a r t l y s u b s i d i z e d by both p r o v i n c i a l and f e d e r a l governments. In one case at Giscome, a community pasture i s developing on an important moose winter range, where land c a p a b i l i t i e s c l e a r l y favor timber and w i l d l i f e . A g r i c u l t u r a l land surrounding P r i n c e George was protected by an A g r i c u l t u r a l Land Reserve (ALR) through l e g i s l a t i o n approved i n 1974. These i n c e n t i v e s are obv i o u s l y encouraged by the a g r i c u l t u r a l s e c t o r and others. The need to preserve arable land i n t h i s , and elsewhere i n the province, i s an obvious one that meets w i t h general p u b l i c approval. A growing urban center l i k e P r i n c e George undoubtedly can support a l o c a l a g r i c u l t u r e , given s o l u t i o n to problems such as i n c r e a s i n g egg production quotas f o r l o c a l producers. A l s o , there i s a d e s i r e to preserve ah a g r i c u l t u r a l l i f e s t y l e although the costs of t h i s needs to be c a r e f u l l y evaluated. This d e s i r e assures the continued presence and development of a g r i c u l t u r e . I t i s a l s o obvious t h a t present-day a g r i c u l t u r e and moose and other w i l d l i f e are l e s s than compatible. Land 23 a l i e n a t i o n , c l e a r i n g and maintenance of forage-producing areas d i r e c t l y remove p o t e n t i a l moose h a b i t a t . Other items such as predator c o n t r o l i n o u t l y i n g , marginal farms pose other d i f f i c u l t problems. These lands o f t e n have high c a p a b i l i t i e s f o r moose. The d i s t u r b i n g aspect of these c o n f l i c t i n g forms of land use i s not only the l o s s of h a b i t a t , but a l s o the apparent lack of r e g i o n a l o b j e c t i v e s to give purpose and d i r e c t i o n to proper land use, and to provide acceptable land use zoning. 1.4.4 F o r e s t r y Although f u r s and, to a l e s s e r extent, minerals provided the i n i t i a l impetus to development and settlement of the Prince George area, timber h a r v e s t i n g has been the main- stay of the economy since the turn of the century. I t s impact on the n a t u r a l systems and human settlements of the region have been complex and extensive. Perhaps the most remarkable feature has been the r a p i d transformation from very simple operations i n the e a r l y 1900"s to a s o p h i s t i c a t e d , i n t e g r a t e d i n d u s t r i a l complex i n the 1970's. P r i o r to 1909, a l l wooden b u i l d i n g s were e i t h e r of log or hand-sawn lumber. The f i r s t sawmill opened i n P r i n c e George i n November, 1909 (Walker 1972). I t s appearance r e f l e c t e d the demand f o r t i e s used i n the c o n s t r u c t i o n of the Grand Trunk Railway, and the concomitant demand f o r b u i l d i n g m a t e r i a l s by s e t t l e r s d u r i n g the 1910-1916 land boom. By 1910 there were three sawmills. 24 In 1912 the Forest Branch was set up and two years l a t e r the f i r s t Crown timber auction was held i n the d i s t r i c t (Glew 1963). Talk of b u i l d i n g pulp m i l l s was a l s o current during t h i s e a r l y period (Runnalls 1946), but these proposals were not r e a l i z e d u n t i l more than 50 years l a t e r . Secondary wood processing began i n 1912 when the f i r s t sash and door m i l l opened i n Prince George. Whitford and C r a i g (1918) summarized the r e s t r i c t e d extent of lumber operations at that time. One l a r g e m i l l was s i t e d at the mouth of the Willow R i v e r , and another small one was located at Giscome. Logging was not e x t e n s i v e , being confined to f o r e s t s adjacent to the m i l l s i t e s . V i r t u a l l y no logging occurred i n the drainages of the S t u a r t , Salmon, Nation, P a r s n i p , Nechako and Blackwater. Lumbering a c t i v i t i e s i n the Upper Fraser area were "not extensive" (Whitford and C r a i g 1918). By 1920, 20 sawmills were operating i n the d i s t r i c t w i t h an annual cut of 97,000 c u n i t s (Figure 1.2, see a l s o Appendix Table C-3). This e a r l y phase of h a r v e s t i n g was t y p i c a l l y done by horse-logging. I t had only a l i m i t e d impact on the f o r e s t ecosystem. Impact on moose h a b i t a t was l i k e l y b e n e f i c i a l . Forest management was minimal although i n t e r e s t i n f i r e c o n t r o l was growing. Harvesting was b a s i c a l l y an e x t r a c t i o n a c t i v i t y with l i t t l e regard to the f u t u r e . Growth of the indu s t r y was slow and s l i g h t l y e r r a t i c u n t i l the end of World War I I . 24a Figure 1.2 Development of f o r e s t r y i n the P r i n c e George Forest D i s t r i c t as i n d i c a t e d by the annual cut and the number of operating sawmills, 1914-1974. Records f o r the sawmills were discontinued a f t e r 1968. Source of data: annual r e p o r t s of the B r i t i s h Columbia Forest S e r v i c e . ANNUAL CUT (CUNITS*104) o o O O o o o 0 0 o o N U M B E R O F S A W M I L L S O P F R A T i M n 26 The post-war era witnessed tremendous expansion (Figure 1.2). This was made p o s s i b l e by a r a p i d t r a n s i t i o n from horse to mechanized logging. Market demands increased as w e l l . Evidence of the r a p i d growth i s given by the t o t a l annual cut which almost doubled from 1945 to 1972 when i t was 252,000 c u n i t s and 407,500 c u n i t s , r e s p e c t i v e l y . S i l v i c u l t u r a l systems evolved r a p i d l y i n response to increased concern f o r ensuring sustained y i e l d s . E a r l i e r logging was p r i m a r i l y diameter l i m i t or commercial c l e a r - c u t t i n g , w i t h large amounts of the o r i g i n a l stand being l e f t . In 1951, s i n g l e tree s e l e c t i o n became o p e r a t i o n a l (Glew 1963). A l t e r n a t e s t r i p c u t t i n g was i n s t i t u t e d i n 1954 and s c a r i f i c a t i o n , as a stand treatment technique, was introduced i n 1956 (Glew 1963). The dominance of the p a r t i a l c u t t i n g d e c l i n e d by the e a r l y 1960's and was replaced by c l e a r c u t t i n g (Table 1.4). Accompanying t h i s t r a n s i t i o n was the i n s t i t u t i o n of ha r v e s t i n g to a c l o s e u t i l i z a t i o n standard. Although the area logged and the annual cut increased, more e f f i c i e n t u t i l i z a t i o n of wood meant that these increases were not p a r a l l e l : the annual area cut grew more slowly than the amount of wood ex t r a c t e d (cf. Figure 1.1 and Table 1.4). The l a s t twenty years a l s o saw the end of the bush- m i l l where small sawmills were s i t u a t e d near the logging operation. Wood processing became h i g h l y i n t e g r a t e d and 27 Table 1.4 Trends i n Logging Methods and Area Cut i n the Pr i n c e George Forest D i s t r i c t , 1950 - 1973* Area logged (km 2) Prop. (%) of s e l e c t i v e methods T o t a l area Year c l e a r c u t p a r t i a l seed tre e d i a . l i m i t s i n g l e t r e e (km 2) 1950 29 122(81%)** 5 78 16 151 1951 25 146(85%) 12 69 18 171 1952 21 172(89%) 14 66 20 193 1953 m*** m 22 67 11 m 1954 m m 25 63 11 m 1955 m m 40 53 7 m 1956 m m m 1957 21 225(91%) 246 1958 34 209(86%) 243 1959 103 226(76%) 329 1960 - 131 194(62%) 315 1961 118 144(55%) 262 1962 182 145(44%) 327 1963 m m m 1964 m m m 1965 m m m 1966 368+ 368+ 1967 335+ 335+ 1968 413+ 413+ 1969 485+ 485+ 1970 442+ 442+ 1971 414 5(1%) 419 1972 368 4(1%) 373 1973 386 t ( l % ) + + 387 *Source of data: annual reports of the B.C. Forest S e r v i c e . - * * P r o p o r t i o n of t o t a l area cut by p a r t i a l c u t t i n g methods. ***Data missing. +Type of logging ( c l e a r c u t or p a r t i a l ) not a v a i l a b l e , but l i k e l y >99% c l e a r c u t . ++t i s l e s s t h a n 0.5%. 23 c e n t r a l i z e d , e s p e c i a l l y i n P r i n c e George. Three pulp m i l l s began operating between 1965 - 1970. The s o c i o l o g i c a l , economic and environmental impact of these developments were fa r - r e a c h i n g . They w i l l c l e a r l y shape the future character and nature of r e g i o n a l development. In f u t u r e , r e g i o n a l f o r e s t r y p r a c t i c e s w i l l demand a s t i l l more i n t e n s i v e use of timber since uncommitted wood supplies are very l i m i t e d (I.P.A. 1976). The C e n t r a l Report 76 (I.P.A. 1976) l i s t e d f i v e probable developments that are paraphrased as f o l l o w s : 1. A change i n the proportions of var i o u s uses of f o r e s t lands. 2. A refinement of logging p r a c t i c e s that w i l l reduce waste and breakage i n ha r v e s t i n g and t r a n s p o r t i n g . 3. An improvement i n wood processing technology. 4. An improvement i n the d i s t r i b u t i o n of raw m a t e r i a l s . 5. An upgrading of products and greater s p e c i a l i z a t i o n of wood products. ". . . the exact s c a l e , t i m i n g and f e a s i b i l i t y . . . w i l l remain to be determined by a c t u a l market c o n d i t i o n s and s i t e (project) s p e c i f i c f e a s i b i l i t y s t u d i e s i n the f u t u r e " (I.P.A. 1976:40-41). The foregoing changes have had major impacts on the h a b i t a t and production of moose. Both the r a t e and nature of logging created and impaired moose h a b i t a t . Logging a l s o produced novel types of h a b i t a t i n the sense that they had no 29 n a t u r a l counterparts. Murray (19 74) examined the impact of logging on h a r v e s t i n g moose through improved access. 1.4.5 W i l d f i r e W i l d f i r e i s the s i n g l e most s i g n i f i c a n t n a t u r a l f a c t o r t h a t modified moose h a b i t a t . Even p r i o r to the a r r i v a l of white man and moose i n n o r t h - c e n t r a l B r i t i s h Columbia, c i r c u m s t a n t i a l evidence i n d i c a t e s t h a t f i r e was a major e c o l o g i c a l f o r c e . Morice (1905) remarked that black (lodgepole) pine was f a i r l y common i n the area. His observations were made oivoa 1860, before the major i n f l u x of Europeans. The region ranks as high to very high on the Canadian Forest F i r e Weather Index maps (Simard 1973). Although f i r e h i s t o r y reports, are l a c k i n g f o r the general study area, p a l y n o l o g i c a l s t u d i e s by Rouse (1973, c i t e d by Smith 1974a) i n the so u t h - c e n t r a l I n t e r i o r revealed p o l l e n g r a i n s of lodgepole pine and charcoal l a y e r s that predate ash l a y e r s deposited a f t e r Oregon's Mt. Mazama erupted, approximately 7,000 years ago. D e t a i l e d f i r e research i n other b o r e a l f o r e s t ecosystems a t t e s t to the i n t e g r a l and long-standing r o l e of f i r e i n northern f o r e s t s (Heinselman 1973, and a review by K e l s a l l e t a l . 