"Land and Food Systems, Faculty of"@en . "DSpace"@en . "UBCV"@en . "Eastman, Donald Sidney"@en . "2010-03-05T21:12:35Z"@en . "1977"@en . "Doctor of Philosophy - PhD"@en . "University of British Columbia"@en . "A study of winter habitat selection and use by moose was conducted in a 11,300 km\u00B2 area of north-central British Columbia from May 1971 to August 1973. The study area was located within the forested sub-boreal spruce biogeoclimatic zone, a zone that is receiving increased development, especially by forestry. Habitat selection and use was examined mainly be pellet group surveys and aerial transects. Wintering moose used partial cutovers and burns more than coniferous forests; deciduous forests and recent clearcuts were used least. Limited data suggested a similar pattern in summer. Winter use typically increased from near zero after a recent disturbance such as clearcutting, to a peak sometime between 10 and 25 years later, then declined to low levels during 25 and 90 years, and then apparently stabilized in the mature forest stage at slightly higher levels. On one intensively surveyed area, moose selected partial cutovers and creek bottoms even though these habitats comprised less than 6 percent of the area. Moose began concentrating on winter ranges at least by mid-November, reached a peak in November-January, and declined steadily thereafter. Food habits and diet were examined by rumen analysis, trailing and post-winter browse surveys. Moose had catholic diets but ate primarily deciduous browse for most of the year. Subalpine fir becomes important in late winter. Diet varied according to season and habitat. Preferred species typically were least common. Tagged twig transects revealed that moose frequently browsed plants more than once but rarely re-browsed a twig. The time of browsing varied by species and by habitat with most use recorded in January and in April. Levels of utilization were all less than 100 percent of the previous year's production. Utilization (weight-basis) ranged from 33 percent on red-osier dogwood to 3 percent on subalpine fir; and from trace amounts in an upland burn habitat to more than 40 percent in deciduous forest, partial cutover and river bottom habitats. Bedding .habits were examined in an attempt to define cover for moose. Moose choose upper slopes that faced south particularly when snow depths became restrictive (> 80 cm). Moose tended to select larger than average trees and to bed on the southerly sides of them. Selection of bed sites varied with snow depth. As snow became deeper, moose bedded closer to larger trees in the denser canopied parts of forest stands. Moose showed greater selection for protected sites as winter conditions became more severe. Secondary seral succession was examined with respect to several attributes for mesic environments on the two commonest substrates, glacial till and lacustrine deposits. Floristics of seral stages from 1-200 years revealed that on lacustrine soils, vegetation was more, diverse and the deciduous phase was prolonged. Species diversity declined around year 25 on till but not on lacustrine. Several major changes occurred in the tree layer: first, a deciduous tree layer developed especially on lacustrine soils; second, after 25 years on till (45 years or more on lacustrine), lodgepole pine became most abundant; third, pine was gradually replaced by white spruce after 150-200 years or more; fourth, subalpine fir would probably become the dominant tree species in the absence of fire. Understory phytomass, though contributing little to the mature forest mass, increased dramatically to peaks early in succession and then remained low. Approximate net primary production of the understory on till was greatest at age 11 with 133 g/m\u00B2/yr produced and least at age 39 with 18 g/m\u00B2/yr produced. Understory production in the mature forest was and estimated 27 g/m\u00B2/yr. The shrubs contributed 70 percent, 26 percent, 44 percent, and 26 percent of annual production at ages 1, 11, 39, and 195 years, post-disturbance. Crude protein and lignin values were determined for 10 species (eight shrubs, one conifer, one lichen) for an annual cycle. Crude protein averaged 7 percent and lignin, 9.8 percent. Crude protein increased abruptly from steady winter values to peaks of 10-15 percent in June-July and then returned to low levels by October. Leaf protein was higher than, and predictable from, stem levels. Crude protein varied by-species, sometimes by substrate and rarely by habitat-type, at least for the species analyzed. The lichen, lungwort, retained a high protein value of approximately 11 percent throughout the year. Lignin levels varied seasonally, though less dramatically than crude protein. Levels were affected by species, substrate and age of seral stage. Protein levels were similar to those reported in the literature. Factors influencing crude protein were difficult to disentangle due to confounding. Winter climate was studied with respect to differences in snow features between habitats. Moose moved into winter ranges before snow depths were limiting. This indicates snow acts to trigger migration. On winter ranges, moose also moved into forested habitats in mid-winter (January) when snow depths approached 80 cm. Snow depths and densities varied between habitats. Snow cover was more variable in partially logged cutovers than in the open or forested stands. The climate of forest, ecotone and adjacent open areas were documented. Compared to adjacent open areas, the forest had higher relative humidity, less wind, more moderate temperatures and approximately 50 percent of the snow depth. The transition zone from open to forest climates appeared to be relatively narrow, less than 50 m. The relationship between carrying capacity, habitat selection and home range are discussed with reference to moose and management of their habitat. Management recommendations and suggestions for future research are provided."@en . "https://circle.library.ubc.ca/rest/handle/2429/21531?expand=metadata"@en . "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 \u00E2\u0080\u00A2 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 . . \u00E2\u0080\u00A2 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 \u00E2\u0080\u00A2 . \u00E2\u0080\u00A2\u00E2\u0080\u00A2 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 \u00E2\u0080\u00A2 FOOD [\"AVAILABLE .GRAZING QUANTITY GRAZING/ BROWSING MODEL FORESMANAGEMENT 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 \u00C2\u00B1 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\u00E2\u0080\u0094I t\u00E2\u0080\u0094I I STUDY AREAS lp 0^ B PRIMARY 1 Eagle 2 Grove 3 Salmon 4 Bowron 5 Found 6 Pyfe 7 Limestone 8 McGregor Major highway KILOMETRES 5\u00C2\u00BB 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\u00C2\u00B0C. 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\u00C2\u00B0C. Annual mean d a i l y minimums and maximums are 2.5\u00C2\u00B0C and 9\u00C2\u00B0C, r e s p e c t i v e l y , w i t h extremes of -50\u00C2\u00B0C recorded i n January and 34.4\u00C2\u00B0C 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\u00C2\u00B0C, while P r i n c e George, 110 km north, has 3.3\u00C2\u00B0C; Vanderhoof and Fo r t St. James (50 km n o r t h ) , have temperature means of 2.7\u00C2\u00B0C and 2.3\u00C2\u00B0C 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- \u00E2\u0080\u00A2-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 \u00E2\u0080\u0094 o a 1932 and oa 1937\u00E2\u0080\u0094.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 \u00C2\u00AB. 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 \u00E2\u0080\u0094 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 \u00E2\u0080\u0094 i ^ , . .. \u00C2\u00BB > Major highway EZD Conifer forest = Mature (>80yr.) Secondary road Sk, \u00E2\u0080\u009E , ' \u00E2\u0084\u00A2 a t u r f ( 2 1 - 8 \u00C2\u00B0 y f J \u00E2\u0080\u0094 S t u d y area boundary KSJ Mixed forest (>21yr.) \u00E2\u0080\u0094 \u00E2\u0080\u0094 \u00E2\u0080\u0094 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 \"\"\"^ ^ t r a n s e c t (Appendix Table A-5) ) . The l e v e l of use at ecotones v a r i e d according to the Table 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 \u00E2\u0080\u009E. . Eagle Grove Salmon McKenzie Torpy/McGregor Distance from \u00E2\u0080\u0094 -ecotone (m) burn f o r e s t burn f o r e s t c l e a r c u t * aspen c l e a r c u t f o r e s t c l e a r c u t f o r e s t 0 - 30 161 161 90 215 144 0 0 0 0 0 31 - 61 108 108 197 269 0 0 0 0 0 65 62 - 91 54 0 144 0 0 36 0 0 0 0 92 - 122 54 161 215 27 36 36 0 54 0 0 123 - 152 54 161 72 108 36 0 0 0 18 0 153 - 183 0 538 90 0 0 0 0 0 184 - 213 0 431 144 0 54 0 0 54 214 - 244 0 108 43 0 0 108 0 0 245 - 274 108 1292 72 0 0 0 0 0 275 - 305 108 0 36 0 108 0 0 108 305 43 22 76 27 ns 79 ;e of stage 37 12 5 2 2-4 *In an aspen stand cleared f o r pasture development. 92a Figure 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 . < I CO DL o DC O LU LU Q_ 200 100 100-1 200 300 200 100-1 EAGLE STUDY AREA a OVERALL MEAN USE b MEAN USE BEYOND 310 M. e ECOTONE 93 ] a \u00E2\u0080\u0094 e 100 200-1 GROVE STUDY AREA -a -b -e -b -a 2 0 0 1 SALMON STUDY AREA 100 H 100 E 3 H -STRIP WIDTH (M) \u00E2\u0080\u00A2n- -a -e -a -b 305 305 94 2. Grove: On t h i s moderate s n o w f a l l area, use i n c r e a s e d up to approximately 230 m from the f o r e s t edge and then d e c l i n e d s l o w l y but s t e a d i l y f o r up to a t l e a s t 305 m i n t o the burn. E x t r a p o l a t i n g beyond t h i s d i s t a n c e suggested t h a t use would reach average l e v e l s recorded f o r the e n t i r e burn at about 500 m from the edge. In the adjacent f o r e s t , use d e c l i n e d over the 305 m sampled but s t i l l remained a t g r e a t e r than the average l e v e l f o r the f o r e s t . Again, e x t r a p o l a t i n g the t r e n d i n d i c a t e d t h a t average l e v e l s would be reached a t 500 - 550 m i n t o the f o r e s t (Figure 3.4) 3. Salmon: In the c l e a r e d aspen f o r e s t , use d e c l i n e d s h a r p l y away from the ecotone to reach the o v e r a l l mean use by approximately 18 0 m. In the u n d i s t u r b e d aspen f o r e s t , use a t the ecotone was v i r t u a l l y n i l . L e v e l of use g r a d u a l l y i n c r e a s e d but had not reached mean use f o r t h i s h a b i t a t even 3 05 m from the ecotone. 4. McKenzie, Torpy, and McGregor: The r e c e n t l y c l e a r c u t areas r e c e i v e d l i t t l e or no use: most use was recorded i n the f o r e s t . No attempts were made to r e l a t e d i s t a n c e from the ecotone and l e v e l of use due to the low usage of open h a b i t a t s . These r e s u l t s on use of ecotones between f o r e s t e d and u n f o r e s t e d h a b i t a t s can be summarized as f o l l o w s : when l i t t l e or no forage i s a v a i l a b l e i n open h a b i t a t s , use of ecotones i s v i r t u a l l y n i l ; when forage i s s u f f i c i e n t i n the open (see S e c t i o n 7) ecotones r e c e i v e h e a v i e r than average use. The d e c l i n e i n use away from ecotones i s g r e a t e s t i n aspen f o r e s t s , i n t e r m e d i a t e i n heavy s n o w f a l l areas and l e a s t i n moderate s n o w f a l l areas. D i s t a n c e from ecotones at which use approaches the average f o r the open h a b i t a t s were 180, 400, and 550 m, r e s p e c t i v e l y , f o r these three types. Levels of use f o r roads and s k i d t r a i l s were d i f f e r e n t from the l e v e l s i n h a b i t a t s i n which these access routes were l o c a t e d (Table 3.5). This d i f f e r e n t i a l use occurred at l e a s t i n c l e a r c u t s and p a r t i a l cutovers, the only types sampled. My impression was that s i m i l a r d i f f e r e n t i a l s occurred i n burn and f o r e s t types, too, although f u r t h e r v e r i f i c a t i o n i s needed. Table 3.5 Winter U t i l i z a t i o n of Roads and H a b i t a t s i n Which they were Located, Based on Accumulated P e l l e t Groups i n 1973 at McKenzie Accumulated p e l l e t groups/ha S i t e s used access h a b i t a t (type) access h a b i t a t Type of access road, s k i d t r a i l 87 road, s k i d t r a i l 221 road 832 s k i d t r a i l 475 164 ( c l e a r c u t ) M17-18 Ml-2 224 ( p a r t i a l ) M19-21 M3-4 1552 ( p a r t i a l ) M22-24 M5-6, M14 262 ( p a r t i a l ) M25-26 M7-8 P e l l e t group counts on roads and s k i d t r a i l s could be employed to estimate use f o r the h a b i t a t i n which they occur. This o f f e r s a very u s e f u l f i e l d technique to assess u t i l i z a t i o n of h a b i t a t s since access roads can be sampled much more e a s i l y than logging s l a s h . The r e l e v a n t p r e d i c t i o n equation was: y = 1.8333 x -0.019, where x = no. of accumulated p e l l e t groups/sq m on an access route, y = estimated accumulated p e l l e t group d e n s i t y / s q m i n the 2 h a b i t a t , n = 4, R = 0.81, s\u00E2\u0080\u009E \u00E2\u0080\u009E = 0.0361, and c a l c u l a t e d F r a t i o = 8.31. Tabulated F r a t i o f o r 1 and 7 df = 5.59 f o r P = 0.05, and 12.25 f o r P = 0.01. A d d i t i o n a l sampling i s needed before t h i s approach could be widely adopted. Although u t i l i z a t i o n surveys were d i r e c t e d p r i m a r i l y at wi n t e r , l i m i t e d data were a l s o c o l l e c t e d on h a b i t a t use i n summer i n 1973 surveys. In the summer, moose pr e f e r r e d p a r t i a l cutovers to f o r e s t s and c l e a r c u t s (Table 3.6). Use of p a r t i a l cutovers appeared to be e s p e c i a l l y heavy at the Eagle study area, where the highest d e n s i t y of summer feces were recorded (131/ha). I t should be remembered th a t these comparisons were made between t e r r e s t r i a l h a b i t a t s : aquatic h a b i t a t s were not sampled. Based on other s t u d i e s of summer h a b i t a t use, aquatic h a b i t a t s would be used at l e v e l s equal to or greater than t e r r e s t r i a l ones. Table 3.6 R e l a t i v e Summer Use of Habitat-types Based on Accumulated Summer Feces Recorded i n the 1973 Synoptic Survey R e l a t i v e summer use (accumulated feces/ha)* Habitat Eagle Grove McKenzie Salmon A l l areas c o n i f e r f o r e s t -(1) 33 (2) 8 (4) 11 (12) 8 p a r t i a l cutover 131(1) ns 26 (5) 25 (8) 25 c l e a r c u t ns** ns 6 (8) ns 9 burn -(1) 32 (4) ns ? ns 22 *Study s t i e s were f o r Eagle: E l - 4 , Grove: Gl-6, McKenzie: M1-M16, and Salmon: S1-S26 except S6. **Not sampled. 3.3.2 Habitat S e l e c t i o n S e l e c t i o n was assessed through p e l l e t group sampling f o r the timber s a l e l i c e n c e area at the Salmon winter range. A t o t a l of 2,522 p l o t s were d i s t r i b u t e d over the 4,288 ha, f o r a t o t a l sample of 2.3 ha or 0.05 percent of the area. In general, few p e l l e t groups were counted (Table 3.7). 