1977). As the number of Europeans increased, i t i s l i k e l y t h at the number of f i r e s and area burned increased. The magnitude of t h i s increase i s d i f f i c u l t to define without d e t a i l e d f i r e h i s t o r y research but again, cursory evidence 30 suggests i t was l a r g e . n The a r r i v a l of moose i n north- c e n t r a l B.C. and t h e i r remarkable southward d i s p e r s a l from 1900-1950 has been a t t r i b u t e d to increased f i r e a c t i v i t y a s s o c i a t e d w i t h white settlement (Hatter 1950). Dawson (1879) reported extensive burns between St u a r t and MacLeod Lakes. Whitford and C r a i g (1918) provided a d d i t i o n a l data t h a t a f f o r d an a p p r e c i a t i o n of w i l d f i r e impact on sub-boreal f o r e s t s (Table 1.5). In t h e i r c l a s s i f i c a t i o n of f o r e s t land v e g e t a t i o n , 53 percent of the major drainages (range of 14-70 percent) i n the n o r t h - c e n t r a l d i s t r i c t was "young growth!" I estimated these stands to be l e s s than 40 years o l d . Thus the f o r e s t s were burned, on an average, at 75-100 year i n t e r v a l s . E a r l y annual r e p o r t s of the Forest Branch r e f e r to the high incidence of f i r e s along the Grand Trunk Railway, and the need f o r f i r e p a t r o l s along the r i g h t of way. Campbell (1920) r e l a t e d t h a t at l e a s t two l a r g e f i r e s burned from the Nechako River north approximately 3 0 km almost to Summit Lake, and westward from the Fraser R i v e r . The f i r s t f i r e occurred 60-70 years p r i o r to 1919, and the second i n 1902 or 1903. The l a t t e r f i r e was l i k e l y very intense since Campbell remarked that s u b s o i l was exposed. There was have been many small f i r e s as w e l l as the very l a r g e and spectacular ones. These types of r e p o r t s suggest th a t the p e r i o d from approximately 1850-1920 was one of unusually high l e v e l s of w i l d f i r e . 31 Table 1.5 Estimated Areas of Broad Vegetation Classes i n F i v e Major Drainages i n North-Central B r i t i s h Columbia (from Whitford and Cr a i g 1918) Propor t i o n s of region i n va r i o u s v e g e t a t i o n c l a s s e s (%) Drainage region Estimated area (km 2) young growth merchantable timber incapable of timber growth above t i m b e r l i n e upper Fraser 19,661 14 27 2 57 Willow - Bowron 8,156 27 58 4 11 Parsnip 11,634 35 38 4 23 Stuart - Salmon - Nation 27,871 59 19 15 6 Nechako - Blackwater 58,350 70 14 8 8 A l l regions 125,672 53 22 8 17 S i n c e 1 1 9 2 0 r w i l d f i r e s h a v e b e e n more c a r e f u l l y a n d s y s t e m a t i c a l l y m o n i t o r e d b y t h e B.C. F o r e s t S e r v i c e ( F i g u r e 1 . 3 ) . The d a t a i n d i c a t e t h a t f o r m o s t y e a r s r e l a t i v e l y s m a l l a r e a s w e re b u r n e d . However, when i d e a l f i r e c o n d i t i o n s d e v e l o p , l a r g e t r a c t s b u r n d e s p i t e f i r e f i g h t i n g e f f o r t s . L a r g e a r e a s w e r e b u r n e d i n 1 9 2 2 , 1 9 4 2 , 1944, 1 9 4 8, 1956, 1961 and 1971 ( F i g u r e 1.3, T a b l e C - 4 ) . P e r h a p s t h e m o s t i m p o r t a n t p o i n t r e l e v a n t t o moose h a b i t a t i s t h e d i m i n i s h i n g n a t u r a l r o l e o f f i r e a s an a g e n t 31a Figure 1.3 Annual area (km2) burned by w i l d f i r e i n the Prin c e George Forest D i s t r i c t , 1910-1975. Source of data: records from the P r o t e c t i o n D i v i s i o n , B r i t i s h Columbia Forest S e r v i c e .  of h a b i t a t c r e a t i o n . This change has long-term consequences on a fi r e - a d a p t e d species such as moose (Geist 1971, 1974) since the t y p i c a l "boom-and^bust" c y c l e of p r i s t i n e environments w i l l become a p r o g r e s s i v e l y r a r e r event. Moose d e n s i t i e s w i l l continue to vary but probably w i t h much reduced amplitude. 2. THE STUDY AREAS 2.1 B i o p h y s i c a l S e t t i n g The general study area i s s i t u a t e d i n c e n t r a l B r i t i s h Columbia, about 8 00 km north of Vancouver (Figure 2.1). I t covers approximately 11,300 km2 roughly defined by a c i r c l e centered on Pr i n c e George, w i t h a rad i u s of 60 km. Most of t h i s area was not studied i n d e t a i l , but the s i t e s studied were s e l e c t e d to a l l o w g e n e r a l i z a t i o n to the l a r g e area. E l e v a t i o n a l range of the study area was 550 to 1220 m, w i t h most of i t 915 ± 150 m. The landscape of the study area i s highly modified by g l a c i a t i o n and associated events such as p r o g l a c i a l l a k e s . The f i n a l g l a c i a t i o n was so "intense" t h a t evidence of previous g l a c i a t i o n s i s scarce ( K e l l e y and Farstad 1946, Tipper 1971) P h y s i o g r a p h i c a l l y , the study area can be c l a s s i f i e d as part of the Fraser Basin of Holland (1964), or the Nechako Plateau of Tipper (1971). The l a t t e r author b e l i e v e d the Fraser Basin d i s t i n c t i o n i s an a r b i t r a r y one, and t h a t t h i s Basin merely represents the lowest p a r t of the er o s i o n surface of the Nechako Plateau (Tipper 1971:10). Regardless of t h i s divergence of o p i n i o n , the physiography can be described as: I t s f l a t or gen t l y r o l l i n g surface l i e s f o r the 34 34a Figure 2.1 Locations of the study areas, and of place names mentioned i n the t e x t . 10 0 I I—I t—I I STUDY AREAS lp 0̂ B PRIMARY 1 Eagle 2 Grove 3 Salmon 4 Bowron 5 Found 6 Pyfe 7 Limestone 8 McGregor Major highway KILOMETRES 5» SECONDARY 9 McKenzie 10 Pineview n Shell 12 Swamp 13 Teardrop 14 Telachick 15 Torpy 16 Whites Secondary roads 36 most part below 3,000 f t . and i s covered w i t h d r i f t and has few exposures of bedrock. On much of the surface the drainage i s poorly organized, and numerous lakes and poorly drained depressions are present. The area was occupied by i c e whose movement created drumlins and d r u m l i n - l i k e forms i n the g l a c i a l d r i f t . . . an eastward and northeastward movement of i c e . (Holland 1964:67). As the C o r d i l l e r a n i c e sheet melted, n a t u r a l drainage channels i n the Nechako Plateau were blocked by i c e or t i l l . Three la r g e lakes subsequently formed around P r i n c e George, Vanderhoof and F o r t St. James (Tipper 1971). Approximately 3035 km2 of the lower l y i n g d r u m l i n i z e d t i l l was covered by l a c u s t r i n e sediments - composed t y p i c a l l y of varved s i l t s , c l a y s and sands. The P r i n c e George pro- g l a c i a l lake was caused by i c e blockage of the Fraser River channel south of P r i n c e George. I t s l e v e l , and t h e r e f o r e the l e v e l of l a c u s t r i n e deposits was approximately 790 m. This l e v e l was determined by a bedrock l i p at Summit Lake, where the p r o g l a c i a l Fraser River flowed northward i n t o the Peace River system v i a the Crooked and Parsnip R i v e r s . (Today i t flows southward.) These l a c u s t r i n e deposits have been c o l l e c t i v e l y termed the Nechako P l a i n (Armstrong and Tipper 1948:285, quoted i n Tipper 1971). Their depth v a r i e s from over 120 m to t h i n overlays (0.3 m) t h a t cap s t i l l obvious drumlins: o u t l i e r s of the Nechako Plateau a l s o are found w i t h i n the Nechako P l a i n . Macroclimate of the study area i s c h a r a c t e r i z e d by abrupt seasonal changes from c o l d , snowy w i n t e r s , to short 37 c o o l summers without a d i s t i n c t dry season. As such i t corresponds to the "Dfc" c l i m a t e type of Koppen, or a microthermal c o n t i n e n t a l sub-boreal type ( K r a j i n a 1965). Chapman (1955) d i s t i n g u i s h e d a Dfb climate i n the v a l l e y s of the Fraser and lower Nechako and Bowron R i v e r s , that i s , a c o o l summer w i t h at l e a s t four months above 10°C. This d i s t i n c t i o n presumably r e f l e c t s the lower e l e v a t i o n and t h e r e f o r e warmer temperatures of these v a l l e y basins. C l i m a t i c data f o r the P r i n c e George a i r p o r t are t y p i c a l of the study area ( B r i t i s h Columbia Department of A g r i c u l t u r e 1976a)(Figure 2.2). Mean annual temperature i s 3.3°C. Annual mean d a i l y minimums and maximums are 2.5°C and 9°C, r e s p e c t i v e l y , w i t h extremes of -50°C recorded i n January and 34.4°C i n J u l y . P r e c i p i t a t i o n averages 621 mm of which 400 mm (63 percent) f a l l s as r a i n . Rain f a l l s i n a l l months of the year and snow has been recorded f o r every month except J u l y and August. However, p r e c i p i t a t i o n i s not evenly d i s t r i b u t e d i n a l l months (Figure 2.2). The n u l l hypothesis of no d i f f e r e n c e i n p r e c i p i t a t i o n between months was r e j e c t e d at P < 0.01 ( x 2 = 100.38, df = 11). The annual m e t e o r o l o g i c a l summary f o r P r i n c e George provides a ready reference to c l i m a t i c records (e.g., Anon 1973). K e l l e y and Farstad (1946) noted the s t r i k i n g v a r i a b i l i t y of the l o c a l c l i m a t e . Strong temperature c o n t r a s t s occur as do abrupt changes i n cloudiness and sequences of wet and dry weather. This v a r i a t i o n i s prob- 37a Figure 2.2 Long term monthly averages of some temperature and p r e c i p i t a t i o n parameters f o r the Pr i n c e George weather s t a t i o n . Source of data: B r i t i s h Columbia Department of A g r i c u l t u r e (1976a). Length of record: 30 years (1941-1970). 38 600 J JAN FEB MAR APR MAY JUNE JULY AUG SEP OCT NOV DEC 39 ably due to i n t e r a c t i o n of moist P a c i f i c a i r from the west and c o o l , dry Po l a r a i r from the north and northeast. Major determinants of t h i s c l i m a t i c regime are l a t i t u d e , e l e v a t i o n , c o n t i n e n t a l l o c a t i o n , and the i n t e r a c t i o n of warm, moist P a c i f i c a i r and c o o l , dry Po l a r a i r . The r e l a t i v e l y even topography r e s u l t s i n n o t i c e a b l e h o r i z o n t a l c l i m a t i c gradients r a t h e r than the marked v e r t i c a l gradients t y p i c a l f o r much of B r i t i s h Columbia. The upland areas to the east ob v i o u s l y are an important determinant of these.gradients, although other f a c t o r s a l s o operate. Of major importance to moose are the gradients of p r e c i p i t a t i o n and c o l d temperature since they i n f l u e n c e c r i t i c a l elements of winter c l i m a t e , v i z . , snow d e p o s i t i o n , c h a r a c t e r i s t i c s and melt. Conventional c l i m a t e records are not a v a i l a b l e f o r these parameters so the gradients must be described i n terms of i n d i c a t o r s such as snow f a l l and mean temperatures. A trend towards c o o l e r temperatures to the north i s evident and p o s s i b l y to the west (Table 2.1). Quesnel has a mean d a i l y temperature of 4.4°C, while P r i n c e George, 110 km north, has 3.3°C; Vanderhoof and Fo r t St. James (50 km n o r t h ) , have temperature means of 2.7°C and 2.3°C r e s p e c t i v e l y . The p r e c i p i t a t i o n gradient i s more pronounced than th a t f o r temperature (Table 2.1). Both r a i n and snow 40 increase from west to east and from south to