2 Approximately 94 percent of the 9.2 m p l o t s had no p e l l e t groups, w h i l e 5 percent had only one group. In the remaining 1 percent of the p l o t s , 27 had two groups, ten had three groups, two had fo u r , and one each had f i v e and ten groups. In t o t a l , 206 p e l l e t groups were encountered f o r an average of 0.08 groups per p l o t . H a b itats v a r i e d i n t h e i r area (Table 3.8). As t h i s area was i n t e n s i v e l y sampled, the types defined f o r the synoptic surveys were expanded as f o l l o w s (the percentages denote the pr o p o r t i o n of the study area occupied by that type) : 1) mature c o n i f e r f o r e s t (77 percent) 2) immature c o n i f e r f o r e s t (7 percent) 3) brush swamp (6 percent) 4) open swamp (2 percent) 5) p a r t i a l cutover (5 percent) 6) c l e a r c u t (2 percent) 7) creek bottom (1 percent) The major a d d i t i o n s were aquatic h a b i t a t s . Coniferous f o r e s t s c l e a r l y dominated the veg e t a t i v e cover w i t h 8 4 98 Table 3.7 D i s t r u 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 In t e n s i v e Salmon Area H a b i t a t s 0 1 2 3 4 5 10 No. of p l o t s per type mature conife r o u s f o r e s t 93 5 1 t * t t t 1857 (74%) immature conife r o u s f o r e s t 99 1 99 ( 4%) open swamp 99 1 99 ( 4%) brush swamp 92 8 99 ( 4%) creek bottom 92 3 5 39 ( 2%) p a r t i a l cutover 88 9 2 2 135 ( 5%) c l e a r c u t 99 1 192 ( 8%) Tot a l s 2361 120 27 10 2 1 1 2522 * t = l e s s than 0.5%. Table 3.8 S e l e c t i o n of Habitats-Types by Moose i n Winter as Indicated by Accumulated P e l l e t Groups on the Int e n s i v e Salmon Area H a b i t a t s No. of p e l l e t groups/type (%) Mean no. of groups/plot name area (%) mature conife r o u s f o r e s t 77 81 0.09 immature coniferous f o r e s t 7 0.5 0.01 open swamp 2 0.5 0.01 brush swamp 6 3.9 0.08 creek bottom 1 2.4 0.13 p a r t i a l cutover 5 10.6 0.16 c l e a r c u t 2 0.9 0.01 Tot a l s (7 types) 7288 ha 206 groups 0.11 percent of the study area i n these types. Brush swamps and p a r t i a l cutovers were the next l a r g e s t types, but each only covered approximately 5 percent. The remaining three h a b i t a t - t y p e s comprised j o i n t l y 5 percent of the timber s a l e area. Moose d i d not use h a b i t a t - t y p e s i n p r o p o r t i o n to t h e i r area (Table 3.8). The n u l l hypothesis of r e l a t i v e use being p r o p o r t i o n a l to areas was r e j e c t e d at P = 0.002 2 (x = 20.51, 6 d f ) . Data that most c l e a r l y r e f l e c t e d s e l e c t i o n of h a b i t a t - t y p e s were average number of groups per p l o t i n each type. Thus, p a r t i a l cutovers and creek bottom types were most p r e f e r r e d w i t h 0.16 and 0.13 groups per p l o t . Less p r e f e r r e d were the mature c o n i f e r f o r e s t and brush swamp types. The remaining h a b i t a t s were l i t t l e used w i t h approximately 0.01 groups per p l o t i n immature c o n i f e r f o r e s t , open swamp and c l e a r c u t types. 3.3.3 The Timing of M i g r a t i o n and Occupancy Periods M i g r a t i o n and occupancy periods were defined p r i m a r i l y by ten s e r i e s of a e r i a l t r a n s e c t s flown between January 1972 and May 1973. The number of moose seen per minute of t r a n s e c t f l y i n g was used as an index of the buildup and d e c l i n e of moose on the three i n t e n s i v e study areas. The value of t h i s approach was l i m i t e d by two f a c t o r s . F i r s t , the success i n s i g h t i n g moose depends l a r g e l y upon adequate snow cover. Thus surveys made i n 100 summer and s p r i n g probably underestimated moose d e n s i t i e s . This e f f e c t was p a r t l y o f f s e t by the very open nature of much of study areas. Second, moose t y p i c a l l y s h i f t from open to f o r e s t e d h a b i t a t s i n mid-winter. Since moose are l e s s observable i n f o r e s t e d h a b i t a t s , t h i s s h i f t would p a r t l y confound a c t u a l migrations away from winter ranges. This e f f e c t was p a r t l y o f f s e t by using t r a c k s to record a c t i v i t y . These shortcomings notwithstanding, pronounced changes i n the moose index were recorded. These changes were broadly s i m i l a r f o r the three areas even though v e g e t a t i v e cover was d i f f e r e n t . In the 1972-73 w i n t e r , moose began concentrating on the winter ranges at l e a s t by mid-November (Figure 3.5). The p a t t e r n of concentration was not documented but probably moose a r r i v e d g r a d u a l l y on the winter ranges (Edwards and R i t c e y 1956, Goddard 1970, Stevens 1970, Houston 1968, Coady 1974). Hunting might have delayed the timing and r a t e of con c e n t r a t i o n . Although I have no data on t h i s p o i n t , general observations by w i l d l i f e b i o l o g i s t s s t r o n g l y suggest that t h i s occurs. Peak d e n s i t i e s occurred i n mid-winter, but i n d i f f e r e n t months f o r the three areas (Figure 3.5). The highest moose i n d i c e s were recorded i n November at Eagle, i n December at Salmon, and i n January at Grove. The i n d i c e s 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 , reaching t h e i r lowest values i n A p r i l or May. Thus moose occupied these ranges f o r 100a Figure 3.5 Number of moose seen/minute of f l y i n g on ' \u00E2\u0080\u00A2' .\" the Eagle, Grove and Salmon study areas during a e r i a l t r a n s e c t s i n the 1972-73 wint e r . 102 approximately s i x months i n 1972-73. 3.4 Dis c u s s i o n 3.4.1 The Importance of Habitat V a r i a b i l i t y Moose used heterogeneous h a b i t a t s more than uniform h a b i t a t s . In p a r t i c u l a r , p a r t i a l cutovers were p r e f e r r e d over a l l others.on most of the study areas sampled. The uniform types that were l e s s h e a v i l y used were mainly immature s i n g l e - s p e c i e s stands and recent c l e a r c u t s . These r e s u l t s were s i m i l a r to many other moose h a b i t a t s t u d i e s . In Minnesota, Peek et a l . (1976:58) st a t e d : \"Apparently, t h i s logging i n a d v e r t e n t l y created i d e a l moose h a b i t a t by removing lar g e acreages of jackpine pulp timber and c r e a t i n g extensive shrub communities, i n t e r s p e r s e d w i t h balsam f i r and aspen and white b i r c h stands.\" In Newfoundland, P i m l o t t (1953) considered t h a t \"good\" moose h a b i t a t was the commercial f o r e s t , e s p e c i a l l y the mixed stands of balsam f i r (Abies balsamea) and white b i r c h (Betula papyrifera) i n t e r s p e r s e d w i t h muskegs and a l p i n e areas. High winter d e n s i t i e s of moose were recorded by Bergerud et a l . (1968) i n logged-over Newfoundland f o r e s t s . These cutovers contained s c a t t e r e d stands of r e s i d u a l c o n i f e r s i n t e r s p e r s e d w i t h regenerating balsam f i r and white b i r c h . In Nova S c o t i a , T e l f e r (1967) found t h a t 15 - 2 0 year o l d pulpwood cutovers provided winter moose yards, and P r e s c o t t (1968) remarked that w i n t e r i n g moose 103 concentrated most f r e q u e n t l y i n p a r t i a l cutovers that were approximately 15 years o l d . In a general review of winter h a b i t a t s of .Quebec moose, Brassard et a l . (1974) concluded that the best h a b i t a t occurred i n the f o r e s t zones that were t r a n s i t i o n a l between c o n i f e r s and broadleaf f o r e s t s . \"Within these . . . moose s e l e c t those areas that have su f f e r e d p e r t u r b a t i o n as a consequence of loggi n g , f i r e or i n s e c t outbreak\" (Brassard et a l . 1974:79). Moose populations of moderate d e n s i t i e s are a l s o found i n the heterogeneous t r a n s i t i o n zone that extends from n o r t h - c e n t r a l A l b e r t a to north-eastern Minnesota (Berg and P h i l l i p s 1974). In western North America, the c r i t i c a l r i p a r i a n winter h a b i t a t of the Shiras moose i s qu i t e d i v e r s e , due to f l o o d i n g , e r o s i o n , changes i n channels, and i c e damage (Peek 1974a). In Alas k a , LeResche et a l . (1974) reviewed important h a b i t a t s used by moose i n winter; these were c h a r a c t e r i z e d by considerable v e g e t a t i v e d i v e r s i t y . The r a p i d increase i n Fennoscandian moose numbers was a t t r i b u t e d p r i m a r i l y to a change from the ol d e r s e l e c t i v e methods to c l e a r c u t t i n g (Lykke and Cowan 1968, Markgren 1974). These c l e a r c u t s were s m a l l , t y p i c a l l y ranging only from 2 ha to 5 ha and so s t i l l represented d i v e r s e h a b i t a t s . Many researchers have demonstrated the value of h a b i t a t v a r i a b i l i t y as an important h a b i t a t component used during winter. Leopold (1933) expressed t h i s concept through h i s 104 Law of I n t e r s p e r s i o n . Given that v a r i a b i l i t y i s a s i g n i f i c a n t h a b i t a t requirement, s e v e r a l questions s t i l l remain unanswered: 1. Does use vary over time, and i f so, how? 2. How much v a r i a b i l i t y i s needed? 3. How does v a r i a b i l i t y vary over time? Results of my study i n d i c a t e that winter use i s minimal i n very young or immature f o r e s t stages. Low u t i l i z a t i o n i s due l a r g e l y to an inadequate forage supply (see Section 7.3.4). Use i s greatest i n s e r a i , stages ranging from 5 - 2 0 years. However, v a r i a t i o n i n t h i s time p e r i o d i s a t t r i b u t e d to such f a c t o r s as s i t e p r o d u c t i v i t y and winter snow depths. The importance of the l a t t e r was shown i n Table 3.3. The time span of 5 ^ 2 0 years a f t e r removal of a mature f o r e s t corresponds w i t h f i n d i n g s of most other researchers, p a r t i c u l a r l y logged-over f o r e s t s . As noted p r e v i o u s l y , cutovers 15 - 20 years o l d provided winter h a b i t a t i n Nova S c o t i a ( T e l f e r 1967, P r e s c o t t 1968). The S i s k i w i t Lake burn on I s l e Royale received heaviest use i n the f i r s t 14 years a f t e r which, use d e c l i n e d ( K r e f t i n g 1974) . However, the Feldtmann burn showed i n c r e a s i n g use up to 34 years afterwards. K r e f t i n g (1974) a t t r i b u t e d t h i s to a b e t t e r i n t e r s p e r s i o n of cover i n the Feldtmann Burn. Obviously, the most u s e f u l p e r i o d of a sere v a r i e s , as shown by the above comparison between the S i s k i w i t and 105 Feldtmann burns on I s l e Royale. A s i m i l a r case was reported i n the P r i n c e George area (K. Sumanink, pers. comm.). The Grove and Tsus Burn both o r i g i n a t e d i n 1961. The former i s an important winter range and has regenerated mainly to w i l l o w and b i r c h , w h i l e the l a t t e r i s an unimportant winter range and regenerated mainly to lodgepole pine. The question of how much v a r i a b i l i t y i s enough v a r i e s according to f a c t o r s s p e c i f i c to the area, and to the nature of the s e r a i v e g e t a t i o n . However, the processes which i n i t i a t e s e r a i development must be s u f f i c i e n t to st i m u l a t e understory production. T e l f e r (1972) found that s t i m u l a t i o n of browse production r e q u i r e d r e d u c t i o n of the 2 stand b a s a l area to 17 m /ha although the r e d u c t i o n must depend upon i n i t i a l s t o c k i n g . The s e l e c t i o n system p r a c t i c e d i n Fennoscandia was probably not i n t e n s i v e enough to a f f e c t browse production since moose populations d i d not increase u n t i l small s i z e d c l e a r c u t t i n g became the dominant c u t t i n g p r a c t i c e (Markgren 1974). In Idaho, Hungerford (19 69) found that commercial f o r e s t t h i n n i n g s were i n s u f f i c i e n t to reduce stand b a s a l area by the 20 - 30 per-cent necessary to s t i m u l a t e understory production. S i m i l a r l y , both c l e a r c u t t i n g and w i l d f i r e s produces more forage i n the e a r l y stages than moose can u t i l i z e . On the McKenzie study area, use of f i v e year o l d c l e a r c u t s was l e s s than the p a r t i a l cutover. A l s o , 10 year o l d 106 p a r t i a l cutovers were more h e a v i l y used than 10 year o l d burns. Thus the type of disturbance required to produce a t t r a c t i v e winter range should be s u f f i c i e n t to s t i m u l a t e browse production but need not be as d r a s t i c as complete removal of the f o r e s t cover. I t appeared that p a r t i a l c u t t i n g as p r a c t i c e d i n n o r t h - c e n t r a l B r i t i s h Columbia approached the i d e a l . A l t e r n a t i v e l y , i t appears t h a t 2 -10 ha c l e a r c u t s achieved a s i m i l a r \"enhancement\" e f f e c t f o r Scandinavian moose. The best s i z e and shape f o r c l e a r c u t s i s d i f f i c u l t to define because even very l a r g e c l e a r c u t s or burns are used by some moose, provided forage i s not snow covered. Moose are very adaptable animals. A more t r a c t a b l e restatement would be: \"what i s the best s i z e and shape f o r c l e a r c u t s i f the management o b j e c t i v e i s to maintain current moose populations?\" A s p e c i f i e d time frame i s an inherent part of t h i s type of question. To maintain present moose d e n s i t i e s on e x i s t i n g ranges the balance between cover - and food-producing areas, and how distance from cover a f f e c t the l e v e l of u t i l i z a t i o n must be determined. The temporal aspect must al s o be addressed. These components r e l a t e p r i m a r i l y to areas where the f o r e s t cover has been completely removed by c l e a r c u t logging or by intense w i l d f i r e . The r e l a t i o n s h i p between edge and use has i n t r i g u e d f o r e s t e r s and w i l d l i f e managers at l e a s t since c l e a r c u t t i n g 107 became the dominant c u t t i n g method. Disputes regarding the best s i z e and shape of c l e a r c u t s became acute as logging companies attempted to reduce u n i t costs by c l e a r c u t t i n g l a r g e r and l a r g e r b l o c k s . Although uncommon now, cutovers of 200 - 30 0 ha can be found i n the Pr i n c e George area. However, the question of s i z e and shape of c l e a r c u t s i s unanswerable unless r e l a t e d to a management goa l . Since moose w i l l use open areas w e l l removed from coniferous cover, i t f o l l o w s that the i n f l u e n c e of s i z e of c l e a r c u t s on the presence or absence of moose i s e s s e n t i a l l y very l i m i t e d , except i n the e a r l y s u c c e s s i o n a l stages when snow covers forage ( S t e l f o x et a l . 