north. Snowfall i n Prince George i s 127 percent that to the east i n Vanderhoof while s n o w f a l l i n A l e z a Lake i s 204 percent that of Vanderhoof. Pr i n c e George and Ale z a Lake are 8 0 km and 130 km east of Vanderhoof, r e s p e c t i v e l y . Snowfall i n Pri n c e George i s 121 percent that i n Quesnel to the South, and sn o w f a l l i n F o r t St. James i s 101 percent that i n Vanderhoof. R a i n f a l l data show s i m i l a r trends. Table 2.1 C l i m a t i c Parameters f o r the Study Area* C l i m a t i c parameter Climate d a i l y d a i l y s n o w f a l l s t a t i o n * * mean (C) minimum (C) (mm) r a i n (mm) Vanderhoof (680) 2.7 -3.8 1,834 273(60%)*** Fort St. James (686) 2.3 -3.5 1,857 284(60%) P r i n c e George (676) 3.3 -2.5 2,334 400(64%) Quesnel (545) 4.4 -1.8 1,928 361(65%) A l e z a Lake (625) 3.1 -3.0 3,744 557(60%) *Source of data: B r i t i s h Columbia Department of A g r i c u l t u r e (1976a). * * E l e v a t i o n i n m. ** * P r o p o r t i o n of t o t a l p r e c i p i t a t i o n f a l l i n g as r a i n . Within the macroclimate there i s an i n t r i c a t e matrix of microclimates or "climate i n a small space" (Geiger 1966). (Microclimate i s defined as l o c a l combinations of atmospheric f a c t o r s which d i f f e r from macroclimate due to v a r i a t i o n s i n o l a n t cover, tocography, slope p o s i t i o n , and pro x i m i t y to lakes (Daubenmire 19.7:4).) Selected microclimates of the sub-boreal spruce b i o g i o c l i m a t i c zone were studied i n d e t a i l by Wali (1969) and Wali and K r a j i n a (1973). Their s i t e s e l e c t i o n s were based on d i f f e r e n c e s i n canopy d e n s i t y , ground cover, species composition, s o i l c o n d i t i o n s and topography. The study area has been comparatively w e l l examined f o r s o i l s . K e l l e y and Farstad (1946) surveyed s o i l s i n a re c t a n g u l a r area centered on P r i n c e George. Farstad and L a i r d (1954) extended the survey area westward towards Burns Lake, while H o r t i e et a l . (1970) surveyed the upper Fraser River v a l l e y from P r i n c e George eastward to approximately McBride. These repo r t s are c o l l a t e d i n a recent compendium (Keser et a l . 1973). E c o l o g i c a l s t u d i e s by Wali (1969) and a co-worker Revel (1972) r e l a t e d s o i l s w i t h ecosystematic u n i t s (sensu K r a j i n a (1965)) i n the sub-boreal spruce b i o g e o c l i m a t i c zone. Their study area included p a r t of mine. The most recent and comprehensive s o i l survey was conducted j o i n t l y by the B r i t i s h Columbia Land Inventory and the S o i l Survey S e c t i o n , Canada Department of A g r i c u l t u r e (A. Dawson, pers. comm.). These surveys i n t e g r a t e d s o i l s , landforms and vegetation i n a b i o p h y s i c a l approach to land mapping, s i m i l a r to two previous r e p o r t s f o r more we s t e r l y regions of n o r t h - c e n t r a l B r i t i s h Columbia (Runka 1972, C o t i c et a l . 1974). Information from these surveys provided a b a s i s f o r c a p a b i l i t y assessments regarding f o r e s t r y , w i l d l i f e and a g r i c u l t u r e . The f o l l o w i n g d e s c r i p t i o n of 42 s o i l s draws upon Keser et a l . (1973), and m a t e r i a l c u r r e n t l y i n prepa r a t i o n (A. Dawson, pers. comm.). S o i l s of the study area mainly belong to the L u v i s o l i c and P o d z o l i c orders, as defined by the N a t i o n a l S o i l Survey Committee of Canada (1970). Smaller acreages of B r u n i s o l s and Regosols occur on mineral s u b s t r a t e s , and Mes i s o l s and Humisols occur on organic d e p o s i t s . S o i l s i n t e g r a t e the f a c t o r s of time, c l i m a t e , r e l i e f , v e g e t a t i o n and substrate (Jenny 1941). Thus they can provide a meaningful way of s t r a t i f y i n g a heterogeneous environment i n t o homogeneous u n i t s . With the advice of A. Dawson and G. Runka, I s t r a t i f i e d my study area i n t o environmental u n i t s based on moisture regime and s u r f i c i a l d e p o s i t . From t h i s s u b d i v i s i o n , major u n i t s were deri v e d to form the b a s i s of much of the sampling i n t h i s p r o j e c t . The general d i s t r i b u t i o n of s o i l a s s o c i a t i o n s and land forms i s presented i n Figure 2.3. Twenty-three u n i t s were d e l i n e a t e d on mineral substrates and two on organic d e p o s i t s (Table 2.2). Of these, the mesic u n i t s on bas a l t i l l and l a c u s t r i n e sub- s t r a t e s were s e l e c t e d f o r d e t a i l e d study. They represented the l a r g e s t map u n i t s i n the study area, received most of the logging a c t i v i t y , and u n d e r l a i d most of the i n t e n s i v e l y studied winter ranges. These s e l e c t e d environmental u n i t s plus seven others of importance to moose are described b r i e f l y below (taken from Dawson, pers. comm.). 42a Figure 2.3 Oblique a e r i a l photographs i l l u s t r a t i n g the general t e r r a i n and vegetation of the P r i n c e George study area. Photographs taken by K. Sumanik and M. Warren. 43 44 Table 2.2 Major S o i l A s s o c i a t i o n s f o r the Study Area and Their R e l a t i o n s h i p to Parent M a t e r i a l s and Moisture Regimes S o i l a s s o c i a t i o n by moisture regime S u r f i c i a l ; deposit x e r i c mesic h y d r i c L a c u s t r i n e - c l a y s Vanderhoof Pineview Bowron - s i l t s Berman Bednesti Coarse outwash A l i x , Mapes Saxton, Giscome Ramsey, Bear Roaring Peta Beach r i d g e Kluk Gunniza Basal t i l l B a r r e t t Deserters Dominion, Twain A b l a t i o n t i l l C r y s t a l (part) C r y s t a l , Cobb Cobb (part) Shallow t i l l / b e d r o c k Pope Ormrod C l u c u l z , A v e r i l Decker Dragon, Oona S o i l a s s o c i a t i o n not ranked by moisture regime Recent a l l u v i u m McGregor and S t e l l a k o G l a c i o - f l u v i a l Fraser and Nechako Organics Chief (sedge peats) Moxley (sphagum peats) The d e t a i l e d features of a l l u n i t s o c c u r r i n g i n the general study area can be found i n Dawson (pers. comm.): 1. Deserters (Figure 2.4): mesic environment on g r a v e l l y and stony g l a c i a l t i l l d e p o s i t s of v a r i a b l e t h i c k n e s s . Medium to moderately coarse t e x t u r e s ( g s l , s i ) . * Predominantly a d r u m l i n i z e d and/or g l a c i a l grooved t i l l p l a i n with r o l l i n g and h i l l y topography, w i t h i n c l u s i o n s of b e d r o c k - c o n t r o l l e d strong to very steep slopes. E l e v a t i o n 745 - 1220 m. S i x sub-group combinations d e l i n e a t e d (Figure 2.4). Also included i n t h i s u n i t i s a h y d r i c subgroup of the d r i e r B a r r e t t s o i l a s s o c i a t i o n (BA 4), an o r t h i c grey l u v i s o l . This u n i t occurs as f a r westward as Smithers. The d r i e r counterpart i s the * L e t t e r s w i t h i n parentheses are standard a b b r e v i a t i o n s used to d e s c r i b e s o i l t e x t u r e . A b b r e v i a t i o n s are: c - c l a y , f - f i n e , g - g r a v e l , 1 - loam, s - sand, s i - s i l t . 4 4 a F i g u r e 2 . 4 A s c h e m a t i c i l l u s t r a t i o n s h o w i n g t h e m a j o r s o i l a s s o c i a t i o n s i n t h e s t u d y a r e a , a n d t h e i r t o p o g r a p h i c r e l a t i o n s h i p t o e a c h o t h e r . D e r i v e d f r o m d a t a p r o v i d e d b y A . D a w s o n , B r i t i s h C o l u m b i a M i n i s t r y o f A g r i c u l t u r e . DESERTERS- •-CRYSTAL- or COBB -ORMROD- or DECKER l b / MOXLEY CHIEF J . BEDROCK 46 B a r r e t t a s s o c i a t i o n ; the wetter, Dominion and Twain. 2. Pineview (Figure 2.4): mesic environment on clayey g l a c i a l lake d e p o s i t s which vary from up to 10 - 15 m i n t h ickness to shallow d e p o s i t s (l e s s than 1.5 m) over g l a c i a l t i l l . The deposits are s i l t y at depth and g r a d u a l l y grade to f i n e (c, s i c ) and very f i n e (hvc) t e x t u r e s near the surface. Undulating to r o l l i n g topography, w i t h the l a t t e r a s s o ciated w i t h the under- l y i n g d rumlinized g l a c i a l t i l l ; moderately to s t e e p l y s l o p i n g adjacent to the major r i v e r s . E l e v a t i o n a l range i s 610 - 790 m. Seven subgroups d i s t i n g u i s h e d i n the P r i n c e George g l a c i a l lake b a s i n . D r i e r counter- pa r t i s the Vanderhoof a s s o c i a t i o n . 3. Bednesti (Figure 2.4): mesic environment on s i l t y g l a c i a l lake d e p o s i t s which vary from up to 10 - 15 m i n t h ickness to shallow deposits (l e s s than 1.5 m) over g l a c i a l t i l l . Medium to moderately f i n e textured ( s i l , s i c l ) . Undulating, r o l l i n g and h i l l y topography; s t r o n g l y to very s t e e p l y s l o p i n g where the deposits are associated w i t h the Stuart and Bednesti eskers; d i s s e c t - ed adjacent to the main r i v e r s and creeks; shallow to deep k e t t l e s occur at random. E l e v a t i o n a l range i s 610 - 790 m. Four subgroups i d e n t i f i e d . The d r i e r counterpart i s the Berman a s s o c i a t i o n , w h i l e the wetter one i s Bowron. 4. Gunniza (Figure 2.4): mesic environment on g r a v e l l y , cobbly and sandy g l a c i a l lake beach deposits which are u n d e r l a i n by g l a c i a l t i l l a t v a r i a b l e depths from 0.3 to 3 m, w i t h i n c l u s i o n s as deep as 4.5 to 6 m. Very coarse textured (g, gs, g l s , s, I s ) . Gently to s t r o n g l y r o l l i n g and moderately r o l l i n g topography which u s u a l l y conforms to the underlying g l a c i a l t i l l . E l e v a t i o n a l range i s from 730 to 790 m. This a s s o c i a t i o n occurs along the margins of the g l a c i a l lake centered at P r i n c e George, and occurs on the Salmon i n t e n s i v e study area. 5. McGregor and S t e l l a k o (Figure 2.4): both u n i t s are recent s i l t y and sandy l a t e r a l l y accreted f l u v i a l ( a l l u v i a l ) t e r r a c e d e p o s i t s . Medium to very coarse textured ( s i l , 1, f s l , s i , I s , s ) ; o f t e n i n t e r - s t r a t i f i e d ; u n d e r l a i n by sands at shallow depths. Undulating topography. Subject to inundation during f r e s h e t and high water t a b l e seasonally. E l e v a t i o n a l range i s 550 - 760 m. These a s s o c i a t i o n s f l a n k major r i v e r s i n the study area, e.g. Salmon, Fr a s e r and Nechako R i v e r s . In combination w i t h o l d e r g l a c i a l f l u v i a l d e p o s i t s described next, they occur on the c r i t i c a l v a l l e y bottom winter ranges of moose. 47 6. Fraser and Nechako (Figure 2.4): both u n i t s are o l d e r a s s o c i a t i o n s than the above. They are s i l t y to sandy v e r t i c a l l y accreted f l u v i a l d e p o s i t s ; moderately coarse to medium (Nechako) or medium to moderately f i n e (Fraser) textured; u n d e r l a i n w i t h coarse t e x t u r e s at v a r i a b l e depths (Fraser) or by sands at depth of l e s s than 1.5 m. Undulating to n e a r l y l e v e l topography. E l e v a t i o n a l range i s 550 to 670 m. (Fraser) and up to 7 60 m. (Nechako). Two subgroups are o u t l i n e d f o r each a s s o c i a t i o n . 