1976). However, the s i z e and shape of c l e a r c u t s w i l l a f f e c t the l e v e l of use. I f the management o b j e c t i v e i s to increase moose, then smaller c l e a r c u t s are i n order. I f the o b j e c t i v e i s to maximize timber production and l e t moose numbers adjust to t h i s , then a d i f f e r e n t set of c r i t e r i a f o r s i z e and shape of c l e a r c u t s w i l l be needed. .Results i n t h i s s e c t i o n demonstrated that use of open areas, p r i m a r i l y burns, d e c l i n e d w i t h distance from cover. The r a t e of d e c l i n e appeared to be i n f l u e n c e d by nature of the cover, age of the s e r a i stage and sno w f a l l regime. Comparatively l i t t l e other research has been done on t h i s t o p i c . T e l f e r (1974) summarized a few reports (Pimlott 1953:577-78, Vozeh and Cumming 1960:2-4, Bergerud and Manuel 1968:733) by saying that c l e a r c u t areas of 130 ha 108 might be u t i l i z e d by moose. None of these reports addressed the f u n c t i o n a l r e l a t i o n s h i p between l e v e l of use and distance from cover. Neu et a l . (1974) found on the L i t t l e Sioux Burn i n northeastern Minnesota that w i n t e r i n g moose used the 0.4 km p e r i p h e r i e s of the burn and the adjacent f o r e s t s i g n i f i c a n t l y more than expected according to t h e i r r e s p e c t i v e a v a i l a b i l i t i e s . I r w i n (1975) recorded s i m i l a r r e s u l t s f o r the subsequent winter of 1971-72. Although snow depths were not given, the p r e f e r r e d 0.4 km periphery i s remarkably s i m i l a r to the s i t u a t i o n recorded f o r the Eagle study area (Figure 3.4). Recent work i n Ontario provides probably the most r e l e v a n t data to date (Hamilton et a l . 1976, Hamilton and Drysdale 1976). Studies were c a r r i e d out i n 1975 and 1976 on three cutovers, aged f i v e and s i x years o l d , and that ranged from 24 ha to 525 ha i n the area. In 1975, 95 per-cent of browsing occurred w i t h i n 80 m of boreal f o r e s t cover. However, i n 1976, s i g n i f i c a n t use was recorded up to 260 m from cover. These d i f f e r e n c e s i n browsing were independent of browse d i s t r i b u t i o n . Causes of annual v a r i a t i o n s were not determined, but such f a c t o r s as increased moose d e n s i t i e s , sampling v a r i a t i o n and age of cutover were suggested. To these reasons could be added v a r i a t i o n i n snow depth and\u00E2\u0080\u00A2hardness. The data showed v a r i a t i o n i n d e t a i l s , but s i m i l a r i t y i n the general r e l a t i o n s h i p of d e c l i n i n g use w i t h i n c r e a s i n g d i stance from cover. 4. FOOD HABITS 4.1 I n t r o d u c t i o n Information on food h a b i t s defines what p l a n t species moose consider as forage. I t a l s o helps to i n t e r p r e t h a b i t a t use and s e l e c t i o n , and to assess r e l a t i o n s h i p s with f o r e s t r y p r a c t i c e s . E s p e c i a l l y important are data that i d e n t i f y d i e t a r y preferences on l o c a l bases (Peek 1974b). In the b o r e a l f o r e s t s of western North America ( i n which my study area l i e s e n t i r e l y ) , food h a b i t s of moose are poorly documented. Peek's (1974b) comprehensive review of moose feeding s t u d i e s on t h i s continent contained no references f o r t h i s biome. Information from p e r i p h e r a l areas i s a v a i l a b l e but of l i m i t e d value (e.g., Hatter 1950, Ri t c e y 1967, R i t c e y and Verbeek 1969, S i l v e r 1976). Studies from other b o r e a l regions have l i t t l e a p p l i c a t i o n due to d i f f e r e n c e s i n v e g e t a t i o n , c l i m a t e , and i n the w i n t e r i n g h a b i t s and h a b i t a t s of moose. Therefore, I conducted a survey to f i l l t h i s gap i n i n f o r m a t i o n . Three main questions were asked: 1) how many species do moose eat?, 2) what seasonal trends e x i s t ? , 3) do d i e t s vary w i t h h a b i t a t - t y p e ? These questions were answered by using three methods, v i z . , rumen analyses, t r a i l i n g and post-winter browse t r a n s e c t s . These 109 110 complementary techniques overcome the biases and l i m i t a t i o n s of each as mentioned by Peek (1974b!: 211-213) . 4.2 Methods 4.2.1 Rumen A n a l y s i s Rumen samples were c o l l e c t e d w i t h i n the general study area from moose k i l l e d by hunters, by poachers and by accide n t s . Information on these samples i s presented i n Appendix E. Approximately f i v e samples per month were c o l l e c t e d : attempts were made to ob t a i n rumens from both sexes and a wide range of age c l a s s e s . One l i t e r samples of well-mixed d i g e s t a were c o l l e c t e d from each rumen, l a b e l l e d and frozen f o r l a t e r a n a l y s i s . Freezing has the advantage over other storage methods of prese r v i n g h e l p f u l c o l o r -based c h a r a c t e r i s t i c s of p l a n t s (Korschgen 1969). The method used was chosen a f t e r t r i a l analyses of 11 i n i t i a l samples (Eastman 1974b). Several approaches were t e s t e d to determine the most s u i t a b l e one, and a l s o to determine the fat e of rumen m a t e r i a l during processing. Standard procedures were followed on completely thawed samples. Oven-dried weights of each sample were determined from t o t a l wet weights ( a f t e r d r a i n i n g to remove excess f l u i d ) and moisture content derived from three subsamples. Samples were washed through standard s i e v e s , 6.35 mm, 4.00 mm, and 2.00 mm, u n t i l l i t t l e or no m a t e r i a l appeared i n the discharge. These s i z e s are commonly used i n analyses of I l l ungulate rumens. M a t e r i a l remaining on the top two sieves was t r a n s f e r r e d to a white enamelled t r a y and separated by hand. This m a t e r i a l was c a l l e d the coarse f r a c t i o n . Separated m a t e r i a l was measured f i r s t v o l u m e t r i c a l l y by water displacement and then weighed a f t e r being oven-dried f o r 4 8 hours at 5 0\u00C2\u00B0C. The m a t e r i a l remaining on the bottom s i e v e , the fine f r a c t i o n was analyzed by p o i n t frame sampling and by manual separation of subsamples. F i r s t , the f r a c t i o n was spread evenly over the bottom of an enamelled t r a y and the fragments sampled w i t h a point-frame. A slanted row of ten p i n s , spaced e q u i d i s t a n t on a wooden frame, was moved across the t r a y at 50 cm i n t e r v a l s . At each i n t e r v a l , the pins were lowered and h i t s recorded u n t i l a t o t a l of one hundred h i t s were obtained. Next, f i v e subsamples of the f i n e f r a c t i o n were separated manually under a b i n o c u l a r d i s s e c t -ing microscope. M a t e r i a l s separated from each subsample were oven-dried and weighed. The unsampled f i n e m a t e r i a l remaining on the t r a y was a l s o oven-dried and weighed. During washing, approximately 7 5 percent of each rumen sample was l o s t (Table 4.1). This represents 154 g of the one l i t e r samples whose mean weight was 193 g. Factors determining the p r o p o r t i o n l o s t were mesh opening, the foods eaten, and the thoroughness of s i e v i n g the samples. The d i g e s t a r e t a i n e d on the screens were almost evenly d i v i d e d between coarse (nine percent) and f i n e (twelve percent) f r a c t i o n s (Table 4.1). A l l of the coarse m a t e r i a l was separated while only f i v e percent of the f i n e f r a c t i o n , or 0.7 percent of the t o t a l rumen sample, was used i n t h i s way. However, a l l the f i n e m a t e r i a l was used f o r the p o i n t frame sampling. Table 4.1 Components of Rumen Digesta A f t e r Sample P r e p a r a t i o n Component amount (oven-dried wt i n g ) * Item coarse f r a c t i o n f i n e f r a c t i o n l o s t i n washing Type of d i g e s t a : i d e n t i f i e d taxa 5.6 \u00C2\u00B1 0.6 0.25 \u00C2\u00B1 0.004 d e c o r t i c a t e d twigs 9.6 \u00C2\u00B1 3.3 0.80 \u00C2\u00B1 0.020 other m a t e r i a l 1.3 \u00C2\u00B1 0.3 0.05 \u00C2\u00B1 0.000 T o t a l weight separated 16.5 \u00C2\u00B1 4.1 1.10 \u00C2\u00B1 0.020 Mean weight of component 16.5 22.4** 153.9 Pr o p o r t i o n of mean sample weight 9% 12% 80% *Mean \u00C2\u00B1 standard e r r o r of mean, where n = 11 except f o r l o s t - i n -washing m a t e r i a l where n = 8. **0nly subsamples of t h i s were separated. Not a l l of the separated p l a n t m a t e r i a l was i d e n t i f i a b l e as to taxon (Table 4.1). Pieces of bark, shoots, p e t i o l e s , e t c . , defy a n a l y s i s unless v a r i o u s micro-techniques are employed (e.g., Hercus 1960, Stewart 1967) -a s o p h i s t i c a t i o n probably unwarranted when the inherent l i m i t a t i o n s of rumen a n a l y s i s are considered. The l a r g e s t s i n g l e item, c l a s s i f i e d as woody twigs, comprised 58 percent 113 and 73 percent of the m a t e r i a l from coarse and f i n e f r a c t i o n s , r e s p e c t i v e l y . In t h i s and other s t u d i e s , the species composition of these twigs i s assumed to be s i m i l a r to that of the i d e n t i f i a b l e m a t e r i a l . To my knowledge, t h i s assumption has not been t e s t e d . Estimates f o r frequency of occurrence v a r i e d according to mesh s i z e and to procedure (Table 4.2). For separation of the coarse f r a c t i o n , only nine (eleven percent) of the p o s s i b l e 7 6 occurrences (sum of the number of times a l l taxons were recorded i n the eleven rumen samples) were missed. These misses c o n s i s t e d mostly of small pieces from uncommon taxons, such as s i n g l e needles of white spruce. S i m i l a r l y , the number of misses recorded i n the a n a l y s i s of the f i n e f r a c t i o n was twenty (26 percent) f o r separation and f o r p o i n t frame sampling 45 (59 percent). With the l a t t e r method, most taxons missed comprised l e s s than ten percent by weight i n a sample, i n d i c a t i n g that the p o i n t analyzer was unsuited f o r l e s s abundant items. Even i f r e s u l t s from separation of both f i n e and coarse f r a c t i o n s were combined, the p r o p o r t i o n of misses would s t i l l be approximately three percent. The make-up of the \"misses\" a l s o v a r i e d . In the coarse f r a c t i o n only one taxon, the l i c h e n Aleotovia spp., was missed; the frequency of occurrence of f i v e others was underestimated. In separation of f i n e m a t e r i a l , s i x taxons were overlooked and e i g h t were under-estimated. With the 114 Table 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 -in Moose Rumen Samples Times recorded No. of times taxa were missed f o r 11 rumen separation p o i n t frame samples coarse f i n e f i n e TREES AND SHRUBS: Abies lasiocarpa (subalpine f i r ) Alnus spp. (alder) Betula papyrifeva (paper b i r c h ) Cornus stolonifera ( r e d - o s i e r dogwood) Pioea glauoa (white spruce) Pinus contorta (lodgepole pine) Populus tremuloides (trembling aspen) P. balsamifeva (black cottonwood) Salix spp. (willows) Sorbus spp. (mountain ash) Vaccinia spp. ( v a c c i n i a ) Rosaceae Salicaceae FORBS: Cornus canadensis (bunchberry) GRAMINEAE: PTERIDOPHYTES: BRYOPHYTES: LICHENS: Alectovia spp. (old man's beard) Lob aria pulmonavia (lungwort) 10 2 4 4 6 1 1 1 11 1 4 3 6 1 4 6 1 1 9 1 1 3 2 1 1 1 1 2 4 4 6 1 5 3 3 5 1 4 5 1 1 1 T o t a l s (19 taxa) 76 8 (11%) 20 (26%) 45 (59%) 115 poi n t frame, eleven of the t o t a l nineteen taxons were not recorded, f i v e were underestimated and only three agreed w i t h the composite i n f o r m a t i o n . Estimates f o r amounts of taxons a l s o v a r i e d according to mesh s i z e and to procedure (Table 4.3). Using a l l g r a v i m e t r i c data as the best q u a n t i t a t i v e estimate of rumen components, the l e a s t d e v i a t i o n was obtained when based on separation of the coarse f r a c t i o n (0.2 percent ignored), and the g r e a t e s t d e v i a t i o n when based on p o i n t frame sampling of the f i n e f r a c t i o n (5.7 percent, s i g n ignored). The d i s c r e p a n c i e s of the p o i n t frame sampling are l a r g e r than those recorded by other observers (e.g. Chamrad and Box 1964, Robel and Watt 1970). This may be due to d i f f e r e n c e s i n screen s i z e s although Chamrad and Box (1964) d i d not mention the s i z e of screens they used. Samples used by these observers and myself were a l i k e , w i t h one hundred h i t s recorded i n a l l cases. Thorough mixing of the m a t e r i a l minimized non-randomness i n the d i s t r i b u t i o n of p l a n t fragments. The most l i k e l y reason f o r d i f f e r e n c e was v a r i a t i o n i n s i z e of fragments i n my samples. Perhaps a n a l y s i s of f i n e r m a t e r i a l would overcome these d i s c r e p a n c i e s . S c o t t e r (1966:241) expressed s u r p r i s e that many rumen analyses are based on volumetric procedures. However, i t i s more s u i t a b l e f o r f i e l d operations because i t i s 116 Table 4.3 The Effect of Analytical Method Plant Taxa Identified in Moose on Amounts of Rumen Samples Proportion (%) Deviation from total wt (%) Plant taxa of total wt* separated separation coarse fine point frame fine TREES AND SHRUBS: Abies lasiocarpa (subalpine f i r ) 33 1 20 49 Alnus spp. (alder) 3 t** -3 -3 Betula papyrifera (paper birch) 3 t -2 -3 Cornus stolonifera (red-osier dogwood) 5 t -1 -5 Picea glauoa (white spruce) 2 t 3 -2 Populus tremuloides (trembling aspen) 4 t -4 -4 P. Balsamifera (black cottonwood) t - t -;t Salix spp. (willows) 37 2 -16 -29 Sorbus spp. (mountain ash) t t - t -t Vaccinia spp. (vaccinia) 2 t 4 -2 Rosaceae t - t Salicaceae 2 t -2 -1 FORBS: Cornus canadensis (bunchberry) t - t -t GRAMINEAE: 1 t -1 PTERIDOPHYTES: 1 t 5 BRYOPHYTES: t - t -t LICHENS: Alectoria spp. (old man's beard) t t -t Lobaria pulmonarea (lungwort) 6 1 -4 Totals 63.75 g Mean deviation, sign ignored t ' 3 6 *Oven-dried weight basis. Includes a l l material that was separated from coarse and fine fraction. * * t = less than 1%. 11.7 f a s t e r under some circumstances and r e q u i r e s l e s s equipment than g r a v i m e t r i c a n a l y s i s . A l s o , the apparent disadvantage of i n c o m p a t a b i l i t y of volumetric and g r a v i m e t r i c data can be overcome since they are h i g h l y c o r r e l a t e d . (r = 0.99 , n = 101, s = 0.343). For the present data, dry weights of separated taxons were estimable from t h e i r volumes by the r e g r e s s i o n equation: y = 74.69x + 276.64, w i t h y = 2 weight i n g and ,x = volume i n ml (s = 0.343, r =0.98, y \u00E2\u0080\u00A2 ^ n = 101) . Based on the foregoing d e t a i l e d examination of the 11 i n i t i a l rumen samples, the most s a t i s f a c t o r y approach was to separate d i g e s t a remaining on a sieve w i t h a mesh opening of 4.00 mm. Analyzing f i n e r m a t e r i a l than t h i s , e i t h e r by separation or p o i n t frame sampling, r e s u l t e d i n missing some taxons and i n underestimating both frequency of occurrence and amounts of others. Although p o i n t sampling was the q u i c k e s t procedure, the r e s u l t i n g data were incomplete. 4.2.2 T r a i l i n g T r a i l i n g provided time- and h a b i t a t - s p e c i f i c i n f o r m a t i o n on foods eaten. T r a i l i n g was done i n conjunction with other f i e l d work. A l l evidence of browsing and grazing was recorded by species, while f o l l o w i n g f r e s h l y made t r a i l s . Each f r e s h stub was considered as one b i t e , and a minimum of one hundred b i t e s f o r each t r a i l was t a l l i e d whenever p o s s i b l e . Information 118 on the abundance of a v a i l a b l e food species was gained from vegetation data c o l l e c t e d f o r the p l a n t succession aspect of t h i s study (see Section 7). 