7. Chief and Moxley (Figure 2.4): two organic a s s o c i a t i o n s , the f i r s t r e p r e s e n t i n g d e p o s i t s composed of sedge and associated hydrophytic v e g e t a t i o n ; the second, w i t h mainly sphagnum moss type of vegetation. Two subgroups d i s t i n g u i s h e d f o r each a s s o c i a t i o n , one f o r g e n e r a l l y land-locked map u n i t s (poor or non- e x i s t e n t drainage) and one f o r those associated w i t h creek drainage. Elevation- range i s wide, from 610 to 137 0 m. Important summer h a b i t a t s , e s p e c i a l l y Chief. These u n i t s , e s p e c i a l l y those r e p r e s e n t i n g mesic environments, formed the b a s i s f o r most of the sampling s t r a t a f o r t h i s study. Vegetation of the study area i s p r i m a r i l y coniferous f o r e s t . A r l i d g e ( i n H o r t i e et a l . (1970:14) provided the f o l l o w i n g appropriate d e s c r i p t i o n : The f o r e s t s are composed of a mixture of spruces (white spruce, Engelmann Spruce and t h e i r i n t e r - grades) and a l p i n e f i r w i t h s c a t t e r e d white b i r c h , o c c a s i o n a l trembling aspen, and Douglas f i r . Stumps and r o t t i n g trunks of Douglas f i r suggest that t h i s species had a greater r e p r e s e n t a t i o n i n the recent past. Lodgepole pine occurs i n pure stands and i n v a r y i n g mixtures w i t h spruce and other species f o l l o w i n g f i r e s . Aspen and b i r c h a l s o form pure stands or sands mixed w i t h other species f o l l o w i n g f i r e s . Black spruce and lodge- pole pine, together or s e p a r a t e l y , are found i n bogs. Black cottonwood i s found on a l l u v i a l bottom s o i l s . Although most workers who have studied t h i s f o r e s t type would agree w i t h the above d e s c r i p t i o n , they have 48 named and c l a s s i f i e d i t d i f f e r e n t l y . Whitford and C r a i g (1918) c l a s s i f i e d t h i s f o r e s t as an Engelmann spruce {Picea glauoa engelmanni) - a l p i n e f i r {Abies lasiooarpa) type, but a l s o c a l l e d i t a lodgepole pine ( Firms oontorta) type. They considered t h a t t h i s species of spruce occurred from the lowest v a l l e y s up to 1,220 - 1,520 m. However, K r a j i n a (1969) st a t e d that white spruce ( Pioea g. glauoa) occurs at lower e l e v a t i o n , while white x Engelmann spruce hybrids occur at intermediate e l e v a t i o n s , and the l a t t e r species at higher e l e v a t i o n s (also Taylor (1959), c i t e d i n K r a j i n a (1969)). Forests of the study area would border on Hare and R i t c h i e ' s (1972) "closed f o r e s t " zone of t h e i r b oreal f o r e s t , and i n Rowe's (1973) montane t r a n s i t i o n s e c t i o n (M.4) of the montane f o r e s t region. I n c l u s i o n i n t h i s region of Rowe's rat h e r than the subalpine region was based on the sca t t e r e d presence of i n t e r i o r Douglas f i r (Pseudotsuga menziesii) (Rowe 1972:76). However, Douglas f i r f a i l s to reproduce i t s e l f except on the d r i e r s i t e s (e.g., beach deposits) and n u t r i t i o n a l l y r i c h L i t h o s o l s . K r a j i n a (1959, 1965) defined a sub-boreal spruce zone of h i s Canadian bor e a l f o r e s t b i o g e o c l i m a t i c region that a l s o encompassed the study area. Van Barneveld ( i n C o t i c et a l . 1974) o f f e r e d the f o l l o w i n g c r i t e r i a f o r separating the sub- bore a l spruce zone from the s i m i l a r northern subzone of the Cariboo-aspen (Populus tremuloides)- lodgepole pine/Douglas f i r b i o g e o c l i m a t i c zone: 49 1. l i t t l e or no p o t e n t i a l f o r climax Douglas f i r stands, 2. absence of pinegrass . {Calamagrostis vubescens) , 3. presence of subalpine f i r regeneration below 915 m., 4. continuous and o f t e n r e l a t i v e l y t h i c k moss l a y e r . These are meaningful c r i t e r i a , based on my observations i n the study area. Based on these a t t r i b u t e s , and the d e s c r i p t i o n s by K r a j i n a (1965), and Revel (1972), I b e l i e v e the most a p p l i c a b l e d e s c r i p t i o n i s the sub-boreal spruce zone. Revel (1972) l i s t e d other authors who have studied the s i m i l a r b oreal f o r e s t , and stated that those by Moss (1953a, 1953b, 1955) were most r e l e v a n t . Work by LaRoi (1967) and Annas (1977) can be added to t h i s l i s t . C l a s s i f i c a t i o n of t h i s f o r e s t zone i n t o vegetation subunits has been attempted. K u j u l a (1945, not seen but c i t e d i n Wali and K r a j i n a (1973)) i d e n t i f i e d f o r e s t types i n the area. Subsequently, I l l i n g w o r t h and A r l i d g e (1960) proposed f i v e s i t e types f o r white spruce - subalpine f i r {Abies lasiooarpa) stands based both on dominant and c h a r a c t e r i s t i c understory species. Their nine s i t e types f o r lodgepole pine f o r e s t s i n s o u t h - c e n t r a l B r i t i s h Columbia have counterparts i n the study area, too. Wali (1969) and Revel (1972) conducted s y n e c o l o g i c a l analyses i n the sub-boreal spruce zone. Their c l a s s i f i c a t i o n f o l l o w s the methods and concepts developed by K r a j i n a (1965, 1969). Most r e c e n t l y , d i s s i m i l a r i t y a n a l y s i s was a p p l i e d to A r l i d g e ' s o r i g i n a l data plus new data from n o r t h - c e n t r a l 50 B r i t i s h Columbia ( J . van Barneveld, pers. comm.). Using t h i s h i e r a r c h i c a l d i v i s i v e approach, 41 f i n a l groups or vegetation types were p r o v i s i o n a l l y i d e n t i f i e d . These were combined i n t o 16 composite groupings that contained up to f i v e v e g e t a t i o n types each. Although t h i s recent work i s not yet completed, the p r o v i s i o n a l u n i t s w i l l probably provide the most u s e f u l s u b d i v i s i o n of sub-boreal f o r e s t s . The study area supports t e r r e s t r i a l fauna t y p i c a l of the sub-alpine f o r e s t b i o t i c area of Cowan and Guiget (1973). Mammalian fauna are very much l i k e t h a t of the boreal f o r e s t . The major ungulate i s moose, w i t h year-round 2 d e n s i t i e s of approximately 0.4/km (K. C h i l d , pers. comm.). Mule deer {Odoooileus hemionus) are l i m i t e d i n numbers, probably because of deep snow. Their c r i t i c a l winter h a b i t a t of steep, south-facing slopes with widely spaced mature Douglas f i r i s confined p r i m a r i l y to major r i v e r drainages. Deer are more numerous to the south, east and west of the area. Mountain caribou {Eangifev tarandus montanus) are t r a n s i e n t s . I observed t r a c k s and p e l l e t groups only on two occasions: at Trapping Lake and at Barney Creek, 3 7 km south and 51 km NNE of P r i n c e George, r e s p e c t i v e l y . Caribou winter to the east, southeast and northeast of P r i n c e George. Winter h a b i t a t u t i l i z a t i o n by t h i s species i s c u r r e n t l y under study approximately 90 km east of P r i n c e George (Bloomfield 1976). Other v e r t e b r a t e herbivores whose d i e t s overlap that of moose are beaver {Castor canadensis), v a r y i n g hare {Lepus americanus), porcupine {Hrethizon dorsatum), and r u f f e d grouse {Bonasa umbellus). During the per i o d of f i e l d work, n o t i c e a b l e hare browsing was l o c a l i z e d and. occurred mainly i n r i p a r i a n h a b i t a t s . Porcupine damage i s b e l i e v e d minimal. Ruffed grouse fed e x t e n s i v e l y on w i l l o w buds i n the tops of trees and t a l l shrubs t h a t were mostly beyond the reach of moose. Thus competition between these species and moose i s considered minimal. Representative c a r n i v o r e s - wolf {Canis lupus), coyote (Canis latvans) , black bear {Ursus americanus) , g r i z z l y bear (U. arctos) , Canada lynx {Lynx canadensis), bobcat (L. rufus) - are comparatively abundant; cougar {Felix concolor) are uncommon. Commercially v a l u a b l e furbearers are a l s o abundant. K e l l y (1976) i s studying h a b i t a t u t i l i z a t i o n by marten {Martes americana) i n r e l a t i o n to logging w i t h i n the study area. The area supports a v a r i e d but l a r g e l y unstudied b i r d fauna. Studies by Munro (1947, 1949 and 1955) of b i r d s and mammals f o r the Vanderhoof l o c a l i t y and surveys by S t a n w e l l - F l e t c h e r and S t a n w e l l - F l e t c h e r (1943) of the Driftwood R i v e r v a l l e y are probably a p p l i c a b l e . E f f o r t s by l o c a l birdwatchers represent the major curre n t c o n t r i b u t i o n s to b i r d study. Many o p p o r t u n i t i e s f o r o r n i t h o l o g i c a l research e x i s t as the area i s t r a n s i t i o n a l f o r many species (e.g., Scott et a l . 1976). A l s o , developments such as land c l e a r i n g , and u t i l i t y c o r r i d o r s have encouraged the northward extension of t y p i c a l l y s o u t h e r l y s p e c i e s , e.g., sparrowhawk .(Faloo sparverius), western meadowlark ( Sturnella neglecta) and l a z u l i bunting ( P a s s e r i n a conoena). A g r i c u l t u r a l land i s used by many mi g r a t i n g waterfowl, e s p e c i a l l y Canada geese (Branta canadensis), but the c a p a b i l i t y r a t i n g s f o r waterfowl are g e n e r a l l y very low, w i t h the major l i m i t a t i o n s being reduced marsh edge, i n a p p r o p r i a t e water depth and low n u t r i e n t s t a t u s . Recent surveys i n d i c a t e t h a t production l e v e l s f o r waterfowl are low (W. A. Munro, pers. comm.). Amphibians, r e p t i l e s and i n v e r t e b r a t e s are even l e s s w e l l studied than the avifauna, except f o r the spruce budworm ( C h o r i s t o n e u r a fumiferana) and s e v e r a l other i n s e c t species t h a t damage commercial t r e e species. 2.2 The Primary Study Areas Three primary study areas were s e l e c t e d w i t h i n the general study area described i n s e c t i o n 2.1 (Figure 2.5). These were c a l l e d Eagle, Grove and Salmon. A l l three are important moose winter ranges (K. Sumanik, pers. comm.). A l l three contained h a b i t a t s t y p i c a l of the general study area, i n c l u d i n g burns, immature f o r e s t s , mature f o r e s t s and cutovers. The Eagle winter range was lo c a t e d approximately 4 0 km ENE of P r i n c e George. I t covered approximately 2 3 90 km and measured 2 0 x 3 0 km at i t s g r e a t e s t width and length, r e s p e c t i v e l y . Western and northern boundaries 52a Figure 2.5 Photographs of the Eagle, Grove and Salmon winter ranges.  54 followed the Fraser River upstream from the Willow River to Mokus Creek, near the mouth of the McGregor Ri v e r . The eastern boundary followed roughly the eastern edge of O g i l v i e and Bearman Creek watersheds. Hay Creek, the north shore of Eaglet Lake and the Willow R i v e r , from i t s confluence w i t h Hay Creek to the Frase r , completed the southern boundary of the Eagle range. The e l e v a t i o n a l range was from 590 to 950 m, wit h most of the area l y i n g below 790 m. S o i l s are mostly s i l t y and clayey l a c u s t r i n e d e p o s i t s , t h a t i s , Bednesti, Pineview and Bowron a s s o c i a t i o n s (Figure 2.6). Scattered beach deposits occur near 790 m. At higher e l e v a t i o n s are small areas of g l a c i a l t i l l s of the Deserters a s s o c i a t i o n and i t s wetter counter- p a r t , Dominion. Both recent' and g l a c i a l - f l u v i a l d eposits border the Fraser and Willow R i v e r s , and Hay Creek. Although few organic s o i l s were mapped, many small pockets occur on the study area. Rock outcrops are uncommon. Lakes are small and s c a t t e r e d , though wetlands are p l e n t i f u l . Vegetation i s complex, v a r i e d and o f t e n l u s h . This i s due to a combination of moisture, undulating topography, f e r t i l e s o i l s , and a wide range and larg e area of logging and w i l d f i r e (Figure 2.7). The most n o t i c e a b l e feature i s 2 the 90 km Eaglet Lake burn. The area was apparently burned t w i c e — o a 1932 and oa 1937—.and coniferous regeneration i s s t i l l s c a t t e r e d . Regeneration f a i l u r e 54a Figure 2.6 A s o i l a s s o c i a t i o n map of the Eagle study area. Derived from maps provided by A. Dawson, B r i t i s h Columbia M i n i s t r y of A g r i c u l t u r e .  55a Figure 2.7 A f o r e s t cover map of the Eagle study area. Derived from f o r e s t cover-type maps of the Inventory D i v i s i o n , B r i t i s h Columbia Forest S e r v i c e .  a t t e s t s to what must have been severe f i r e s . Mature spruce-subalpine f i r f o r e s t s cover the northern o n e - t h i r d of the area. The remainder i s cutover land. Among the c u t t i n g p r a c t i c e s represented are d i a m e t e r - l i m i t and s i n g l e t r e e s e l e c t i o n c u t t i n g , cut and leave s t r i p s , c l e a r c u t and burn, c l e a r c u t w i t h pre- and p o s t - s c a r i f i c a t i o n and seed block logging. A small black cottonwood {Populus balsamifera tviehoeavpa) cutover occurs near the confluence of the Willow and Fraser R i v e r s . Some of the e a r l i e s t logging i n the Pri n c e George Forest D i s t r i c t occurred here, and i t i s one of the few places i n c e n t r a l B r i t i s h Columbia t h a t was r a i l - logged. The Eagle contained a wide v a r i e t y of moose h a b i t a t s . I t supported both r e s i d e n t and migrant herds, w i t h at l e a s t one reported m i g r a t i o n route from the McGregor Pl a t e a u , and the watersheds of A v e r i l , Limestone, and Olsson Creeks, southward across the Fraser River onto the study area. T y p i c a l herd s t r u c t u r e and a r e l a t i v e estimate of abundance are presented i n Table 2.3 (K. C h i l d , pers. comm.). The dominant land use i s timber h a r v e s t i n g . Winter logging i s commonest as the wet s o i l s create access problems f o r mechanized equipment i n summer. The small forestry-based communities of Willow River and Giscome are near to the mouth of the Willow River and the western end of Eaglet Lake, r e s p e c t i v e l y . A l a r g e sawmill operated i n Table 2.3 58 Estimated Relative Abundance and Herd Structure f o r W i n t e r i n g Moose on the I n t e n s i v e Scudv A r e a s , 1964-65 to 1975-76 Study area Winter Relative abundance* Herd structure (%) Sample size b u l l s cows calves unclass moose time** Eagle 1964-65*** 1965-66 1.22 41 42 17 «. 93 76(D) 1966-67 2.80 29 56 16 70 25(D) 1967-68 1.15 28 48 24 - 82 71(D) 1968-69*** 1969-70*** 1970-71 2.60 23 - - 77 13 5(J) 1971-72 0.97 26 47 27 1 90 93(D) 1972-73*** 1973-74 0.69 12 56 33 — 52 rri-1974-75 m 19 61 19 - 36 1975-76 1.00 14 57 29 - 14 14 (J) Grove 1964-65 1.47 - 16 16 68 25 17 (J) 1965-66 0.53 39 43 18 - 28 53(D) 1966-67 0.02 100 1 45(M) 1967-68 0.54 31 41 27 - 51 94(D) 1968-69 1.41 32 49 18 1 82 58(J) 1969-70*** 1970-71 0.82 22 - - 78 9 H ( J ) 1971-72 0.98 26 48 25 - 87 89(D) 1972-73*** 1973-74 1.09 45 39 3.6 _ 140 m 1974-75*** 1975-76 0.30 21 50 .29 - 24 80(J) Salmon 1964-65 1.93 _ 10 10 81 31 16(J) 1965-66 0.85 20 46 29 6 35 41(D) 1966-67*** 1967-68 0.94 18 54 29 - 61 65(D) 1968-69*** 1969-70*** 1970-71 m 33 66 - - 9 m(J) 1971-72 1.42 38 40 18 4 50 35(D) 1972-73*** 1973-74 0.90 21 55 19 2 85 ID 1974-75 m 29 54 15 1 m 1975-76 1.00 37 40 23 - 30 30(J) Summary : E3gle 1.49(2.11)^ '24(29) 52(19) 24(17) Grove 0.80(1.39) 31(24) 41(34) 21(13) Salmon 1.17(1.08) 28(20) 47(56) 20(19) *No. of moose seen/min. helicopter survey. **Time i n min. with month of survey in parenthesis (J - Jan., M - March). ***Data missing or surveys not conducted. +Missing. -H-Mean (range). Giscome u n t i l 1974 a f t e r which both i t and the v i l l a g e c l o s e d . Logs are now hauled to m i l l s i n Pri n c e George and Upper Fraser. A g r i c u l t u r a l development i s l i m i t e d . Several small farms border the north and west shores of Eaglet Lake. The burn has been t r a d i t i o n a l l y u t i l i z e d by a small number of c a t t l e . A community pasture development f o r the burn i s c u r r e n t l y i n progress. Pasture development and maintenance could prove d e t r i m e n t a l to moose. The burn and logged over lands are t r a d i t i o n a l , well-used hunting areas. This r e s u l t s from the r e l a t i v e abundance of animals, good access, e s p e c i a l l y a f t e r f r e e z e - up, and pro x i m i t y to P r i n c e George. Few other r e c r e a t i o n a l past-times occur except f o r some nature-viewing, berry- p i c k i n g and l i m i t e d bathing and boating along Eaglet Lake. The second i n t e n s i v e study area was Grove, so-named a f t e r a f o r e s t f i r e i n August, 1961 that burned 2 approximately 320 km of mostly cutover f o r e s t s . Grove l i e s 30 km due east of Pri n c e George (Figure 2.1) and covers 2 approximately 450 km . The area i s oval-shaped w i t h the long a x i s (27 km) o r i e n t e d north-south and the short a x i s (19 km), east-west. The western boundary roughly f o l l o w s the 762 m contour (the east boundary of the Pri n c e George S p e c i a l Sale Area, and a l s o the upper l i m i t of the pro- g l a c i a l lake) from Buckhorn Lake northwards to Tsadestsa Creek; then e a s t e r l y and southerly up the Willow River to 60 the Willow-Kale Forest Development Road; then w e s t e r l y along t h i s road to Buckhorn Lake. This southern boundary, l i k e the others f o l l o w s a topographic/physiographic break - an abandoned outwash channel t h a t separates the Mt. George upland from the Tabor Mountain upland. E l e v a t i o n a l range i s from 640 to 1,260 m. Compared to Eagle, most of Grove i s above the p r o g l a c i a l lake b a s i n . Thus s o i l s belong mostly to the Deserters a s s o c i a t i o n - g l a c i a l t i l l i n a moist environment (Figure 2.8). Upland areas to the west and south have shallow t i l l over bedrock (Decker a s s o c i a t i o n ) , and rock outcrops make up to 20% of the high e l e v a t i o n map u n i t s . G l a c i a l - f l u v i a l d e p o s i t s are e s p e c i a l l y common along the southern boundary but are a l s o found at higher, more c e n t r a l l o c a t i o n s . Presumably, these were meltwater deposits o r i g i n a t i n g from i c e wastage. L a c u s t r i n e deposits occupy the northern one-quarter of the study area, b a s i c a l l y that land s i t u a t e d north of Highway 16 and below 790 m. These are mainly Bowron s i l t s and Pineview c l a y s . Beach depo s i t s and d e l t a s mark the boundary between l a c u s t r i n e and t i l l s u b s t r a t e s . Recent a l l u v i u m , McGregor and S t e l l a k o a s s o c i a t i o n s , i s present along the Willow R i v e r . Organics are more common than at Eagle but are confined mainly to the southwest corner of the study area. Many streams d r a i n Grove but lakes are s c a t t e r e d and s m a l l , except f o r Buckhorn and Tabor Lakes. 60a Figure 2.8 A s o i l a s s o c i a t i o n map of the Grove study- area. Source of data: see Figure. 2.6. 61 GROVE STUDY AREA SOILS 62 Most of the p l a n t cover i s s e r a i , due to the lar g e f i r e i n 1961 (Figure 2.9). Willow {Salix spp.) and paper b i r c h {Betula papyvifera) are the major woody p l a n t s i n the burn but c o n i f e r s are regenerating. In the southern part of the burn, dense stands of immature lodgepole pine occur. Mature f o r e s t s of mixed pine, spruce and subalpine f i r are found at the north and e a s t - c e n t r a l p a r t s . Compared w i t h Eagle, the Grove area has few cutover h a b i t a t - t y p e s as most of them were burned i n 1961. For moose, the Grove Burn i s p r i m a r i l y a winter range. Resident animals are present but not as abundant as on Eagle. Although tagging data are l a c k i n g , most of the w i n t e r i n g moose probably come from the upper part of the Willow River v a l l e y and the surrounding uplands southward. P o s s i b l y before the lands surrounding the Fraser River were cl e a r e d and s e t t l e d , moose wintered there i n s t e a d of on Grove. T y p i c a l composition and r e l a t i v e abundance of the winter moose herd are presented i n Table 2.3. Winter d i s t r i b u t i o n of moose on Grove was obviously non-random. One of the main reasons f o r s e l e c t i n g t h i s study area was to determine why t h i s was so. Recreation i s the p r i n c i p a l land-use on t h i s area apart from i t s value as a moose winter range. As access i s good throughout much of the burn, i t i s w e l l used by hunters. A d o w n h i l l s k i f a c i l i t y i s lo c a t e d along Highway 16, and cross-country s k i i n g i s commonly pursued throughout the 62a Figure 2.9 A f o r e s t cover map of the Grove study area. Source of. data: see Figure 2.7. 6 3 GROVE STUDY AREA VEGETATION r — i ^ , . .. » > Major highway EZD Conifer forest = Mature (>80yr.) Secondary road Sk, „ , ' ™ a t u r f ( 2 1 - 8 ° y f J — S t u d y area boundary KSJ Mixed forest (>21yr.) — — — Deciduous forest A Aspen B Birch Li^] Shrub type : Burn ESi] Swamp * S j Shrubby vegetation Cultivated Cutover = Clearcut Partial cut 2 1 o IHHMHHI KILOMETRES burned-over area. T r a i l - r i d e r s , snowmobilers, and n a t u r a l i s t s a l s o use the area. D i f f i c u l t i e s i n accommodating such a d i v e r s e range of outdoor a c t i v i t i e s have been encountered. An o r g a n i z a t i o n has been formed to t r y to harmonize the various r e c r e a t i o n a l demands and s t i l l minimize disturbance to w i n t e r i n g moose. The t h i r d i n t e n s i v e study area was c a l l e d the Salmon, t a k i n g i t s name from the r i v e r that flowed through t h i s important winter range (Figure 2.1). S i t u a t e d approximately 3 0 km north of Prince George, i t measures approximately 3 0- x 10-km, s t r a d d l i n g the Salmon River and Merton Creek from the Highway 97 road bridge at Salmon V a l l e y to approximately 2 5 km up Merton Creek. This rectangle i s o r i e n t e d northwest to southeast. The Salmon 2 study area covers about 300 km and ranges i n e l e v a t i o n from 610 to 780 m. Compared w i t h Eagle and Grove, which are r e l a t i v e l y d i s c r e t e geographic u n i t s , the Salmon area i s considered as a subsample of a l a r g e r area. From a s o i l s and landform p e r s p e c t i v e , the Salmon area i s more heterogeneous than Eagle and Grove (Figure 2.10). Beginning at the lowest e l e v a t i o n i n the Salmon River v a l l e y at 610 m are the recent a l l u v i a l m a t e r i a l s that form the r i v e r banks. Above these S t e l l a k o deposits are beach r i d g e s intermixed w i t h g l a c i a l t i l l and clayey l a c u s t r i n e sediments. A t r a n s i t i o n a l zone occurs above t h i s , between the Deserters and Pineview s o i l a s s o c i a t i o n s . 