4.2.3 The Browsed Stem Survey These surveys were done i n conjunction w i t h post-winter p e l l e t group counts. Thus they were a r e a - s p e c i f i c but d i d not r e v e a l at what time during the previous winter browsing occurred. They complemented the data from rumen samples. These l a t t e r data were t i m e - s p e c i f i c but not n e c e s s a r i l y a r e a - s p e c i f i c since a moose may have fed i n a d i f f e r e n t h a b i t a t from the one i n which i t was k i l l e d . The method used was to not the species present and whether or not they were browsed i n the previous winter. 2 These observations were made i n c i r c u l a r 4 m (mil-acre) p l o t s spaced at e i t h e r 20 m or 30 m i n t e r v a l s along the p e l l e t group t r a n s e c t s . At l e a s t 20 p l o t s were sampled f o r each h a b i t a t v i s i t e d . 4.3 Results 4.3.1 The Range of Species Taken Moose used at l e a s t 64 p l a n t species ranging from l i c h e n s to d i c o t s , based on a l l the data (Table 4.4). One-t h i r d of the species were shrubs, one-quarter were f o r b s , one-quarter were l i c h e n s and mosses, w i t h the remainder mostly c o n i f e r s and f e r n s . While t h i s adequately r e v e a l s 119 Table 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 Their North American Range No. of species by forage c l a s s (%) Forage c l a s s C e n t r a l B.C. Wyoming Minnesota Alaska Ontario Co n i f e r s 9 6 14 2 13 Shrubs 33 41 58 31 43 Forbs 23 42 14 39 20 Graminoids 2 9 11 11 9 Ferns 6 3 10 Clubmosses Live r w o r t s 2 2 H o r s e t a i l s 2 3 3 1 Mosses 13 2 Algae 3 Lichens 11 3 4 Mushrooms 3 1 T o t a l no. species 64 69 36 62 80 Reference This study Houston Peek Le Resche Peter (1968) (1971a) and Davis (1955) (1973) the range of forage c l a s s e s taken, i t undoubtedly under-estimates the a c t u a l number of species taken. Many aquatics and other species g e n e r a l l y recognized as moose foods were not found (e.g., i n R i t c e y and Verbeek 1969), and few rumen were c o l l e c t e d i n summer when food h a b i t s were l i k e l y the most v a r i e d . Despite these shortcomings, the r e s u l t s show that moose eat a v a r i e t y of p l a n t species. This compares favo r a b l y w i t h other s t u d i e s except that moose i n t h i s study 120 appeared to take more mosses and l i c h e n s (Table 4.4). Whether or not a l l species were d e l i b e r a t e l y taken or a c c i d e n t a l l y ingested w i t h others was not always determinable. Probably most of the mosses and l i c h e n s were eaten a c c i d e n t a l l y by f o r a g i n g moose since these p l a n t s grew on or very c l o s e to s t a p l e food p l a n t s . 4.3.2 The Seasonal Trends This study supported the general p a t t e r n of seasonal trends i n moose food h a b i t s as summarized by Peek (1974). I t a l s o supported Peek's observations that moose of c e n t r a l B r i t i s h Columbia have s i m i l a r food h a b i t s to those i n eastern North America. The sp r i n g d i e t c o n s i s t e d of 91 percent deciduous shrubs, mainly S i t k a a l d e r {Alnus orispa sinuata) , Douglas maple (Acer glabrum) , w i l l o w and paper b i r c h . The high p r o p o r t i o n of S i t k a a l d e r (31 percent) i s unusual f o r moose. I t s occurrence i n the spr i n g d i e t and that of other shade t o l e r a n t , m o i s t u r e - l o v i n g species such as D e v i l ' s club (Oplopanax horridus) and r e d - o s i e r dogwood {Cornus stolonifeva) , suggested that moose i n the study area fed more commonly i n mesic r a t h e r than i n x e r i c f o r e s t s . In summer, use of w i l l o w increased to 49 percent while use of S i t k a a l d e r dropped to 4 percent. During both s p r i n g and summer seasons, forbs and graminoids formed l e s s than 10 percent of the d i e t . This f i g u r e l i k e l y under-estimates the importance of these forage c l a s s e s since three 121 rumens c o l l e c t e d during these seasons contained <1 percent, 72 percent and 89 percent f o r b s ; 50 percent, 11 percent and 2 percent graminoids; and 5 percent, 11 percent and 0 per-cent h o r s e t a i l s (Equisetum spp.), r e s p e c t i v e l y (also c f . R i t c e y and V.erbeek 1969) . Rumen samples taken i n e a r l y f a l l contained s i g n i f i c a n t amounts of non-browse species (Table 4.5, Figure 4.1). A d d i t i o n a l l y , woody p l a n t s would tend to have more recognizable items i n the rumen than succulents since they degrade l e s s r e a d i l y . Deciduous shrubs continued to form the l a r g e s t forage c l a s s i n the f a l l d i e t (63 p e r c e n t ) , w i t h w i l l o w and r e d - o s i e r dogwood as the main species. Grasses rank as the second most important forage c l a s s (25 p e r c e n t ) , w i t h ferns a l s o important (10 percent). P o s s i b l y the increase i n f e r n s , e s p e c i a l l y the evergreen lady iAthyvium filix-femina) and grape-ferns (Botrychium multifidum), r e f l e c t s the d e c l i n i n g p a l a t a b i l i t y and a v a i l a b i l i t y of t e r r e s t r i a l and aquatic forbs due to f r o s t and lowered water l e v e l s . E a r l y and l a t e winter d i e t s were examined to see i f they changed when moose s h i f t e d from open h a b i t a t s i n e a r l y winter to coniferous f o r e s t s i n l a t e winter (Table 4.5). In e a r l y w i n t e r , moose ate p r i m a r i l y w i l l o w , paper b i r c h , and r e d - o s i e r dogwood. In l a t e w i n t e r , when moose used f o r e s t e d h a b i t a t s most h e a v i l y , sub-alpine f i r comprised 26 percent of the d i e t , w i t h paper b i r c h and w i l l o w making up almost comparable p r o p o r t i o n s . Thus, the r a t i o of c o n i f e r s to Table 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) Species eaten* May - June J u l y - Aug Sept - Oct Nov - Jan Feb - Apr t r a i l i n g t r a i l i n g rumen t r a i l i n g rumen t r a i l i n g rumen CONIFERS: Abies lasiocarpa (subalpine f i r ) Pinus eontovta (lodgepole pine) Pseudotsuga menziesii (Douglas f i r ) Thuja plicata (western red cedar) 17 t 1 t 23 t 26 t DECIDUOUS TREES & SHRUBS: Acer glabrium (Douglas maple) Alnus arispa ( S i t k a alder) Ametanehier alnifoHa (Saskatoon) Betula glandulosa (bog b i r c h ) B. papyrifera (paper b i r c h ) Cornus stolonifera ( f e d - o s i e r dogwood) Lonicera involuorata (black twinberry) Oplopanax. horridus ( d e v i l ' s club) Populus tremuloides (trembling aspen) P. balsamifera (black cottonwood) Rosa spp. (rose) Salix spp. (willow) Sorbus sitchensis ( S i t k a mountain ash) Spiraea spp. (spirea) Vaecinium spp. ( v a c c i n i a ) Viburnum edule (squashberry) 13 4 1 4 t 4 31 4 1 4 1 t 1 5 6 t 6 1 1 1 1 2 9 9 4 19 27 32 12 7 4 12 23 11 3 3 4 8 4 t t 1 2 1 6 1 3 5 8 5 3 1 t 1 3 2 1 4 1 t 2 t 10 49 28 27 29 21 36 2 1 t 3 1 2 t 2 t t 1 t t 1 2 6 4 5 1 Table 4.5, Continued May - June J u l y - Aug Sept Oct Nov - Jan Feb - Apr. Species eaten* FORBS: Aster spp. (aster) Disporwn oreganum (Oregon f a i r y b e l l s ) Epilobium angustifolivcm (fireweed) Streptopus amplexifolius ( t w i s t e d s t a l k ) t r a i l i n g t r a i l i n g rumen t r a i l i n g rumen t r a i l i n g rumen GRAMINOIDS: Graminae (grasses) Typha latifolia ( C a t - t a i l ) 25 1 FERNS: Athyrium filix-femina (lady fern) Botrychium multifidum ( l e a t h e r y grape-fern) Dryopteris austriaca (spiny wood-fern) 1 1 LICHENS: Ldbaria pulmonaria (lungwort) U n i d e n t i f i e d l i c h e n s T o t a l number species recorded Data- base 27 15,844 b i t e s 12,116 23 b i t e s 36 21 1 t 33 6 1 21 22 12(105 g ) * * 10,543 b i t e s 33(321 g) 4,447 17(143 g) b i t e s *Includes only species recorded w i t h more than 1%. **No. samples ( t o t a l oven-dried wt). 123a Figure 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. Derived from data i n Table 4.5. I 125 deciduous shrubs changed from 1:63 i n f a l l , to 1:18 i n e a r l y w i n t e r , and to 1:3 i n l a t e w inter. This s h i f t probably r e f l e c t e d the e f f e c t s of snow on the a v a i l a b i l i t y of forage species. Lungwort {Lobaria pulmonaria) was more abundant i n the l a t e winter d i e t than f o r any other season. No signs of moose pawing through snow were .seen, although t h i s apparently occurs i n the study area (K. Sumanik, pers. comm.), i n northeastern and south c e n t r a l B r i t i s h Columbia (R. S i l v e r and R. R i t c e y , pers. comm.), and i n Alaska (Le Resche and Davis 1973). 4.3.3 The E f f e c t of Habitat-Type on D i e t Diet v a r i e d according to h a b i t a t . This was shown by data from rumen analyses (Table 4.6), and from post-winter browse surveys (Table 4.7). For example, y e a r l y winter d i e t s d i f f e r e d between three adjacent h a b i t a t : logged, o l d burn and coniferous f o r e s t (Table 4.6). Although paper b i r c h and w i l l o w were major d i e t a r y items, the p r o p o r t i o n of subalpine f i r v a r i e d . I t comprised 43 percent of the d i e t i n the o l d burn, 10 percent i n the logged type and was i n s i g n i f i c a n t i n the f o r e s t . Trembling aspen and lady f e r n were w e l l represented i n the samples from the f o r e s t but were uncommon or absent i n moose sampled from the other two h a b i t a t s . Samples from the Grove study area i n the l a t e winter p e r i o d revealed d i e t a r y d i f f e r e n c e s between recent burn and f o r e s t h a b i t a t s . Moose i n the former type ate 126 Table 4.6 Comparisons of Food Habits of Moose between D i f f e r e n t H a b i tats i n E a r l y and Late Winter* P r o p o r t i o n of species i n d i e t (%)** Area 1: e a r l y winter Area 2: l a t e winter Species recorded logged o l d burn f o r e s t f o r e s t recent burn CONIFERS AND DECIDUOUS SHRUBS: Abies lasioaarpa (subalpine f i r ) 16 43 t 31 1 Acer glabrum (Douglas maple) 3 6 1 Alnus crispa (alder) 2 t 1 Be tula papyrifera (paper b i r c h ) 34 24 24 2 25 Cornus stolonifera ( r e d - o s i e r dogwood) 8 5 1 2 Populus tremuloides (trembling aspen) 5 1 22 7 Populus balsamifera (black cottonwood) 1 7 Salix spp. (willow) 30 25 21 16 72 Sorbus spp. (mountain ash) 2 Viburnum edule (squashberry) 5 OTHER SPECIES: Athyrium filix-femina (lady fern) 1 17 t Lobaria pulmonaria (Lungwort) t t 10 No. spp. recorded 15 11 16 13 7 No. rumen samples 10 4 4 7 4 * E a r l y winter i s September to December and l a t e winter i s January to A p r i l . Habitat-types are logged - cutovers from 1 to 20 years o l d , f o r e s t - coniferous f o r e s t s , recent burn - 13 years o l d , and o l d burn - 60 to 70 years o l d . **0nly species w i t h at l e a s t 1% i n d i e t . 127 Table A.7 Winter Food P r e f e r e n c e s of Noose i n N o r t h - C e n t r a l B r i t i s h Columbia, by Hab i t a t - T y p e Percent occurrence/percent browsed** coniferous clear p a r t i a l recent Species recorded* forest cut cutover burn CONIFERS: Abies lasiocarpa (subalpine f i r ) Picea glauca (white spruce) Pinus contorta (Lodgepole pine) Pseudotsuga menziesii (Douglas f i r ) DECIDUOUS TREES & SHRUBS: Acer gldbrum (Douglas maple) Alnus crispa (Sitka alder) Amelanahier alnifolia (Saskatoon) Betula papyrifera (paper birch) Chimaphila umbellata (prince's pine) Cornus stolonifera (red-osier dogwood) Lonicera involucrata (black twinberry) Oplopanaz horridus (devil's club) Populus tremuloid.es (trembling aspen) Ribes spp. (gooseberry, currant) Rosa spp. (rose) Rubus idaeus (raspberry) R. parviflorus (thimbleberry) Salix spp. (willow) Sambucus racemosa (elderberry) Sorbus spp. (mountain ash) Spiraea luaida (flat-top spirea) S. douglasii (hardhack) Symphoricarpus alba (waxberry) Vaceiniwn spp. (vaccinia) Viburnum eduls (squashberry) No. of plots Prop, by type No. spp. present Prop, of spp. browsed 35/3 12/8 51/8 21/37 11 14/8 19 32/33 t 4/57 7 3/50 2 14/17 1 4/33 2 6/22 12/50 17 4 26/2 1/10C 9 10/35 41 4/13 17/15 13/37 24/44 10 2 .5/75 3/25 4/67 7/53 5/75 30 19/10 32/4 33/8 5 12 5 3 t 14/7 4/36 3 22 23 31/1 29/9 26 32/2 39/2 20 4 23 21 4 16 28 31 11 3/27 20/17 7/61 79/38 1/20 8 2 1 17 1 16/22 1 29 14 49 7 17 13 20 5 2 49/1 14/22 49/2 5 13 2 25/14 367 159 253 76 43% 19% 30% 9% 35 24 34 23 21% 58% 45% 48% *0nly species with 5% frequency of occurrence are l i s t e d . **Percent occurrence = (no. plots with spp. present) (100)/ho. plots sampled; Percent browsed = (no. plots with spp. browsed) (100)/no. plots with spp. present. 128 almost e n t i r e l y w i l l o w (72 percent) and paper b i r c h (25 per-cent) , the two most abundant browse species. In the f o r e s t , moose ate paper b i r c h and subalpine f i r i n n e a r l y equal p r o p o r t i o n s , 27 percent and 31 percent, r e s p e c t i v e l y . Considerably l e s s w i l l o w was taken i n the f o r e s t than i n the burn (16 percent versus 72 percent). Trembling aspen, squashberry ( Viburnum edule) and lungwort were taken i n the f o r e s t , but were missing from the d i e t s of moose i n the recent burn even though aspen was common i n the burn. H a b i t a t - r e l a t e d d i f f e r e n c e s i n food h a b i t s were a l s o i n d i c a t e d by the r e s u l t s from the post-winter browse surveys. Although these surveys provided only frequency of occurrence data, they d i d a l l o w l i m i t e d comparisons between the a v a i l a b i l i t y and u t i l i z a t i o n of browse on a h a b i t a t b a s i s . In the f o r e s t , w i l l o w , r e d - o s i e r dogwood, e l d e r b e r r y {Sambuous racemosa) and paper b i r c h were browsed most frequent-l y but each occurred i n l e s s than f i v e percent of the 367 p l o t s examined. The most common taxons encountered, v a c c i n i a (Vaooinium spp. ), regenerating subalpine f i r , black twinberry '(Loniaera involuorata), f l a t - t o p s p i r e a (Spiraea tuoida) and rose (Rosa spp.), were e i t h e r very l i g h t l y browsed or not touched. In a l l , only 21 percent of the 35 species recorded i n f o r e s t types were taken. A s i m i l a r p a t t e r n occurred i n the p a r t i a l cutover type. The most f r e q u e n t l y browsed species of w i l l o w , r e d - o s i e r dogwood, Douglas maple, trembling aspen, Saskatoon (Amelanohier alnifolia) and paper 129 b i r c h were among the l e a s t f r e q u e n t l y encountered species. S i m i l a r l y , regenerating subalpine f i r , v a c c i n i a , f l a t - t o p s p i r e a and rose occurred f r e q u e n t l y ( i n more than 3 9 percent of the p l o t s ) but were unbrowsed or only taken i n l i m i t e d amounts. In c o n t r a s t w i t h the f o r e s t - t y p e , however, 45 percent of the 3 4 recorded species were browsed. Results f o r both the c l e a r c u t and recent burn types d i f f e r e d from the two foregoing types and between themselves (Table 4.7). In the c l e a r c u t , the commonest species were the deciduous shrubs, rose, thimbleberry (Rubus parviflorus), raspberry (Rubus idaeus), w i l l o w , black twinberry and paper b i r c h : the most f r e q u e n t l y browsed were r e d - o s i e r dogwood, lodgepole pine, v a c c i n i a and Douglas maple. Over 50 percent of the l i s t of 2 4 species recorded were browsed. In the recent burn, w i l l o w , Saskatoon, black twinberry, goose-b e r r i e s / c u r r a n t s (Ribes spp. ) , and white spruce were commonest but of these, only w i l l o w was browsed f r e q u e n t l y . Other important browse species were a l d e r , p r i n c e ' s pine (Chimaphila umbellata) and r e d - o s i e r dogwood. S i m i l a r to the c l e a r c u t , about 50 percent of the l i s t of 23 recorded species were taken by moose. Several p l a n t species showed c o n s i s t e n t p a t t e r n s , whatever the h a b i t a t - t y p e . For example, r e d - o s i e r dogwood was always uncommon but always f r e q u e n t l y used; gooseb e r r i e s / c u r r a n t s , rose, thimbleberry and f l a t - t o p s p i r e a were always common and always l i g h t l y used. Other 130 species were browsed more f r e q u e n t l y i n the open h a b i t a t -types than i n the f o r e s t . Examples of t h i s i n c l u d e black twinberry, subalpine f i r and paper b i r c h . A l s o , a greater p r o p o r t i o n of the species found growing i n open s i t e s were browsed (45-50 percent) than, i n the f o r e s t (21 percent). Thus species growing i n f u l l s u n l i g h t may be more p a l a t a b l e to moose, p o s s i b l y due to higher l e v e l s of carbohydrates and other n u t r i e n t s (Laycock and P r i c e 197 0). 