64a Figure 2.10 A s o i l a s s o c i a t i o n map of the Salmon study- area. Source of data: see Figure 2.6.  66 This v a r i e t y i s expressed i n smaller map u n i t s than are found at the Eagle and Grove study areas. Vegetation on Salmon i s a l s o d i v e r s e (Figure 2.11). Stands of white spruce and black Cottonwood i n various developmental stages grow on the f l o o d p l a i n of the Salmon Rive r . At the southeastern end of the area i s an extensive aspen and aspen-conifer stand t h a t developed a f t e r a w i l d f i r e approximately 75 years ago. The northeastern area i s p r i m a r i l y pine and pine-spruce mixtures w i t h smaller areas of immature lodgepole pine. Old cutovers occur i n the v a l l e y bottom, mostly at the northeast end of the area. Other commercial f o r e s t stands are c u r r e n t l y being logged so that p r o p o r t i o n of cutovers w i l l i n c r ease. The Salmon River v a l l e y i s one of the major moose ranges i n the study area. Complexes of summer and winter h a b i t a t s occur over much of i t , but winter range predominates. A major mig r a t i o n that crosses Highway 97 south of Summit Lake i n d i c a t e s most w i n t e r i n g moose come from the McGregor uplands. The a v a i l a b l e data f o r w i n t e r i n g moose are summarized i n Table 2.3. Logging occurs on the study area. Formerly, timber was rough sawn at a m i l l near the mouth of Merton Creek, and trucked to Pri n c e George f o r d r y i n g , p l a n i n g , and marketing. This small m i l l settlement i s abandoned and logs are now hauled to P r i n c e George f o r processing. A small summer ranch i s s i t u a t e d approximately 3 km west of 66a Figure 2.11 A f o r e s t cover map of the Salmon study area. Source of data: see Figure 2.7. S A L M O N STUDY A R E A 6 8 Highway 97, along the north side of the Salmon Ri v e r . To develop pasture, the present trembling aspen f o r e s t i s being cut down and the growth of shrubs and r e s p r o u t i n g aspen c o n t r o l l e d . These a c t i v i t i e s w i l l reduce the area's s u i t a b i l i t y f o r moose. As w i t h the other study areas, the Salmon range provides many r e c r e a t i o n a l o p p o r t u n i t i e s . I t i s a w e l l known and a c c e s s i b l e hunting l o c a l e . The f a l l m i g r a t i o n of moose across Highway 97 onto the area i s almost legendary w i t h P r i n c e George r e s i d e n t s . The many stands of v a c c i n i a provide f i n e b e r r y - p i c k i n g , w h i l e the Salmon River o f f e r s canoeing and f i s h i n g . One a c t i v e t r a p l i n e i n cludes p a r t of the study area. 2.3 The Secondary Study Areas In a d d i t i o n to the i n t e n s i v e l y studied winter ranges (primary study a r e a s ) , other l o c a t i o n s were sampled f o r various components of the p r o j e c t (Table 2.4). S e l e c t i o n of these secondary study areas was governed by the type of components, the analyses conducted, the lack of c e r t a i n c h a r a c t e r i s t i c s on the i n t e n s i v e areas, and by the need to gain a broad p e r s p e c t i v e f o r the study. Thus a wide range of s o i l s , c l i m a t i c regimes, h a b i t a t - t y p e s and cutovers was v i s i t e d . Locations of both primary and secondary areas are shown i n Figure 2.1. Data f o r food h a b i t s were c o l l e c t e d throughout the e n t i r e study area, i n c l u d i n g the primary areas. S p e c i f i c l o c a t i o n s of rumen samples are described i n Appendix E. Table 2.4 Types of Analyses Conducted on the Study Areas Topic s t u d i e d * Study area Habitat P l a n t Browsing, Names use succession n u t r i e n t s Bedding Climate Primary study areas: Eagle X X X X X Grove X X X X X Salmon X X X X X Secondary study areas: Bowron X X Found Lake X Fyfe X Limestone X McGregor X McKenzie X X X Pineview X S h e l l X Swamp X Teardrop X Te l a c h i c k X X Torpy X Whites X *Food h a b i t s were based on samples c o l l e c t e d over the general study area. 70 2.4 H i s t o r y of Moose D i s t r i b u t i o n and Abundance The f o l l o w i n g h i s t o r i c a l account was ex t r a c t e d from Hatter's (1950) comprehensive treatment of the t o p i c . Apparently, moose were absent or very low i n numbers p r i o r to and i n c l u d i n g the f i r s t h a l f of the 19th century. MacKenzie (190 3) saw moose on the east slope of the Rockies but not i n the Parsnip, Fraser or Blackwater drainages to the west. Harmon (19 03), who t r a v e l l e d e x t e n s i v e l y between t r a d i n g posts on S t u a r t , Fraser and MacLeod Lakes, reported a few moose. Hatter (1950:28) concluded that these s i g h t i n g s r e f e r r e d to the northeastern p a r t of New Caledonia adjacent to the Rocky Mountains. Brooks (1928) r e l a t e d t h a t r e l i a b l e r e p o r t e r s b e l i e v e d moose to be absent i n the Prince George area " i n t h e i r f a t h e r ' s time"--ca. 1800 to 1850. Moose d e f i n i t e l y i n h a b i t e d the n o r t h - c e n t r a l region by the l a s t h a l f of the 1800's. In 1862, Anderson (1867) knew of moose s t r a y i n g i n the area between F o r t George and Bowron Lake, and r a r e l y as f a r west as Fo r t George. E v i d e n t l y , a moose was k i l l e d at Fraser Lake and another at Ch i n l a c , near the j u n c t i o n of the Stuart and Nechako R i v e r s . McCabe and McCabe (1928a, b) summarized the h i s t o r y and statu s of moose i n the Bowron Lake Game Reserve. They described moose being hunted i n the 1870's and 1890's i n t h i s Reserve. Cast a n t l e r s and sig n were a l s o found i n the 1890's around Ahbau Lake. 71 In the e a r l y 1900's, records of moose seen and shot became more p l e n t i f u l than p r e v i o u s l y (Hatter 1950). Walker (1972:17) r e c a l l e d that the f i r s t moose was shot at Princ e George i n 1914. The increased number of s i g h t i n g s c o r r e l a t e s w i t h increased settlement by Europeans. Therefore i t i s p o s s i b l e t h a t moose were as common p r i o r to 1900 as they appeared to be i n the e a r l y 1910's. Nevertheless, t h e i r r a p i d range extension and population i r r u p t i o n from o a . 1900 to 1950 was a remarkable b i o l o g i c a l phenomenon. The general p a t t e r n of extension was from headwaters of the Fraser River west to Francois Lake and subsequently southward and westward. Both the appearance and abundance of moose i n the n o r t h - c e n t r a l region were a t t r i b u t e d to f o r e s t succession caused mainly by increased man-induced f o r e s t f i r e s (Hatter 1950). I n i t i a l expansion was l i k e l y due to these v a s t l y increased s e r a i shrub communities. The l a t t e r spread was due to a "forced expansion": moose ranged f u r t h e r a f i e l d as forage became scarce i n regenerating f o r e s t s . Hatter (1950) i d e n t i f i e d that peak populations probably occurred i n the 1920's and 1930's, based on many accounts of l o c a l abundance th a t occurred at t h i s time. S i m i l a r l y , h i s choice of f i r e as the main causative agent was based on a c o l l a t i o n of h i s t o r i c a l accounts from the period and the preceding 40-50 years. In a d d i t i o n to n a t u r a l f i r e s were those set by 72 Indians, to a t t r a c t game and provide forage f o r horses; by s e t t l e r s , to c l e a r land f o r farming; by r a i l w a y and road c o n t r a c t o r s , to remove s l a s h and d e b r i s ; and by miners, to f a c i l i t a t e prospecting. Although a l l the foregoing evidence i s c i r c u m s t a n t i a l , the r o l e of f i r e i n the expansion and abundance of moose i s the most p l a u s i b l e e xplanation. Moose numbers d e c l i n e d a f t e r the 1920-1930 peak. However, the harvest records from 1950 onwards suggest that the d e c l i n e has not been to those l e v e l s of the e a r l y 1900's. Current estimate d e n s i t i e s of moose i n the general 2 study area are estimated at 0.4/km (K. C h i l d , pers. comm.). 3. HABITAT USE AND SELECTION 3.1 I n t r o d u c t i o n H a b i t a t i s defined as the place where an animal or p l a n t l i v e s (Odum 1971). Occupation of the h a b i t a t i s h a b i t a t use. Since moose occur throughout the sub-boreal f o r e s t , v i r t u a l l y a l l of i t can be considered moose h a b i t a t . Moose do not occupy these f o r e s t s uniformly. They use the various s e r a i stages and types of f o r e s t stands to vary i n g degrees. By sampling defined u n i t s of sub-boreal f o r e s t s , r e l a t i v e use of these u n i t s i s determinable. The r e l a t i v e importance of these various u n i t s , or h a b i t a t - types, can be assessed from t h i s d e s c r i p t i v e i n f o r m a t i o n . Assessments then all o w w i l d l i f e b i o l o g i s t s and f o r e s t e r s to i d e n t i f y h a b i t a t u n i t s that may r e q u i r e s p e c i a l considera- t i o n i n management. However, data on h a b i t a t use are o f t e n inadequate to i d e n t i f y f a c t o r s r e s p o n s i b l e f o r d i f f e r e n t i a l use of h a b i t a t - t y p e s . Sometimes these data are i n t e r p r e t a b l e but data on h a b i t a t s e l e c t i o n are, i n the long run, most u s e f u l i n t h i s regard. S e l e c t i o n d i f f e r s from use. Habitat use records r e l a t i v e occupancy of h a b i t a t - t y p e s . I t does not de f i n e s e l e c t i o n since the a v a i l a b i l i t y of h a b i t a t - t y p e s i s not 74 u s u a l l y measured. S e l e c t i o n i m p l i e s choice. I t can therefore provide i n s i g h t i n t o what f a c t o r s t h a t moose r e l a t e to when occupying h a b i t a t s . I t i s these f a c t o r s that managers must address to manage e f f e c t i v e l y . Use i s d e s c r i p t i v e while s e l e c t i o n i s p r e d i c t i v e . The term " h a b i t a t - t y p e " has s e v e r a l meanings. In t h i s t h e s i s , i t i s used to describe u n i t s of the landscape that are c h a r a c t e r i z e d by obvious d i f f e r e n c e s i n v e g e t a t i v e cover or by obvious d i f f e r e n c e s i n p o s i t i o n on the land- scape. Common ha b i t a t - t y p e s are mature coniferous f o r e s t , upland burn, p a r t i a l cutover, deciduous f o r e s t , swamp, and c l e a r cut. D e f i n i n g these types c a r r i e s the i m p l i c i t assumption t h a t they are meaningful to moose. The term h a b i t a t - t y p e i s not used i n the sense of Daubenmire (1959), that i s , to describe a land u n i t t h a t i s capable of producing a c e r t a i n kind of climax v e g e t a t i o n . Patterns of h a b i t a t use have been described f o r most of the circumpolar range of moose. North America and Scandinavia have received p a r t i c u l a r a t t e n t i o n . The Quebec symposium on moose ecology (Bedard et a l . 