4.4 Dis c u s s i o n 4.4.1 Methodology The strengths and weaknesses of these three, commonly-used methods to c o l l e c t food h a b i t information can be seen since a l l were used i n the same area over the same time p e r i o d . Rumen a n a l y s i s provided t i m e - s p e c i f i c but u s u a l l y not s i t e - s p e c i f i c data and so food preferences were not determinable. Rumen a n a l y s i s s u f f e r s from other w e l l -known weaknesses (e.g., Bergerud and R u s s e l l 1964). However, a n a l y s i s of pr o p e r l y taken samples probably provides the best way of d e s c r i b i n g the range of p l a n t species eaten. Rumen a n a l y s i s i s a l s o valuable where moose feed i n p l a n t communities that are too complex or too d i f f i c u l t to allo w d i r e c t observation. T r a i l i n g can be both time- and s i t e - s p e c i f i c , e s p e c i a l l y i n winter when the occurrence of snowfalls can be used to age t r a c k s . Food preferences can a l s o be 131 determined, although t h i s becomes very time-consuming i f t r a i l s go through many p l a n t communities or wander along ecotones. The major weakness of t r a i l i n g i s i n overlooking p l a n t s , e s p e c i a l l y aquatic and ar b o r e a l species. In t h i s study, the v a r i e t y of l i c h e n s taken, and the p r e v i o u s l y unrecorded use and importance of lungwort would have been overlooked i f only t r a i l i n g had been used. Post-winter p l o t s t u d i e s are s i t e - s p e c i f i c but not t i m e - s p e c i f i c except to one winter. General information on a v a i l a b i l i t y of foods can be c o l l e c t e d but t h i s must be , c a r e f u l l y i n t e r p r e t e d , c o n s i d e r i n g over-winter snow co n d i t i o n s and depths. S i m i l a r to t r a i l i n g s t u d i e s , some pl a n t species can be overlooked or t h e i r importance underrated. Thus a l l three methods have l i m i t a t i o n s i n d e s c r i b -ing food h a b i t s . The best approach i s to use at l e a s t two or p r e f e r a b l y more techniques that w i l l enable a b e t t e r d e s c r i p t i o n of the v a r i e t y of p l a n t s eaten, t h e i r r e l a t i v e importance, t h e i r preference i n r e l a t i o n to a v a i l a b i l i t y , and the a c t u a l amounts eaten. 4.4.2 V a r i a t i o n s i n the Di e t This study has shown tha t the d i e t s of moose not only vary s e a s o n a l l y , but a l s o s p a t i a l l y . Seasonal v a r i a t i o n s are documented i n e a r l i e r s t u d i e s , e.g., Peek (1974b), but published accounts of d i f f e r e n c e s between 132 h a b i t a t f o r the same general area and the same winter are uncommon. These s p a t i a l v a r i a t i o n s probably are due to d i f f e r e n c e s i n both the p a l a t a b i l i t y and a v a i l a b i l i t y of food species. Shrubs growing i n open h a b i t a t s g e n e r a l l y have comparatively high carbohydrate l e v e l s (Oldenmeyer 1974, Laycock and P r i c e 1970). Since carbohydrate l e v e l s are r e l a t e d to p a l a t a b i l i t y , open-grown shrubs of the same species can be u s u a l l y expected to be browsed more h e a v i l y than p l a n t s i n the f o r e s t . (Of course, other f a c t o r s i n a d d i t i o n to carbohydrates determine p a l a t a b i l i t y of forages.) This p a t t e r n was recorded f o r a l l species i n Table 4.7 except f o r e l d e r b e r r y . A v a i l a b i l i t y a l s o i n f l u e n c e s d i e t . This u s u a l l y r e f e r s to absolute abundance of a species (e.g., cover, phytomass, p r o d u c t i o n ) , but a l s o should r e f e r to other, l e s s obvious d i f f e r e n c e s . One of these i s the h a b i t a t -r e l a t e d d i f f e r e n c e s i n a species growth form. A Saskatoon p l a n t i n the f o r e s t o f t e n c o n s i s t s of one or two, t a l l s p i n d l y s t a l k s ; i n the open, the same species i s bushy wi t h many s t a l k s . I f moose s e l e c t shrubs on the b a s i s of form, then these shape d i f f e r e n c e s may be important. Use of species may a l s o depend on a t h r e s h o l d of abundance, th a t i s , a c e r t a i n amount of a species must be present before a feeding moose w i l l f i n d or \"choose\" i t . Type-related c l i m a t i c e f f e c t s a l s o modify a species' a v a i l a b i l i t y . For example, subalpine f i r regeneration i n a f o r e s t may be above 133 the snow, but the t i p s can be covered by ice-encrusted snow caps formed by water and snow d r i p p i n g from the canopy and f r e e z i n g on the understory p l a n t s . This may help to e x p l a i n why some i n d i v i d u a l s of a species are h e a v i l y browsed while adjacent ones are unbrowsed. 4.4.3 Some Management Im 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 Moose eat a wide v a r i e t y of foods. Thus they are able to use many stages of the su c c e s s i o n a l vegetation that develop a f t e r logging. For example, lodgepole pine s i t e s provide forage i n the e a r l y s u c c e s s i o n a l stages a f t e r logging when deciduous species f l o u r i s h . As the f o r e s t develops, the understory decreases almost to the p o i n t where no forage i s produced. As the stand matures, the canopy opens, enabling shrubs and l i c h e n s to develop. These su c c e s s i o n a l trends are t r e a t e d more f u l l y i n Section 7. Because moose have a d i v e r s e d i e t does not mean tha t c u t t i n g v i r t u a l l y any stand w i l l b e n e f i t moose. This i s an o v e r s i m p l i f i c a t i o n because i t overlooks the concept of forage a v a i l a b i l i t y ; the changing n u t r i t i o n a l needs of moose, both seasonally and annually; the r e l a t i v e balance between q u a l i t y and q u a n t i t y of forage; the changes i n p l a n t n u t r i e n t l e v e l s due to s o i l moisture (Peek et a l . 1976) and canopy c l o s u r e (Cowan et a l . 1950); and the use of seasonal ranges by moose. A l l these f a c t o r s must be considered when 134 assessing the impacts of timber h a r v e s t i n g and post-logging treatments on forage production. Although moose eat many p l a n t species, r e l a t i v e l y few make up most of t h e i r d i e t (Table 4.5). For n o r t h -c e n t r a l moose, these s t a p l e species are subalpine f i r , a l d e r ( i n s p r i n g ) , paper b i r c h , r e d - o s i e r dogwood, and w i l l o w . I t seems s e n s i b l e to emphasize the e f f e c t s of timber management p r a c t i c e s on the production of these species. The v a r i a b i l i t y i n the d i e t r a i s e s concerns about the nature and r o l e of f o r e s t s reserved from logging. In f o r e s t development plans, immature stands are of t e n reserved to meet the needs of moose f o r winter cover. While these stands may provide cover, they u s u a l l y do not have commonly eaten food species. Immature stands t y p i c a l l y l a c k p l a n t species g e n e r a l l y found i n o l d e r f o r e s t s such as subalpine f i r , squashberry, and lungwort. How v i t a l these winter foods are to moose i s unknown\u00E2\u0080\u0094our knowledge of moose n u t r i t i o n a l needs are too incomplete to provide d e c i s i v e data. However, r e l y i n g p r i m a r i l y on immature f o r e s t s f o r food and cover may be l i m i t i n g options f o r moose production. Another c o n s i d e r a t i o n r e l a t e s to h a b i t a t management. In n o r t h - c e n t r a l B r i t i s h Columbia, complete c l e a r c u t t i n g i s the main h a r v e s t i n g method p r a c t i c e d . The same can be s a i d f o r most of b o r e a l North America and Scandinavia. I t i s ther e f o r e the major method of manipulating moose h a b i t a t . While t h i s technique may be the best way of har v e s t i n g wood f i b r e i t i s not always b e n e f i c i a l f o r moose. As Peek (1974b) and t h i s study pointed out, l o c a l v a r i a t i o n s i n forage preferences are e s p e c i a l l y r e l e v a n t i f h a b i t a t management p r a c t i c e s are to favour the l o c a l l y p r e f e r r e d species. I f logging i s the main management p r a c t i c e and moose production i s the major aim i n an i n t e g r a t e d f o r e s t management pl a n , h a r v e s t i n g systems and s i l v i c u l t u r a l techniques should be s e l e c t e d to create post-logging v egetation that w i l l be most productive f o r moose. This type of p r e d i c t i v e i n f o r m a t i o n i s c r i t i c a l , yet i s l a c k i n g i n many place s . 4.4.4 Future Research This study r a i s e s s e v e r a l questions f o r f u r t h e r research. The s p a t i a l v a r i a t i o n i n food h a b i t s r a i s e s the question of the r e l a t i o n s h i p between d i e t , home range and over-winter s u r v i v a l of moose. I f moose i n wi n t e r , e s p e c i a l l y l a t e w i n t e r , are sedentary as suggested by van Ballenberghe and Peek (1971), then the h a b i t a t mosaic of these home ranges i s important. Moose r e s t r i c t e d to an immature lodgepole pine stand would probably f a r e more poo r l y than moose i n an o l d stand; and both would be worse o f f than moose i n a heterogeneous home range c o n s i s t i n g of both cover- and food-producing h a b i t a t - t y p e s . The important 136 questions then become how big are home ranges, what mix of habitat-types w i l l produce a sustained moose population s u f f i c i e n t to meet public demands, and w i l l t h i s mix be compatible with other resource users? Pood habit studies provide part of the information needed to answer these questions, provided, of course, that .these studies are properly interpreted. Gullion (1966) offered some s t r i k i n g examples of how improperly interpreted food habits data can lead to inadequate and counter-productive forest management guidelines. A second area of research relates to factors influencing the a v a i l a b i l i t y and p a l a t a b i l i t y of food species. Disentangling the complex of factors such as s o i l , l i g h t , plant competition, species d i v e r s i t y , chemical composition, browsing history, plant strategies and moose behaviour requires long-term, well-designed experiments. Many studies e x i s t for domestic stock in temperate regions but w i l d l i f e - o r i e n t e d projects, e s p e c i a l l y for boreal regions, are v i r t u a l l y non-existent. A t h i r d area of research i s to improve ways of vassessing the n u t r i t i v e values of forage i n terms that are meaningful to the animal, that i s , b i o l o g i c a l l y interpret-able. This research need i s discussed more f u l l y i n Section 8. 5. DYNAMICS OF WINTER BROWSING 5.1 I n t r o d u c t i o n The previous s e c t i o n described the v a r i e t y of foods taken by moose, and t h e i r r e l a t i v e preference f o r s p e c i f i e d periods of time and l o c a l i t i e s . However, i t d i d not provide answers to questions about browsing dynamics. When were c e r t a i n winter h a b i t a t s used by moose f o r feeding? Were twigs on a p a r t i c u l a r p l a n t browsed more than once? Were p l a n t s browsed repeatedly? Did l e v e l s and r a t e s of browsing vary between p l a n t s and between h a b i t a t s ? How oft e n was more than c u r r e n t annual growth removed by moose? What l e v e l s of u t i l i z a t i o n were browse species experiencing? To answer these questions, the course of i n d i v i d u a l -l y marked twigs of four major food species was followed i n four h a b i t a t s through a winter season. Not a l l twigs of a pl a n t were tagged. Instead, the tagged twigs were considered as a sample of the a v a i l a b l e browse f o r a p a r t i -c u l a r species i n a p a r t i c u l a r h a b i t a t . The a v a i l a b l e browse was put on a weight b a s i s by measuring twig diameters and then converting them to oven-dried weights by appropriate r e g r e s s i o n equations (cf. Peek 1970, T e l f e r 1969). Thus i f 50 percent of the tagged twigs of paper b i r c h i n a burn h a b i t a t were browsed, then the estimated l e v e l of browsing 137 138 on paper b i r c h i n that h a b i t a t was a l s o 50%. The e v a l u a t i o n of browsing can be based on s e v e r a l c r i t e r i a . For t h i s s e c t i o n , browsing i s examined w i t h respect to i n c i d e n c e , time, and l e v e l of u t i l i z a t i o n . Incidence i s concerned w i t h whether or not a twig or p l a n t was browsed, and how many times were they used (Section 5.3.1). Time of browsing deals w i t h the temporal aspect of incidence, that i s , i n what months were twigs and p l a n t s browsed by moose (Section 5.3.2). Level of u t i l i z a t i o n d e s cribes how much browse was removed both by month and over the e n t i r e w i n t e r , November to A p r i l 1972-73 (Section 5.3.3) . U t i l i z a t i o n a l s o tends to be an ambiguous term. In the context of t h i s s e c t i o n , i t r e f e r s to browse removed on the b a s i s of oven-dried weight. I t does not r e f e r to time spent i n a h a b i t a t , although moose obviously spent time i n a h a b i t a t w h i l e browsing. I t does not r e f l e c t d e n s i t i e s of moose, since a given l e v e l of use can be s a t i s f i e d by many combinations of occupancy p e r i o d , age/sex s t r u c t u r e and moose d e n s i t y . 5.2 Methods Study s i t e s were lo c a t e d on the i n t e n s i v e study areas, v i z . , Eagle, Grove and Salmon. The major h a b i t a t -types were coniferous f o r e s t , deciduous f o r e s t , burn, p a r t i a l cutover. A l l were represented by one or two s i t e s 139 at each study area. S i t e d e s c r i p t i o n s are given i n Section 2.2. T y p i c a l l y , each s i t e c o n s i s t e d of f i v e s t a t i o n s spaced at i n t e r v a l s of approximately 30 m along a t r a n s e c t . At each s t a t i o n , u s u a l l y f i v e twigs on each of ten p l a n t s were marked w i t h individually.numbered aluminum tags, f o r a t o t a l of 250 twigs per s i t e (Figure 5.1). The species tagged were paper b i r c h , w i l l o w , subalpine f i r and r e d - o s i e r dogwood -the major winter foods of moose i n the region - w i t h the most abundant species tagged at each s i t e . In most cases only one or two species were sampled as other shrubs were uncommon. The i n i t i a l sampling was done i n October 1972, except f o r the r i v e r v a l l e y . s i t e at Salmon which was set up i n November. At t h i s time, the b a s a l diameters of curren t annual growth were measured to the nearest 0.1 mm, w i t h v e r n i e r c a l i p e r s , and entered on standard data sheets s u i t a b l e f o r d i r e c t key punching. A l l s i t e s were r e - v i s i t e d at monthly i n t e r v a l s . I f twigs were browsed, the diameter at p o i n t of browsing, the DPB of Peek (1971), was measured (Figure 5.1) and recorded. Monthly v i s i t s concluded i n e a r l y May, a f t e r snow melt was l a r g e l y completed. Moose were d i s p e r s i n g to summer ranges and the s w e l l i n g and new growth of twigs prevented f u r t h e r a p p l i c a t i o n of the method. For the twig data, diameters were converted to oven-d r i e d weight ( i n grams) by using the f o l l o w i n g l i n e a r r e g r e s s i o n equations developed p r e v i o u s l y : 140a Figure 5.1 Photographs i l l u s t r a t i n g methods of tagging twigs and measuring diameters at p o i n t of browsing. subalpine f i r : l o g (y) = 1.908 l o g (x) + 1.883, S = 0.118, n = 60, r 2 = 0.82 y. x wi l l o w : l o g (y) = 3.19 l o g (x) - 4.01, S = 0.034, n = 120, r 2 = 0.99 y .x paper b i r c h : l o g (y) = 2.95 log (x) - 3.34, S = 0.040, n = 90., r 2 = 0.98 y. x re d - o s i e r dogwood: l o g (y) = 0.92 l o g (x) + 0.22, S = 0.85, n = 37, r 2 = 0.84 y. x where, y = oven-dried weight i n grams, and x = diameter of twig or p o i n t of browsing as measured i n the f i e l d i n mm. The converted data were then analyzed by c a l c u l a t i n g the a v a i l a b l e browse, monthly amounts browsed, t o t a l amount browsed, and the percent u t i l i z a t i o n f o r the winter. The a v a i l a b l e browse was defined by the diameters measured at the i n i t i a l sampling. The monthly amount browsed was c a l c u l a t e d by s u b t r a c t i n g the current month's dry weight from the previous month's dry weight. For example, the amount browsed i n December equalled the estimated dry weight i n November, minus the estimated dry weight i n December. The t o t a l amount browsed was c a l c u l a t e d as the sum of the monthly amounts browsed, and the percent u t i l i z a t i o n was c a l c u l a t e d as the pr o p o r t i o n of the t o t a l weight browsed to the a v a i l a b l e weight of browse. A l l c a l c u l a t i o n s were done using a program c a l l e d 143 TTT (tagged twig transect) w r i t t e n i n F o r t r a n 10-G f o r use on an IBM/3 60 computer. The program l i s t i n g w i t h explanatory notes i s on f i l e at the o f f i c e of the W i l d l i f e Research and Technical Services 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. 5.3 Results 5.3.1 The Incidence of Use The f o l l o w i n g data-base was e s t a b l i s h e d on the three study areas. T h i r t e e n s i t e s were e s t a b l i s h e d , each w i t h f i v e s t a t i o n s . At these 65 s t a t i o n s , 638 p l a n t s were sampled f o r a t o t a l of 2,953 twigs tagged. A l l s i t e s were checked e i g h t times, except f o r one s i t e on the Salmon area which was sampled seven times. A t o t a l of 25,116 observations were made during the 1972-73 winter. Incidence of use on a t w i g - b a s i s v a r i e d widely between study areas (Table 5.1). Combining a l l species and a l l s i t e s , the Eagle area had the highest r a t e of browsing, w i t h 3 9 percent of the twigs browsed at l e a s t once. For Salmon the p r o p o r t i o n was 3 4 percent, followed by 15 percent f o r Grove. A p a r a l l e l trend was noted f o r the p r o p o r t i o n of twigs browsed twice w i t h values of 3 percent, 2 percent and 1 percent f o r the three study areas, r e s p e c t i v e l y . Combining the data f o r a l l three areas, 7 0 percent of the twigs were unbrowsed, 3 0 percent were browsed once and 2 percent were browsed twice. 144 Table 5.1 Proport i o n s of Twigs that were Browsed Once and Twice i n Major H a b i t a t s on the Eagle, Grove and Salmon Winter Ranges During the 1972-73 Winter STUDY AREA Habitat-type No. of twigs browsed (%) Month of second use* No. of twigs once twice N D J F M A EAGLE: Conifer f o r e s t 4 2 X X 226 Burn ecotone 39 1 X X X 274 Burn centre 58 5 X X X 240 P a r t i a l cutover 54 4 X X X X 236 39 3 GROVE: Conifer f o r e s t 1 286 Burn ecotone 19 1 X 310 Burn centre 30 1 X X X X 251 Upland burn 1 150 15 1 SALMON: Conifer f o r e s t 10 231 Deciduous f o r e s t 47 2 X X X 250 P a r t i a l cutover 38 4 X X X 250 River bottom 76 1 X X X X 249 34 1 ALL AREAS 30 2 X X X X X X 2,953 *Month i n which twigs browsed p r e v i o u s l y i n the 1972-73 winter were browsed again. 145 Incidence of use on a p l a n t - b a s i s a l s o v a r i e d widely between study areas (Table 5.2). A higher p r o p o r t i o n of p l a n t s was u t i l i z e d on the Salmon and Eagle study areas (58 percent and 49 percent) than on the Grove area (25 percent). A s i m i l a r ranking occurred w i t h p r o p o r t i o n of p l a n t s browsed twice w i t h 13 percent, 13 percent and 4 percent, r e s p e c t i v e l y . Repeated browsing on a p l a n t - b a s i s was much higher than repeated browsing on twi g - b a s i s . This i n d i c a t e d t h a t moose returned at l e a s t once to as many as 17-33 percent of p r e v i o u s l y browsed p l a n t s , but r a r e l y took twigs browsed.earlier t h a t winter. The p r o p o r t i o n of sampled p l a n t s that were browsed v a r i e d between species (Table 5.2). Subalpine f i r was browsed l e a s t f r e q u e n t l y on a l l study areas, w i t h an average .of 8 6 percent p l a n t s unbrowsed and 14 percent p l a n t s browsed once. I n d i v i d u a l p l a n t s of t h i s species were used l e a s t at Grove (8 percent browsed once), and approximately e q u a l l y at Salmon and Eagle- (18.5 percent browsed once). P r o p o r t i o n a t e l y more w i l l o w p l a n t s were browsed than subalpine f i r p l a n t s . Approximately 44 percent were browsed once, 11 percent were browsed twice and 3 per-cent were taken three or four times. Fewer w i l l o w s were browsed at Grove than at the other two winter ranges. For paper b i r c h , approximately 56 percent of the p l a n t s were browsed at l e a s t once, 16 percent browsed at l e a s t twice and 2 percent at l e a s t three times. Paper b i r c h p l a n t s were 146 Table 5.2 Number of Times B l a n t s of Subalpine F i r , Eaper 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 SPECIES No. of times p l a n t s were browsed (%)* No. p l a n t s Study area 0 1 2 3 4 i n sample SUBALPINE FIR: Eagle 82 18 50 Grove 92 8 60 Salmon 81 19 35 Means 86 14 145 PAPER BIRCH: Eagle 37 63 17 1 120 Grove 74 26 5 2 57 Salmon 27 73 24 5 55 Means 44 56 16 2 232 RED-OSIER DOGWOOD: Salmon (mean) 22 78 12 50 WILLOW: Eagle 54 46 17 35 Grove 65 35 7 1 85 Salmon 41 59 13 4 2 46 Means 56 44 11 2 1 166 STUDY AREA SUMMARY: Eagle 51 49 13 205 Grove 75 25 4 1 202 Salmon 42 58 13 3 196 Means 56 44 10 1 t 603 *e;g., 1 = p l a n t s browsed at l e a s t once, some browsed more than only once. * * t i s l e s s than 1 percent. used most f r e q u e n t l y at the Salmon study area, l e s s f r e q u e n t l y at Eagle, and l e a s t f r e q u e n t l y at Grove. Results f o r r e d - o s i e r dogwood showed that 7 8 percent of the p l a n t s were browsed at l e a s t once, and 12 percent at l e a s t twice. Thus incidence of browsing on p l a n t s i n descending i n t e n s i t y was Salmon, Eagle and Grove. A comparable ranking f o r the browse species was r e d - o s i e r dogwood, paper b i r c h , w i l l o w , and subalpine f i r . Thus v a r i a t i o n s i n evidence of use was great between the species on i n d i v i d u a l study areas. The v a r i a t i o n i n incidence of use was s l i g h t l y l e s s w i t h i n each species. Incidence of browsing on p l a n t s was a l s o compared between h a b i t a t s (Table 5.3). The n u l l hypothesis t e s t e d was that among the species or h a b i t a t s t e s t e d , the proportions browsed were the same. The p r o p o r t i o n of subalpine f i r p l a n t s browsed (16 percent) was e s s e n t i a l l y the same on a l l three study areas ( x 2 = 0.97, 2 df, P = 0.62). Moreover, the p r o p o r t i o n of w i l l o w and paper b i r c h p l a n t s browsed d i d not d i f f e r s i g n i f i c a n t l y w i t h i n h a b i t a t -types nor were there s i g n i f i c a n t d i f f e r e n c e s between study areas ( x 2 = 4.09, 9 df, P = 0.91). Thus i t appeared th a t moose on a l l areas t r e a t e d paper b i r c h and w i l l o w s i m i l a r l y w i t h respect to p r o p o r t i o n of p l a n t s browsed. Comparisons between a l l s i t e s w i t h paper b i r c h and w i l l o w w i t h red-o s i e r dogwood were not s i g n i f i c a n t l y d i f f e r e n t (x 2 = 5.88, df = 11, P = 0.88). S i g n i f i c a n t l y fewer subalpine f i r 148 Table 5.3 Pr o p o r t i o n of P l a n t s Browsed, by Species and by Habitat i n the 1972-73 Winter Number of p l a n t s that were browsed (%) STUDY AREA + Habitat-type Abies Betula Cornus Salix EAGLE: Conifer f o r e s t 18% (50)* Burn ecotone 60% (30) 44% (25) Burn centre 64%* (50) P a r t i a l cutover 63% (40) 50% (10) GROVE: Conifer f o r e s t * * 12% (51) Burn ecotone** 35% (60) Burn centre 4.4% (25) 36% (25) Upland burn 7% (30) SALMON: Conifer f o r e s t 20% (35) 7% (15) Deciduous f o r e s t 76% (25) 68% (25) P a r t i a l cutover 64% (25) 52% (25) River bottom 78%* (50) HABITAT SUMMARY: Coni f e r f o r e s t 16% (137) 7% (15) Deciduous f o r e s t 76% (25) 68% (25) Burn ecotone 60% (30) 38% (85) Burn centre 57% (75) 36% (25) Upland burn 7% (30) P a r t i a l cutover 63% (65) 51% (35) River bottom 78% (50) ALL DATA MEANS 16% (137) 55% (225) 62% (65) 45% (170) *No. of p l a n t s i n sample. * * S i t e s 1 and 2 included f o r e s t and burn o r i g i n a l l y . They were r e -grouped as i n d i c a t e d above f o r t h i s , p r e s e n t a t i o n . + F i e l d s i t e numbers are 1, 2, 4, 3; 1 and 2, 1 and 2, 4, 3; 2, 1, 3, 5, r e s p e c t i v e l y . 149 p l a n t s were browsed than r e d - o s i e r dogwood p l a n t s ( x 2 = 28.29, 3 df, P < 0.001), and than paper b i r c h and w i l l o w p l a n t s ( x 2 = 32.01, 13 df, P < 0.001). 5.3.2 The Time of Use Data on the time of use i n d i c a t e d s e v e r a l general p o i n t s . F i r s t , browsing occurred at a l l study areas i n a l l months (Table 5.4). Second, a l l species were browsed i n a l l months, except f o r r e d - o s i e r dogwood which was h i g h l y p r e f e r r e d yet taken only i n January and February. T h i r d , only the deciduous f o r e s t at the Salmon area was browsed i n every month. A l l other h a b i t a t s were used f o r two-five of the s i x winter months. Each study area showed some d i f f e r e n c e s from the foregoing g e n e r a l i t i e s (Table 5.4). At Eagle, the c o n i f e r f o r e s t was browsed p r i m a r i l y i n the l a s t h a l f of the w i n t e r , but both burn s i t e s experienced browsing almost e n t i r e l y from November to February. The p a r t i a l cutover was used f o r feeding i n a l l months except March. At Grove, the c o n i f e r f o r e s t was used i n November and i n A p r i l r a t h e r than mostly i n l a t e winter as at Eagle. These d i f f e r e n c e s may r e f l e c t s n o w f a l l p a t t e r n s . A lso d i f f e r e n t from Eagle was the occurrence of browsing at the burn s i t e s i n a l l w i n t e r months rather than i n e a r l y w inter. At Salmon, s i m i l a r i t i e s e x i s t e d w i t h the two other areas. S i m i l a r to Grove, the burn or deciduous f o r e s t was used i n a l l months. S i m i l a r to Table 5.4 Time of Browsing and Lev 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 STUDY AREA % t o t a l use removed, by month Over-winter Browse Habitat Spp.* Nov. Dec. Jan. Feb. Mar. Apr. use (%) a v a i l a b l e EAGLE: Con i f e r f o r e s t S 1 66 22 12 4 8,825 Burn ecotone B 32 62 1 6 23 60 W 20 30 12 38 23 89 Burn center B 42 49 10 t 29 105 P a r t i a l cutover B 1 80 5 15 47 90 W 92 3 6 1 25 31 GROVE: Conifer f o r e s t s 2 98 2 26,700 Burn ecotone w 2 8 61 6 23 14 193 Burn center B 9 71 8 2 9 17 58 w 6 94 17 46 Upland burn B 50 t 67 SALMON: Conifer f o r e s t R 100 11 212 S 62 38 10 4,486 Deciduous f o r e s t B 3 46 9 22 2 17 36 96 W 6 32 18 16 1 27 63 47 P a r t i a l cutover B 51 31 10 8 43 53 W 38 26 5 30 38 70 River bottom R NS** 96 4 36 1,059 Table 5.4, Continued STUDY AREA % t o t a l use removed, by month Over-winter Browse Habi t a t Nov. Dec. Jan. Feb. Mar. Apr. use (%) a v a i l a b l e AREA SUMMARY: Eagle 5 17 2 51 16 10 5 9,199 Grove 2 1 7 1 t 90 2 27,061 Salmon t 5 69 6 18 2 16 6,024 SPECIES SUMMARY: Abies lasiooarpa (subalpine f i r ) 1 t 23 18 20 37 3 40,010 Betula papyrifera (paper b i r c h ) 12 55 14 6 2 10 29 528 Cornus stolonifera ( r e d - o s i e r dogwood) NS 96 4 33 1,271 Salix spp. (willow) 5 30 32 13 t 19 25 476 HABITAT SUMMARY: Co n i f e r f o r e s t 1 t 23 20 20 36 3 40,222 Deciduous f o r e s t \u00E2\u0080\u00A25 39 13 19 2 22 45 143 Burn ecotone 15 27 31 17 11 18 341 Burn center 26 34 36 2 t > 2 23 209 Upland burn 50 50 t 69 P a r t i a l cutover t 63 16 2 2 16 41 244 R i v e r bottom NS 96 4 36 1,059 OVERALL SUMMARY: 2 7 38 15 13 25 4 42,284 *Species abbreviated as S = subalpine f i r , B **NS - s i t e not set up t h i s month. = paper b i r c h , W = w i l l o w , R = re d - o s i e r dogwood. 152 Eagle, the c o n i f e r f o r e s t was used i n l a t e w i n t e r , and the p a r t i a l cutover used throughout the period. The r i v e r bottom type, s i t e d only at Salmon, was used f o r feeding only during January and February. Thus the time of browsing i n h a b i t a t s was g e n e r a l l y s i m i l a r on the three study areas. 