1974) p r e s e n t l y provides the most recent and comprehensive summary on t h i s t o p i c since Peterson's (1955) book. Hab i t a t use by moose i n B r i t i s h Columbia has been v i r t u a l l y unstudied. Much u s e f u l , general information was thoroughly c o l l a t e d and synthesized by Hatter (1950). Baynes (1956) contained general remarks on h a b i t a t use. Several r e p o r t s such as those by R i t c e y (1967), Sumanik and Warren (1968), and Eastman (1974a) d e a l t w i t h h a b i t a t use but t h e i r data have not been published. The major exception i s the recent t h e s i s by S i l v e r (1976) on h a b i t a t use by moose i n northeastern B r i t i s h Columbia. Therefore, the f i r s t o b j e c t i v e of t h i s phase of the study was to describe h a b i t a t use by moose, w i t h p a r t i c u l a r reference to cutovers, burns and f o r e s t s . Given the dearth of i n f o r m a t i o n , a synoptic approach was judged to be the best approach. Emphasis was on sampling a wide v a r i e t y of ha b i t a t - t y p e s w i t h i n the study area boundaries so th a t use patterns could be g e n e r a l i z e d . The primary technique used to determine h a b i t a t use was p e l l e t group counting. The second o b j e c t i v e of t h i s phase of the study was to examine h a b i t a t s e l e c t i o n . This o b j e c t i v e r e q u i r e d complete sampling of an area without regard to where moose occurred. This approach was undertaken through i n t e n s i v e study on the Salmon winter range. The s i t e was s e l e c t e d as t y p i c a l of moose h a b i t a t i n the study area. I t included a major winter h a b i t a t , the bottomlands of the Salmon River v a l l e y , w i t h an adjacent upland that was l a r g e l y unlogged. S p e c i f i c boundaries of the i n t e n s i v e study were those of a timber s a l e h a r v e s t i n g l i c e n s e . Thus I t r i e d to assess h a b i t a t s e l e c t i o n i n an area before logging. I t would be u s e f u l to conduct follow-up surveys a f t e r l ogging. 75a Figure 3.1 Photographs i l l u s t r a t i n g logged h a b i t a t s i n sub-boreal f o r e s t s : a) s e l e c t i v e , cut and leave, and c) c l e a r c u t .  77 3.2 Methods 3.2.1 The Synoptic Survey The method of counting p e l l e t groups was developed from a review of r e l e v a n t a r t i c l e s (e.g. Neff 1968, Smith et a l . 1969) and a p r e l i m i n a r y t r i a l i n the f i e l d . The two standard p l o t shapes, c i r c l e s and s t r i p s , were tes t e d using three commonly used diameters and widths of b e l t s (Table 3.1). The checking time, recorded to the nearest second, was measured from the s t a r t of one p l o t to the s t a r t of the next. Thus the time included both t r a v e l l i n g and p l o t - reading. With the b e l t t r a n s e c t s , each p l o t was re-checked f o r any missing p e l l e t groups. The t r i a l was located i n the Grove Burn study area. A t o t a l of 1,524 m of t r a n s e c t or 50 c i r c u l a r p l o t s was t a l l i e d f o r each p l o t shape-area combination. Based on these sources of in f o r m a t i o n , I s e l e c t e d a 1.5- x 30.5-m b e l t p l o t f o r each person on the synoptic surveys. I t was the qui c k e s t of a l l combinations te s t e d (Table 3.1) and, of the b e l t p l o t s , was the only one i n which no groups were missed. C o e f f i c i e n t of v a r i a t i o n (CV) compared favourably w i t h that of other t r i a l p l o t s i z e s and shapes (24 percent vs. a mean CV of 24.3 percent). To determine winter h a b i t a t use, p e l l e t groups were counted a f t e r snow melt u n t i l ground vegetation obscured p e l l e t s , u s u a l l y from mid-May to the end of June, i n 1972 and 1973. P e l l e t groups were t a l l i e d i n contiguous 78 Table 3.1 Results from T r i a l P e l l e t Group Survey; Time/Plot and Number of Groups/Plot.* P l o t s i z e and shape width x le n g t h of b e l t p l o t s (m) r a d i i of c i r c u l a r p l o t s (m) C r i t e r i a examined 1.5x30.5 3x30.5 6.1x30.5 1.1 1.7 3.6 mean time/ p l o t * * 44(10.6)*** 168(33.2) 205(56.6) 49(11.3) 53(11.3) 85(25.5) mean no. groups/ p l o t 0.65 1.47 3.06 0.10 0.21 0.66 c o e f f i - c i e n t of v a r i a t i o n mean no. groups/ha 23 26 27 40 37 27 no. of groups missed 0 3 9 n d v nd nd + p l o t area (sq/m) 44.5 93.1 186 4.05 8.09 40.5 sample s i z e 49 49 49 50 50 50 *Data i n Appendix Table D - l . **Includes t r a v e l time. ***Mean (sd). + n d = no data. 3- x 15.2-m b e l t p l o t s (46 sq m), w i t h two people each surveying a 1.5- x 15.2-m area. U s u a l l y , two or more tr a n s e c t s were run f o r a t o t a l of at l e a s t 305 m each i n each homogeneous type, y i e l d i n g a t o t a l sample area of 1,830 sq m per type. The s i t e and d i r e c t i o n of t r a n s e c t s were s e l e c t e d p r i o r to f i e l d work to minimize b i a s i n t r a n s e c t l o c a t i o n . Data from both years were combined and expressed as mean number of accumulated p e l l e t groups per ha i n each h a b i t a t . U s u a l l y p e l l e t groups deposited during the preceding winter were e a s i l y aged as such. The autumn l e a f f a l l u s u a l l y covered p e l l e t s from previous w i n t e r s . Also p e l l e t s from the preceding winter were co l o r e d d i f f e r e n t l y from those deposited i n previous w i n t e r s . F r e e z i n g , thawing, and decomposition speeded up the r a p i d d i s i n t e g r a t i o n of p e l l e t s and l i k e l y c o n t r i b u t e d to the observed d i f f e r e n c e s i n c o l o r . Summer and s p r i n g feces were r e a d i l y d i s t i n g u i s a b l e from f a l l and wi n t e r p e l l e t s . Thus p e l l e t group data were assumed to r e f l e c t h a b i t a t use during the approximately preceding e i g h t months. P e l l e t group data were expressed on a per hectare b a s i s r a t h e r than as moose-days per hectare. The conversion to moose-days assumes th a t d a i l y d e f e c a t i o n r a t e s are known, but the published l i t e r a t u r e shows th a t the conventional r a t e of 13 groups per day i s i n v a l i d . Timmermann (1974: 616-617) documented d a i l y r a t e estimates that ranged from 80 10.3 (Le Resche 1970) to 32.2 (Le Resche and Davis 1971). Reasons f o r these d i f f e r i n g estimates probably r e l a t e to i n c o r r e c t aging of p e l l e t groups (Le Resche and Davis 1971), d i e t (Smith 1964), and s e x - s p e c i f i c r a t e s of d e p o s i t i o n (Des Meules 1968).. C a r e f u l study by (Franzmann et a l . 1976) documented l a r g e d i f f e r e n c e s i n d e p o s i t i o n r a t e s between male and female moose. Because sex and age s t r u c t u r e of a sampled moose herd i s not always known (and t h i s can vary w i t h i n a s i n g l e w i n t e r season), and since the r e l a t i o n s h i p between d i e t and d e f e c a t i o n r a t e i s not e s t a b l i s h e d , the conversion of p e l l e t group data to moose-days i s misleading. For these reasons, p e l l e t group data were not converted to moose-days. 3.2.2 P e l l e t Group Counting Methods f o r the D e t a i l e d Survey c l u s t e r s of three c i r c u l a r p l o t s (r = 1.1 m) at s t a t i o n s along t r a n s e c t s . The layout of t r a n s e c t s and spacing of s t a t i o n s along t r a n s e c t s followed methods o u t l i n e d by Smith et a l . (1969). The number of s t a t i o n s per t r a n s e c t was estimated using the formula: For t h i s survey, p e l l e t groups were counted i n m m number of s t a t i o n s per t r a n s e c t , average time i n hours to read a s t a t i o n , C 2 average t r a v e l l i n g time i n hours, 81 2 Sw = variance among s t a t i o n s on the same t r a n s e c t , 2 and S = variance among a l l s t a t i o n s . Variance estimates were obtained from p r e l i m i n a r y f i e l d t r i a l s . Time estimates were based on Smith et a l . (1969), w i t h an upward r e v i s i o n of C 2 to four h^due to the i n a c c e s s i b i l i t y of the study area. The number of t r a n s e c t s was determined by the f o l l o w i n g formula: n = G/(C 2 + C^m), where C^, and m are as above, G = t o t a l number of a v a i l a b l e man-hours, and n = number of s t a t i o n s per t r a n s e c t . Transects were randomly s e l e c t e d from a l l p o s s i b l e t r a n s e c t s , given a separation of 161 m to minimize overlaps. North and south ends of these t r a n s e c t s were permanently marked as were s t a t i o n centers along each t r a n s e c t . D e t a i l s are provided i n Bonar et a l . (1975) . 3.2.3 The A e r i a l Surveys D e t a i l e d monitoring of h a b i t a t use was accomplished through a e r i a l surveys across the Grove study area from January.1972 to May 1973. Using a Cessna 185 a i r c r a f t , t r a n s e c t s were flown monthly at 3.2 km i n t e r v a l s across the study areas at 90 - 150 m above the ground. The same f l i g h t paths were followed each month. The s t a r t and end of each t r a n s e c t were l o c a t e d to the nearest 7 5 m, and i t s . d u r a t i o n 82 was timed with a stopwatch, to the nearest second. Two observers, seated on e i t h e r side of the a i r c r a f t recorded moose and t h e i r t r a c k s on p o r t a b l e tape recorders. Observations were recorded to the nearest second from the s t a r t of each t r a n s e c t . Moose were c l a s s i f i e d as e i t h e r young, a d u l t - unknown sex, cow, l a r g e - , medium-, or s m a l l - a n t l e r e d b u l l , or u n c l a s s i f i e d . Tracks were d i s t i n g u i s h e d as o l d (at l e a s t one week or before the l a s t snowfall) or f r e s h (made since the previous s n o w f a l l ) . Aging of t r a c k s was f a c i l i t a t e d by f l y i n g w i t h i n one or two days a f t e r a s n o w f a l l (Figure 3.2). Track abundance was recorded s u b j e c t i v e l y as few, moderate or p l e n t i f u l . Examples of f l i g h t summary sheets and t r a n s e c t data are appended i n Tables D-2 and D-3. A l l data are on f i l e at the W i l d l i f e Research S e c t i o n , F i s h and W i l d l i f e Branch, Parliament B u i l d i n g s , V i c t o r i a , B.C. Several precautions were taken to standardize the surveys. Most f l i g h t s were made i n the morning during calm, s t a b l e a i r c o n d i t i o n s w i t h a high overcast cloud cover. Stable a i r reduced the problem of d r i f t , w h i l e shadowless or faint-shadow l i g h t c o n d i t i o n s minimized the problems of both g l a r e and poor v i s i b i l i t y . Transect t i e - p o i n t s were conspicuous, d i s c r e t e and e a s i l y defined from the a i r , e.g., road j u n c t i o n s , bridge c r o s s i n g s , boundaries and corners of logged areas. A d d i t i o n a l l y , s e v e r a l conspicuous p o i n t s were noted along the t r a n s e c t s to check on d r i f t . 