5.3.3 The L e v e l of Use The t i m i n g and amount of browse eaten by moose .varied c o n s i d e r a b l y between study areas, shrub species and h a b i t a t s (Table 5.4). The Salmon range received the heaviest l e v e l of browsing, where moose ate an estimated 16 percent.of the tagged 1972 annual growth. Moose took much l e s s of the a v a i l a b l e browse at the Eagle and Grove ranges where only 5 percent and 2 percent, r e s p e c t i v e l y , of the tagged twig phytomass was removed. These d i f f e r e n c e s between study areas were not a r e s u l t of comparable d i f f e r e n c e s i n populations of w i n t e r i n g moose. Rather, they probably r e f l e c t e d s h i f t s i n the h a b i t a t s where browsing occurred. For example, only one c o n i f e r f o r e s t h a b i t a t was sampled at each study area. Other c o n i f e r stands were a v a i l a b l e and presumably used by moose. The l e v e l and timing of use of these l i k e l y v a r i e d between the study areas. In f u t u r e s t u d i e s , the sampling of the v a r i o u s h a b i t a t - t y p e s should be r e p l i c a t e d and the number of twigs sampled, reduced. U t i l i z a t i o n on a species b a s i s g e n e r a l l y r e f l e c t e d 153 changes i n d i e t as described i n Section 4. Subalpine f i r was browsed l i g h t l y e a r l y i n the w i n t e r , but by January when moose occupied coniferous f o r e s t s , use reached 23 percent. This l e v e l was maintained during the remainder of the wint e r , ranging from 2 0 percent to 3 6 percent (Table 5.4). O v e r a l l use of subalpine f i r was very l i g h t , at 3 percent of the 197 2 growth. The l i g h t usage coupled w i t h the importance of t h i s species i n the winter d i e t (Section 4.3), demonstrates i t s abundance i n the h a b i t a t s used by moose i n l a t e w i n t e r . Paper b i r c h and w i l l o w were browsed throughout the w i n t e r , though s l i g h t l y more i n e a r l y winter than l a t e r on. For the two species, monthly l e v e l s of use were s i m i l a r . T o t a l winter use was a l i k e at.29 percent and 25 percent f o r paper b i r c h and w i l l o w , r e s p e c t i v e l y . The data f o r r e d - o s i e r dogwood were considered f o r the Salmon area as no comparable s i t e s were studied at Eagle and Grove. Apparently, r e d - o s i e r dogwood was u t i l i z e d very h e a v i l y for. a short period. V i r t u a l l y a l l use i n the r i v e r bottom type occurred i n January, when moose were d r i v e n by deep snow to low e l e v a t i o n s . A f t e r t h i s month, c o n t i n u i n g snowfalls covered most p l a n t s of t h i s species. Compared to w i l l o w and paper b i r c h , r e d - o s i e r dogwood i s low-growing i n most s i t e s . T o t a l use of dogwood was 33 percent, the highest recorded f o r the four species studied. H a b i t a t s were used d i f f e r e n t i a l l y (Table 5.4). Generally, r e s u l t s f o r the three areas were s i m i l a r . Thus reference i s only made to study areas to p o i n t out s p e c i f i c s : otherwise h a b i t a t - t y p e s are described g e n e r i c a l l y . Three broad l e v e l s of use were d i s c e r n i b l e , based on winter u t i l i z a t i o n . Deciduous f o r e s t , p a r t i a l cutover and r i v e r bottom types were used most h e a v i l y w i t h 45 percent, 41 percent and 36 percent of the 1972 twig production browsed by moose, r e s p e c t i v e l y . Next were the burn h a b i t a t - t y p e s , w i t h u t i l i z a t i o n l e v e l s of approximately 20 percent. The o l d e r burn at Eagle was used s l i g h t l y more than the burn at Grove, although I considered Grove to have more a v a i l a b l e browse than Eagle. Least browsed were the coniferous f o r e s t and upland burn, w i t h over-winter u t i l i z a t i o n l e v e l s of 3 percent and l e s s than 0.5 percent, r e s p e c t i v e l y . The c o n i f e r o u s f o r e s t was represented at a l l three study areas, and a l l three s i t e s were used very l i g h t l y . The previous sub-section on incidence of use (5.3.2) showed th a t obvious d i f f e r e n c e s were apparent between h a b i t a t - t y p e s . The u t i l i z a t i o n data showed f u r t h e r that l e v e l s of use v a r i e d widely even between two months i n which browsing was recorded. For example, the f o l l o w i n g h a b i t a t -types were used i n a l l s i x winter months: c o n i f e r f o r e s t , deciduous f o r e s t , burn center and p a r t i a l cutover. Yet, the l e v e l s of use d i f f e r e d c o n s iderably. S i m i l a r t o the p a t t e r n of subalpine f i r , the c o n i f e r f o r e s t type was used very 155 l i g h t l y i n November and December, and then at approximately 25 percent f o r each of the f o l l o w i n g months. The deciduous f o r e s t was browsed h e a v i l y i n December, when migrat i n g moose l i k e l y f i r s t moved onto the winter range, and then at approximately 14 percent per month t h e r e a f t e r . The burn ecotone and burn center types were used most h e a v i l y i n November, December and January, and then l i g h t l y f o r the d u r a t i o n of the w i n t e r , averaging about 5 percent per month. The l e v e l of use at the p a r t i a l cutover type p a r a l l e l e d t h a t of the burn h a b i t a t - t y p e s . Since r e d - o s i e r dogwood was the only species i n the r i v e r bottom type, l e v e l s of use were the same f o r both p o i n t s of view, that i s , v i r t u a l l y a l l use i n January. A d d i t i o n a l r e p r e s e n t a t i o n of t h i s h a b i t a t - t y p e would have been very u s e f u l . The upland burn type occurred only at Grove. I t was v i r t u a l l y unbrowsed except f o r a few twigs taken i n December and March. Such low use occurred d e s p i t e the abundance of paper b i r c h , a key winter browse species. This s i t e c l e a r l y demonstrated the need to consider more than browse supply when e v a l u a t i n g a h a b i t a t - t y p e ' s p o t e n t i a l . The o v e r a l l summary of browsing e x h i b i t e d the f o l l o w i n g p a t t e r n . Levels of use were low i n e a r l y winter but increased sharply i n December. This upswing presumably r e f l e c t e d the increased d e n s i t y of moose on the winter ranges. Levels d e c l i n e d by more than one-half i n February and March. Perhaps t h i s r e f l e c t e d a v o l u n t a r y r e d u c t i o n i n 156 intake as has been demonstrated f o r b l a c k t a i l deer (Wood et a l . 1962) and w h i t e t a i l deer (French et a l . 1956). U t i l i z a t i o n rose sharply again i n A p r i l as snow depths d e c l i n e d and n u t r i t i v e l e v e l s i n forage improved. The o v e r a l l l e v e l of use was only 4 percent, based on a l l data. 6. BED SITE SELECTION BY MOOSE IN WINTER 6.1 I n t r o d u c t i o n M i n i m i z i n g energy l o s s e s i s a major challenge> c o n f r o n t i n g moose i n winter. Forage i s o f t e n l i m i t e d i n amount and t y p i c a l l y wanting i n n u t r i t i v e value. Movement i s r e s t r i c t e d by snow. The d i f f e r e n t i a l s between ambient and body temperatures reach t h e i r annual extremes. C h i l l f a c t o r s are o f t e n high. That moose occur so widely i n bore a l environments demonstrates t h e i r success i n adapting to these adverse winter c o n d i t i o n s . One adaptation to winter i s moose's a b i l i t y to f i n d m i c r o - s i t e s that minimize energy l o s s e s . This a b i l i t y can be seen as an attempt to l o c a t e thermoneutral environments. Obviously, such events as w i l d f i r e and logging modify the c h a r a c t e r i s t i c s and d i s t r i b u t i o n of these m i c r o - s i t e s . A f u l l a p p r e c i a t i o n of the impact of these events f i r s t r e q u i r e s an a p p r e c i a t i o n of what moose s e l e c t f o r i n choosing s h e l t e r e d l o c a t i o n s . From a management standpoint, t h i s type of information i s a p r e r e q u i s i t e to p r e s c r i b i n g logging p r a c t i c e s that w i l l minimize t h e i r adverse e f f e c t s on p o t e n t i a l s h e l t e r areas. The s h e l t e r s i t e s s e l e c t e d by moose l i k e l y vary during the year, as do t h e i r reason f o r s e l e c t i o n . In 157 158 summer, K e l s a l l and T e l f e r (1974) suggested t h a t moose seek c o o l s i t e s to avoid heat s t r e s s . However, i t i s probably i n winter t h a t moose face the gr e a t e s t need to avoid thermal extremes, e s p e c i a l l y when l y i n g down. Thus i f moose se l e c t e d p a r t i c u l a r s i t e s f o r s h e l t e r i n wi n t e r , i t would be expected to be most obvious i n t h e i r s e l e c t i o n of bedding s i t e s . More s p e c i f i c a l l y , bedding behaviour and bed s i t e f eatures can be used to t e s t the f o l l o w i n g f i v e p r e d i c t i o n s : 1) beds w i l l be lo c a t e d under denser ra t h e r than more open t r e e canopies. S e l e c t i o n w i l l be more pronounced during periods of low temperatures or deep snow or both, than i n more moderate temperatures or snow c o n d i t i o n s . 2) beds w i l l be lo c a t e d under tr e e s a f f o r d i n g g r e a t e s t s h e l t e r i n g e f f e c t s , i . e . , under white spruce and subalpine f i r r a t h e r than under lodgepole pine and Douglas f i r . Deciduous species w i l l be used only o c c a s i o n a l l y . As snow c o n d i t i o n s become p r o g r e s s i v e l y adverse, l a r g e t r e e s w i l l be se l e c t e d since they provide greater s h e l t e r than small ones. 3) i n drumlinized terrain., moose w i l l bed on upper r a t h e r than lower slopes to escape the c o o l a i r t h a t d r a i n s i n t o depressions. Beds w i l l have sout h e r l y and wes t e r l y exposures to maximize exposure to sun. 4) beds w i l l be i n wind-protected l o c a t i o n s such as behind t r e e s or shrubbery; o r i e n t a t i o n w i l l be \u00C2\u00BB towards the wind to detect p o s s i b l e predators. 5) moose w i l l s e l e c t s o f t e r snow than average since i t w i l l a f f o r d greater i n s u l a t i o n (based on Des Meules 1965). The f o l l o w i n g o b j e c t i v e s were set f o r the bed s i t e study. F i r s t , to desc r i b e g e n e r a l l y the features of beds and t h e i r s e t t i n g . Second, to q u a n t i f y snow depth and 159 hardness of the beds and the surrounding area. T h i r d , to describe the species composition, s i z e , and crown canopy-c l o s u r e of tr e e s at the bed s i t e s and the surrounding area. Fourth, to estimate the period and d u r a t i o n of occupation of beds. To ensure sampling a wide range of environments, f i e l d work was conducted i n three major h a b i t a t - t y p e s and i n three c l a s s e s of snow depths (Table 6.1). The h a b i t a t - t y p e s were coniferous f o r e s t (undisturbed canopy), s e l e c t i v e l y logged ( p a r t i a l l y a l t e r e d f o r e s t canopy), and c l e a r c u t and burns ( l i t t l e or no overstory canopy). Most s i t e s were on drumlinized t i l l m a t e r i a l s , the dominant landform and sub-s t r a t e i n the regi o n . A few were s i t u a t e d on l a c u s t r i n e d e p o s i t s , the second commonest substrate. Snow c l a s s e s were defined on the b a s i s of the ge n e r a l i z e d e f f e c t s of snow depth on moose movements as reported i n the l i t e r a t u r e (Coady 1974, Formosov 1946, K e l s a l l and P r e s c o t t 1971, T e l f e r 1970, R i t c e y 1967). In shallow snow (0-40 cm), moose experience l i t t l e or no hindrance to movement; i n medium snow (41-8 0 cm), moderate impediment to movement; and i n deep snow (greater than 81 cm), movement i s d i f f i c u l t and r e s t r i c t e d . 6.2 Methods Beds were found by f o l l o w i n g f r e s h l y made t r a i l s . Information f o r each bed was recorded on a standard data sheet to secure uniform and complete data c o l l e c t i o n 160 Table 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 Study areas by snow depth c l a s s (cm) Habitat shallow (0-40) moderate (41-80) deep (> 81) Coniferous f o r e s t Mackenzie Mtn. Te l a c h i c k Grove Limestone Creek Found Lake P a r t i a l cutover Mackenzie Mtn. Grove Found Lake Open (burn, c l e a r c u t ) Mackenzie Mtn. T e l a c h i c k Grove (Table 6.2). The f o l l o w i n g l i s t o u t l i n e s the type of data and methods used: a) d e s c r i p t i v e notes on area, h a b i t a t , date, landform, v e g e t a t i o n and snow cover. b) crown canopy c l o s u r e immediately above a bed and three m away from i t , i n a randomly s e l e c t e d d i r e c t i o n . Closure was estimated from dot g r i d counts of v e r t i c a l photographs taken w i t h a 3 5 mm camera equipped w i t h a 17 mm or f i s h - e y e lens (Brown and Worley 1965). Dot g r i d counts were d u p l i c a t e d on each photograph to estimate percent canopy c l o s u r e . c) species, number and DBH of a l l t r e e s w i t h i n a 28 m2 c i r c u l a r p l o t (R = 3m) centered i n the middle of the bed. Distance from the edge of the bed to the c l o s e s t t r e e stem was a l s o recorded to the nearest cm. This t r e e was. defined as the \" s h e l t e r \" t r e e . d) l o c a t i o n of the bed w i t h respect to the s h e l t e r t r e e (compass bearing and whether or not i n a quamaniq), p o s i t i o n on slope (1 = c r e s t of drumlin, 2 = upper h a l f , 3 = lower h a l f , 4 = swale), aspect and slope. e) snow depth i n , adjacent t o , and three m away from the bed, to the nearest cm. 161 Table 6.2 Example of Data Sheets Used to Study Bed Sites MOOSE-FORESTRY: Bed survey MOOSE NO. BED NO. AREA HABITAT-TYPE LOCATION DATE AREA FEATURES (general description) PHOTO DATA bedsite away a) vegetation b) physiography^ c) snow No. F-stop Speed d) recent weather Film type BED FEATURES - c i r c l e units of measurement a) physiography: slope % aspect \u00C2\u00B0 bed position on slope b) snow depth (cm i n ) : i n bed ajd. away_ c) distance to closest tree (cm in) ( c i r c l e closest tree below) d) i n quamaniq: yes no e) excretion: urine p e l l e t s none f) age: before a f t e r g) vegetation: TREES SKETCH (N) No. Spp. DBH 1. 2. _ 3. _ 4. 5. 6. cm. i n . REGEN Spp. No. stems SHRUBS Spp. No. stems 9.. 10. Notes: Distance to Forest cover, etc. 162 f) l e n g t h , width and depth of bed to the n e a r e s t cm. g) o r i e n t a t i o n of moose i n bed. T h i s was determined r e a d i l y by the low and wide d e p r e s s i o n produced i n the bed by the moose's h i n d q u a r t e r s . The p o s i t i o n i n g of f r o n t and hind l e g s i n the snow provided a check on the d e p r e s s i o n f e a t u r e s , f e c e s and u r i n e were usu a l l y , found o n l y a t one end of the bed. h) observed f e c e s and u r i n e . i ) estimated time o f . f o r m a t i o n , based on s n o w f a l l r e c o r d s . j) a sketch of the bed, drawn to s c a l e . Unusual f e a t u r e s or o b s e r v a t i o n s on moose behaviour were a l s o recorded on these sheets. 