82a Figure 3.2 The r e l a t i o n s h i p between d a i l y s n o w f a l l and the t i m i n g of the a e r i a l t r a n s e c t surveys, January 1972 to May 1973. 83 A TRACE SNOWFALL ' f FLIGHT DATE 104 84 A i r photo mosaics (approximately 1:50,000) proved e s p e c i a l l y h e l p f u l i n keeping the a i r c r a f t on course. The t r a n s e c t s were p o s i t i o n e d on the study area to inclu d e as much v a r i a t i o n as p o s s i b l e w i t h respect to e l e v a t i o n , landforms, types and ages of logged over stands, and n a t u r a l h a b i t a t - t y p e s . The f l i g h t l i n e s used f o r the Grove study area are shown as an example i n Figure 3.3. 3.3 Results 3.3.1 Habitat Use Synoptic survey covered 12 6 ha on a t o t a l of e i g h t areas during May and June of 1972 and 1973. S i x areas were sampled i n 197 3 wit h 56 t r a n s e c t s , and f i v e areas were sampled i n 1973 w i t h 67 t r a n s e c t s . In both years, Eagle, Grove and Salmon were surveyed w i t h some s i t e s checked i n both w i n t e r s . The remaining f i v e areas were d i f f e r e n t . Data summaries are l i s t e d i n Appendix Tables D-4 and D-5. Results of the synoptic survey are summarized b r i e f l y below (Table 3.2): 1. Eagle: Winter use was gr e a t e s t i n the coniferous f o r e s t cutover, w i t h 108 p e l l e t groups/ha (pg/ha) recorded. This was f i v e times the l e v e l of use determined f o r the b i r c h f o r e s t . Two types w i t h i n the Eagle burn were used d i f f e r e n t i a l l y . The open shrubby v e g e t a t i o n had almost three times the use of the b i r c h f o r e s t . Although both types o r i g i n a t e d from the same f i r e , the b i r c h f o r e s t had l e s s browse a v a i l a b l e i n wint e r . The cut and leave s t r i p cutover received about the same use as the b i r c h type. 84a Figure 3.3 A map showing f l i g h t l i n e s used f o r the a e r i a l t r a n s e c t surveys on the Grove, study area. 85 86 2. Salmon: The most heterogeneous h a b i t a t s of aspen, mixed f o r e s t , and p a r t i a l cutover, received heaviest use. The coniferous f o r e s t was used at an intermediate l e v e l , w hile the c l e a r e d aspen type was used the l e a s t . Compared w i t h Eagle, d i f f e r e n c e s i n use of the various h a b i t a t s was not great. 3. Torpy: This s i t e was i n a heavy snow b e l t . The f o r e s t received e s s e n t i a l l y a l l the winter use. 4. Whites: The two c l e a r c u t s sampled experienced s i m i l a r l e v e l s of winter use although they were logged seven years apart. 5. Grove: The importance of burn h a b i t a t s was emphasized on t h i s winter range. Use of the 12 year o l d burn was double that of the coniferous f o r e s t a t 91 pg/ha. The mature f o r e s t received heavier use than the immature f o r e s t , presumably due to i t s b e t t e r forage supply and p o s s i b l y greater a b i l i t y to i n t e r c e p t snow. 6. McGregor: The p o s s i b l e e f f e c t of s l a s h burning i s i l l u s t r a t e d at t h i s study area. The unburned cutover received heavier use than the burned one. Apparently, the coniferous f o r e s t had l i t t l e or no winter use. This may have occurred because the sampling s i t e had l i t t l e understory vegetation and because i t was s i t u a t e d adjacent to the burned cutover. 7. McKenzie: A t o t a l of seven h a b i t a t - t y p e s were sampled, a l l w i t h i n c l o s e p r o x i m i t y of each other. As at the Eagle and Salmon study areas, the heterogeneous p a r t i a l cutover was very h e a v i l y used w i t h 323 pg/ha. This was the highest d e n s i t y recorded during the e n t i r e study. The i n c r e a s i n g use of o l d e r cutovers was a l s o shown at t h i s area w i t h group d e n s i t i e s i n c r e a s i n g from 0, 16 and 49 f o r c l e a r c u t s aged 1, 3 and 5 years. S i m i l a r to the Grove area, immature f o r e s t s were used l e s s than mature f o r e s t s . Mean l e v e l s of use v a r i e d between the study areas, although sample s i z e s were small f o r some areas (Table 3.2). Table 3.2 R e l a t i v e Winter Use of A v a i l a b l e Habitats on Selected Study Areas, Based on P e l l e t Group Surveys Study Area Winter use Study Area Winter use Habitat-type ( p e l l e t groups/ha) Habitat-type ( p e l l e t groups/ha) EAGLE: b i r c h f o r e s t ( 2 ) * 22(0. 2) * * c oniferous f o r e s t (4) 108(0. 3) p a r t i a l cutover (4) 73(0. 4) cut and leave cutover (2) 22(0. 2) burn - 35 yr (5) 57(0. 4) Mean (17) 63(1. 5) SALMON: mixed f o r e s t (3) 42(0. 3) aspen f o r e s t (3) 43(0. 3) cleared aspen f o r e s t (3) 23(0. 3) p a r t i a l cutover (11) 41(1. 3) coniferous f o r e s t (14) 35(3. 3) Mean (34) 36(5.6) GROVE: burn - 12 yr (13) imm. c o n i f e r , f o r e s t c oniferous f o r e s t (7) (4) 91(2.2) 22(0.4) 39(0.9) Mean (24) 69(3.5) MCGREGOR: c l e a r c u t , burn - 3 yr (1) 0(0.1) c l e a r c u t - 3 yr (1) 14(0.1) cut and leave cutover (1) 22(0.2) coniferous f o r e s t (2) 0(0.05) Mean (5) 12(0.5) Table 3.2, Continued Study Area Winter use Study Area Winter use Habitat-type ( p e l l e t groups/ha) Habitat-type ( p e l l e t groups/ha) TORPY: McKENZIE: c l e a r c u t - 1 yr ( 2 ) * * * 0(0.1) c l e a r c u t , burn - 1 y r (3) 0(0.3) conifer o u s f o r e s t (3) 36(0.1) c l e a r c u t , burn - 3 yr (2) 16(0.2) c l e a r c u t , burn - 5 y r (2) 49(0.2) Mean (5) 22(0.2) p a r t i a l cutover - 5 yr (2) 65(0.2) p a r t i a l cutover - 10 y r (3) 323(0.2) imm. c o n i f e r , f o r e s t (1) 11(0.1) WHITES: coniferous f o r e s t - 200 yr (3) 29(0.3) c l e a r c u t - 1 y r (1) 43(0.1) Mean (16) 67(1.4) c l e a r c u t - 8 y r (2) 34(0.2) Mean (3) 34(0.3) *No. of t r a n s e c t s . **No. ha i n sample. ***Unburned unless noted as burned. CO 00 89 Three l e v e l s were d i s t i n g u i s h a b l e : high at Eagle, Grove and McKenzie; moderate at Salmon and Whites; and low at McGregor and Torpy. Winter use a l s o v a r i e d between h a b i t a t - t y p e s , based on combined information from a l l areas (Table 3.3). Burns were most h e a v i l y used w i t h 8 5 pg/ha. P a r t i a l cutovers were a l s o h e a v i l y used at 69 pg/ha. Deciduous, mixed and coniferous f o r e s t s r e c e ived intermediate u t i l i z a t i o n , averaging 39/ha. Least used were c l e a r c u t s l e s s than 10 years o l d and immature f o r e s t s , where approximately 20 pg/ha were recorded. R e l a t i v e l e v e l s of h a b i t a t use ranged widely, w i t h the highest recorded use 4.5 times the lowest. Table 3.3 R e l a t i v e Winter Use of Major Habitat-types i n the Sub-boreal F o r e s t , Based on P e l l e t Group Surveys Habitat Winter use by moderate s n o w f a l l c l a s s * heavy O v e r a l l mean use Burn 91(2.2) 57(0.4) 85 P a r t i a l cutover 76(1.7) 51(0.6) 69 Mixed f o r e s t 42(0.3) ns 42 Mature coniferous f o r e s t 35(4.5) 68(0.7) 39 Deciduous f o r e s t 43(0.3) 22(0.2) 35 Clearcuts 26(1.0) 0 21 Immature coniferous f o r e s t 19(0.5)** ns*** 19 Means 52(10.5) 48(2.1) 51 *Use expressed as pg/ha. **Area sampled i n ha. ***Not sampled. Snowfall c l a s s modified s t r o n g l y the l e v e l s of use (Table 3.3). Thus i n heavy s n o w f a l l areas such as McGregor and Torpy, the p a r t i a l cutovers and mature coniferous f o r e s t s were the most h e a v i l y used of the sampled h a b i t a t s . Burns and p a r t i a l cutovers r e c e i v e d l e s s use, w h i l e deciduous f o r e s t s were used at an even lower l e v e l . C l e a r c u t s appeared to be avoided during w i n t e r . Compared w i t h i n areas of moderate s n o w f a l l , burns were c l e a r l y the most h e a v i l y u t i l i z e d . P a r t i a l cutovers ranked next, followed by deciduous and mixed f o r e s t s . Mature coniferous f o r e s t s ranked f i f t h r a t h e r than f i r s t as they d i d i n the heavy s n o w f a l l c l a s s . disturbance. This was i n d i c a t e d by comparing the r a t i o of p e l l e t group d e n s i t i e s i n a s e r a i stage to d e n s i t i e s i n an adjacent mature f o r e s t . Using the r a t i o s enables comparisons to be made between the study areas. These r a t i o s are as f o l l o w s (derived from Table 3.2): S e r a i age Ratio of s e r a i (yr) Habitat stage: f o r e s t Study areas 1-3 c l e a r c u t 0.3 Torpy, McGregor, The l e v e l of use changed w i t h time since McKenzie 5 c l e a r c u t p a r t i a l cut-over 1.7 2.2 McKenzie McKenzie 10-11 burn p a r t i a l cut-over 2.3 Grove McKenzie 11 35 burn b i r c h f o r e s t 0.5 0.2 Eagle Eagle 91 S e r a i age (yr) Habitat Ratio of s e r a i stage: f o r e s t Study areas 50 mixed wood, aspen lodgepole pine lodgepole pine 1.2 0.6 Salmon Grove 90 0.4 McKenzie Use of s e r a i stages v a r i e d from 0.20 to 11 times t h a t of mature f o r e s t s . Recent c l e a r c u t s and immature lodgepole pine and b i r c h f o r e s t s were used comparatively the l e a s t , w i t h a l l r a t i o s l e s s than or equal to 0.4:1. Older c l e a r c u t s , burns and e s p e c i a l l y p a r t i a l cutovers received heavier use than mature f o r e s t s except at Eagle. However, the v a r i a t i o n between study areas i n d i c a t e s t h a t the f o r e - going g e n e r a l i z a t i o n must be a p p l i e d c a r e f u l l y to s p e c i f i c study areas - other f a c t o r s such as p r o x i m i t y to roads a l s o i n f l u e n c e r e l a t i v e use of s e r a i stages. age and type of s e r a i stage, type of adjacent f o r e s t and d i s t a n c e from the ecotone (Table 3.4, Figure 3.4). Several case s t u d i e s i n d i c a t e d t h i s : 1. Eagle: On t h i s heavy s n o w f a l l study area, winter use of the open burn d e c l i n e d s t e a d i l y from the ecotone out i n t o the burn u n t i l approximately 300 m (Figure 3.4) from the f o r e s t edge. Beyond t h i s , use approached the average recorded f o r the e n t i r e burn. In the f o r e s t , use again d e c l i n e d but only u n t i l about 150 m from the edge a f t e r which use increased s t e a d i l y . This p a t t e r n r e s u l t e d p r i m a r i l y from the very high p e l l e t group counts made on one t r a n s e c t . ( E l i n 1973, where 32 p e l l e t groups were recorded i n a 305 m "&