6.3 R e s u l t s A t o t a l of 94 beds were examined d u r i n g January, February, March and December, 1973. In s i z e , they averaged 145 \u00C2\u00B1 26 (sd) cm long by ,94 \u00C2\u00B1 11 cm wide (n = 46) . The time spent i n a bed c o u l d not be documented d i r e c t l y . R e l a t i v e d i f f e r e n c e s were determined, however, by assuming t h a t the amount of e x c r e t a i n a bed was d i r e c t l y r e l a t e d to time spent t h e r e . Thus i t appeared t h a t time spent i n beds v a r i e d a c c o r d i n g to snow depth (Figure 6.1). The ' r e g r e s s i o n of mean numbers of p e l l e t groups p l u s u r i n a t i o n s , and snow depth was s i g n i f i c a n t ( c a l c u l a t e d F r a t i o = 6.78, t a b u l a t e d F f o r 1 and 17 df a t 95 percent p r o b a b i l i t y l e v e l = 4.45). The a p p r o p r i a t e equation was: y = 0.03 + 0.01 x, where y = mean number of p e l l e t groups and u r i n a t i o n s per 2 bed, x = mean snow depth i n cm, r =0.30 and SE of estimate =0.62. 162a Figure 6.1 The r e l a t i o n s h i p between snow depth and 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 u r i n e . 2.604 163 2.30. 2.00. 170 y -0.03+0.01 x Sy.x\u00C2\u00AB0.62 r 2 -0.30 n -18 F = 6.78 HABITAT T Y P E \u00E2\u0080\u00A2 O P E N o PARTIAL A CLOSED 140 A 1.10 0.80 \u00E2\u0080\u00A2o\u00C2\u00BB 0.50-J \u00E2\u0080\u00A2 m 0.20 20 40 60 80 \"ioo\" \u00E2\u0080\u0094 r -120 M E A N S N O W D E P T H IN C M . 164 Time spent i n beds a l s o apparently v a r i e d according to habitat-type (Table 6.3). Moose remained longest i n beds i n p a r t i a l l y logged h a b i t a t s , next longest i n open h a b i t a t s and l e a s t i n coniferous f o r e s t s . This r e s u l t showed an i n c o n s i s t e n t r e l a t i o n s h i p with snow depth; moose might be p r e d i c t e d to bed longer i n open h a b i t a t where snow packs were deeper than i n p a r t i a l l y logged stands. An explanation f o r t h i s might be that i n p a r t i a l cutovers, moose were able to secure p r o t e c t i o n adjacent to food. Thus they would not be o b l i g e d to move as f r e q u e n t l y as i n the open (food but l i m i t e d cover) and i n the f o r e s t (cover but l i m i t e d food). A l s o , i t i s l i k e l y t hat other f a c t o r s i n f l u e n c e the time spent i n beds. Thus snow depth alone i s probably an i n s u f f i c i e n t p r e d i c t o r . Moose stayed longer i n t h e i r beds during January and February than i n e i t h e r December or March (Table 6.3), perhaps due to c o l d e r temperatures. Longest stays e v i d e n t l y occurred i n February, when the mean number of p e l l e t groups and u r i n a t i o n s was 1.16, or 55% higher than the lowest mean recorded f o r March. In drumlinized t e r r a i n , moose appeared to choose p a r t i c u l a r slope p o s i t i o n s (Table 6.4A). Gene r a l l y , upper slopes were taken r a t h e r than c r e s t s , lower slopes or swales. The n u l l hypothesis of equal numbers of beds i n the four categories of slope p o s i t i o n s was r e j e c t e d ( x 2 = 10.68, 6 df, P < 0.05). The preference f o r upper slopes was most 165 Table 6.3 Time Spent by Moose i n Beds as Indicated by Feces and Urine, According to Habitat and Month A. Monthly d i f f e r e n c e i n time spent bedded down. Ratios/bed Month p e l l e t groups u r i n a t i o n s both No. of beds Dec. 0. 61 0. 18 0.79 33 Jan. 0. 87 0. 21 1.08 24 Feb. 0.79 0. 37 1.16 19 Mar. 0.50 0. 25 0.75 8 B. Habitat-\u00E2\u0080\u00A2related d i f f e r e n c e s on time spent bedded down. Ratios/bed Habitat p e l l e t groups u r i n a t i o n s both No. of beds Forest 0. 61 0. 11 0.71 28 P a r t i a l cutover 0.86 0. 45 1.32 34 Open 0.74 0.21 0. 95 22 Means/total 0.71 0. 24 0. 95 84 166 Table 6.4 Locations of Moose Beds w i t h Respect to P o s i t i o n on Slope, and Aspect A. Bed l o c a t i o n i n r e l a t i o n to slope Snow depth P r o p o r t i o n of beds . (%) by slope p o s i t i o n No. Of c l a s s c r e s t upper h a l f lower h a l f swale beds shallow 33 47 20 15 moderate 35 27 31 8 26 deep 7 50 43 14 Habitat type f o r e s t 17 52 26 4 23 p a r t i a l cutover 53 17 30 15 open 18 38 38 6 17 Mean/total 27 38 31 4 55 B. Aspect of b e d s i t e P r o p o r t i o n of beds (%) by compass p o i n t No. of N NE E SE S SW W NW beds A l l samples 5 8 14 6 22 8 8 29 63 167 pronounced f o r deep snow c o n d i t i o n s . Type of h a b i t a t d i d not appear to i n f l u e n c e s e l e c t i o n of slope p o s i t i o n (Table 6.4A). The n u l l hypothesis of equal d i s t r i b u t i o n of bed s i t e s by slope p o s i t i o n f o r the three h a b i t a t - t y p e s was not r e j e c t e d ( x 2 = 9.42, 6 d f ) . Moose and other ungulates p r e f e r s o u t h e r l y and westerly aspects i n temperate and b o r e a l l a t i t u d e s , p a r t i c u l a r l y i n areas of pronounced r e l i e f ( S t e l f o x and Taber 1969). Data from t h i s study i n d i c a t e d t h a t a s i m i l a r preference e x i s t e d but on a smaller s c a l e where r e l i e f was not pronounced ( x 2 = 11.13, 7 df, p = 0.14). I t should be r e c a l l e d t h a t the range of e l e v a t i o n i n drumlinized t e r r a i n commonly does not exceed 150 m. Based on a l l data, beds on so u t h e r l y and w e s t e r l y aspects (S ->\u00E2\u0080\u00A2 NW) made up 57 percent of a l l beds (Table 6.4). S i m i l a r to slope p o s i t i o n , the aspect preference v a r i e d w i t h snow depth. At low depths (0-40 cm), 59 percent of the beds had n o r t h e r l y aspects and none had s o u t h e r l y aspects. At intermediate depths (41-80 cm), the p r o p o r t i o n of beds on n o r t h - f a c i n g aspects d e c l i n e d to 4 8 percent and s i t e s on south-facing aspects represented 33 percent. At r e s t r i c t i v e depths (> 81 cm), only 17 per-cent of the beds were on n o r t h e r l y exposure and 72 percent on south slopes. Within a f o r e s t or logged stand, moose s e l e c t e d c e r t a i n t rees as s h e l t e r t r e e s . Choice of species v a r i e d according to species present, a v a i l a b l e t r e e diameters, and 168 p r e v a i l i n g snow c o n d i t i o n s . These f a c t o r s notwithstanding coniferous species were s e l e c t e d over deciduous species i n a l l cases (n = 8). This r e s u l t i s not s u r p r i s i n g since the s h e l t e r i n g e f f e c t of deciduous species i s l e s s than that of c o n i f e r s (cf. Des Meules 1965). .Snow depths i n paper b i r c h stands on the Eagle study area, and i n trembling aspen stands at Salmon were s i m i l a r to.those i n the adjacent open areas but deeper than those i n adjacent f o r e s t s (see Section 9.3.2). Moose d i d not s e l e c t amongst the a v a i l a b l e c o n i f e r species w i t h i n the sample p l o t ( x 2 = 4.30, 4 df) (Table '6.5A). This was unexpected since spruce and subalpine f i r were p r e d i c t e d to be b e t t e r s h e l t e r trees than Douglas f i r and lodgepole pine on the b a s i s of t h e i r crown c h a r a c t e r i s -t i c s : the two former species have dense, low canopies, while the l a t t e r species have higher, more open canopies. Beds tended to be under the l a r g e s t t r e e of the t r e e s w i t h i n the v e g e t a t i o n p l o t (Table 6.5B). This choice was most pronounced when only 2-4 t r e e s occurred i n the p l o t . This trend presumably r e f l e c t e d the more pronounced s h e l t e r i n g e f f e c t of i n d i v i d u a l t r e e s i n open-spaced stands than i n densely stocked, closed canopy f o r e s t s . F i e l d observations support t h i s : quamaniqs were r e a d i l y i d e n t i f i a b l e i n open stands but d i f f i c u l t to d i s t i n g u i s h i n denser stands such as even-aged lodgepole pine. The s h e l t e r t r e e s averaged 2 9 cm i n diameter. Mean 169 Table 6.5 Comparison of Conifer Species A v a i l a b l e as S h e l t e r Trees, With Those Used by Moose A. Tree species s e l e c t i o n Tree species (% b a s i s ) i r e e \u00E2\u0080\u00A2 -status white spruce subalpine f i r lodgepole pine Douglas f i r T o t a l s A v a i l a b l e 46% 46% 7% 1% 132 Used 44% 49%. 5% 2% 43 T o t a l s 80 82 11 2 175 B. Diameter s e l e c t i o n (only samples w i t h > two trees) No. of trees S i z e of s h e l t e r t r e e , DBH b a s i s , w i t h 1 = l a r g e s t No. of i n p l o t 1 2 3 4 5 6+' p l o t s 2 3 2 5 3 5 3 1 9 4 3 1 1 5 5 1 1 1 3 6-8 2 1 2 2 7 Totals 14 8 2 3 2 29 170 diameters of the two major species were: white spruce: 36 \u00C2\u00B1 21 cm, n = 15 subalpine f i r : 25 \u00C2\u00B1 16 cm, n = 24. These means were s i g n i f i c a n t l y d i f f e r e n t at P = 0.10 (t = 1.83, df = 37). These data were f u r t h e r analyzed by h a b i t a t . In the f o r e s t , white spruce tr e e s were l a r g e r than subalpine f i r (t = 2.46, df = 19). Diameters of the two species were not s i g n i f i c a n t l y d i f f e r e n t i n p a r t i a l cutovers (t = 0.48, df = 10) nor i n open types (t = 1.61, df = 4). Other tr e e species were not commonly used as s h e l t e r t r e e s . Two lodgepole pine s h e l t e r t r e e s were 28 and 48 cm i n diameter, and one Douglas f i r was 33 cm. The d i r e c t i o n of beds r e l a t i v e to the s h e l t e r t r e e s was not random. The n u l l hypothesis of no s i g n i f i c a n t d i f f e r e n c e i n l o c a t i o n frequencies f o r each of the ei g h t c a r d i n a l compass p o i n t s was r e j e c t e d ( x 2 = 22.25, 7 df. Tabulated x 2 = 14.1 at 95% p r o b a b i l i t y l e v e l ) . Beds were s i t e d mainly i n so u t h e r l y d i r e c t i o n s from the t r e e s ; n o r t h e r l y and we s t e r l y l o c a t i o n s occurred l e s s f r e q u e n t l y than expected (Table 6.6C). The o r i e n t a t i o n of moose i n t h e i r beds was examined. No compass po i n t was p r e f e r r e d , based on a l l a v a i l a b l e data (Table 6.6B). However, d e t a i l e d a n a l y s i s i n d i c a t e d that preferences i n e a r l y winter (Dec.-Jan.) d i f f e r e d s i g n i f i c a n t l y from that i n l a t e winter (Feb.-Mar.). For the n u l l hypothesis which po s t u l a t e d no d i f f e r e n c e s between the 171 Table 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 She l t e r Tree A. O r i e n t a t i o n w i t h respect to eight compass po i n t s Months Frequencies of occurrence (%) of o r i e n t a t i o n T o t a l compared N NE E SE S SW W NW beds Dec. 9 21 15 12 18 12 12 33 Jan. 9 4 26 4 9 9 39 23 Feb. 21 42 11 11 16 19 Mar. 25 13 13 25 13 13 8 B. O r i e n t a t i o n w i t h respect to h a b i t a t - t y p e H a b i t a t - Frequencies of occurrence (%) of o r i e n t a t i o n T o t a l type N NE E SE S SW W NW beds f o r e s t 7 10 14 14 14 31 10 29 open 9 15 15 18 3 15 12 15 34 p a r t i a l cutover 10 15 15 25 25 10 20 T o t a l no. 5 10 11 15 10 9 15 8 83 C. D i r e c t i o n w i t h respect to s h e l t e r t r e e (% b a s i s ) 12 23 12 29 5 15 3 65 172 two winter p e r i o d s , the c a l c u l a t e d c h i - s q u a r e was 14.88. As the t a b u l a t e d c h i - s q u a r e i s 14.1 at the 95 percent p r o b a b i l i t y l e v e l , the n u l l h y p o t h e s i s was r e j e c t e d . O r i e n t a t i o n a l s o v a r i e d w i t h h a b i t a t - t y p e (Table 6.6B), with, p a r t i a l c u t o v e r s d i f f e r i n g from f o r e s t s ( x 2 = 12.64, df = 7, P = 0.10) and from open types ( x 2 = 11.78, d f = 7, P = 0.10). These d i f f e r e n c e s i n o r i e n t a t i o n may have r e f l e c t e d responses of moose to wind d i r e c t i o n . P r o t e c t i o n from wind a f f o r d e d by bed s i t e s was t r e a t e d very b r i e f l y . Wind v e l o c i t i e s averaged 34 percent of those away from the s i t e (n = 4). To be c o n c l u s i v e , more data are needed. The average d i s t a n c e between a moose bed and the nea r e s t t r e e was 92 \u00C2\u00B1 51 cm (n = 3 means of 14 measurements). Although mean d i s t a n c e s v a r i e d between f o r e s t (106 cm), p a r t i a l c utover (36 cm) and open (13 5 cm) h a b i t a t types, these d i f f e r e n c e s were not s t a t i s t i c a l l y s i g n i f i c a n t based on a one way a n a l y s i s of v a r i a n c e ( c a l c u l a t e d F 2 = 0.8 66, t a b u l a t e d F a t P Q 5 = 3.98). The need f o r s h e l t e r from snow was p r e d i c t e d t o i n c r e a s e as snow pack i n c r e a s e d . T h i s p r e d i c t i o n was t e s t e d by examining four r e l a t i o n s h i p s : 1. stem diameter of s h e l t e r t r e e s and snow depth, 2. d i s t a n c e between bed and s h e l t e r t r e e and snow depths, 3. d i f f e r e n c e between canopy coverage above bed and average coverage and, 173 4. snow depth and the pr o p o r t i o n of beds i n quamaniqs. The r e l a t i o n s h i p s should be d i r e c t f o r a l l cases except number two. The stem diameters of s h e l t e r t r e e s was not r e l a t e d 2 to snow depth (r = 0.0, n =11, F r a t i o = 0.12). This r e s u l t was unexpected. I thought that moose would have chosen l a r g e r diameter t r e e s as snow depths increased. However, t h i s may not be the case f o r one or more of the f o l l o w i n g reasons: 1. snow-shielding a b i l i t y of t r e e s may not be r e l a t e d to stem diameter ( d i r e c t comparisons of crown canopy may be more r e v e a l i n g ) ; 2. moose may have s e l e c t e d f o r e s t stands r a t h e r than i n d i v i d u a l t r e e s f o r bedding; 3. i n the areas sampled, a l l c o n i f e r s may have been s u i t a b l e as s h e l t e r t r e e s . As snow pack increased, the d i s t a n c e between beds and the s h e l t e r t r e e s decreased (Figure 6.2). Again t h i s i s expected since the s h i e l d i n g e f f e c t of a tre e i s l i k e l y g r e a t e s t at i t s stem and l e a s t at the crown perimeter. The r e l a t i o n s h i p i s s t a t i s t i c a l l y s i g n i f i c a n t (P = 0.10) and i s defined by y = 142.38-1.OOx, where y = mean d i s t a n c e i n cm between a bed's edge and the s h e l t e r t r e e ' s trunk, and x = 2 mean snow depth i n cm (r = .32, S = 52.12, n = 11). The e f f e c t i v e n e s s of reduced snow pack was d r a m a t i c a l l y demonstrated at the Limestone Creek survey s i t e . Average 173a Figure 6.2 The r e l a t i o n s h i p between snow depth and the di s t a n c e between bed s i t e s and t h e i r associated s h e l t e r t r e e s . 225 174 y = 142.4+ 100 X M E A N S N O W D E P T H (CM.) i 175 snow depths i n the openings between tr e e s was approximately 12 0 cm, but the bases of subalpine f i r s were exposed. Moose t r a v e l l e d i n s t r a i g h t l i n e paths between quamaniqs r a t h e r than the t y p i c a l meander. The only browse a v a i l a b l e was i n quamaniqs and most stems were h e a v i l y u t i l i z e d . As snow depths increased, moose chose s i t e s w i t h crown c l o s u r e s that were p r o g r e s s i v e l y more dense than the stands' average. In other words, snow depth and the d i f f e r e n t i a l between crown c l o s u r e of the bed s i t e s and that of the stand, were p o s i t i v e l y r e l a t e d (Figure 6.3). The r e l e v a n t r e g r e s s i o n equations are: f o r e s t s : y = 0.12 + 0.004x, n = 21, S = 0.21 2 y. x F = 5.62, r 2 = 0.23, p a r t i a l cutovers: y = -0.40 + O.Olx, n = 16, S = 0.17, F = 33.49, r 2 = 0.71, y. x ' where y = the d i f f e r e n c e i n crown canopy c l o s u r e as assessed p h o t o g r a p h i c a l l y and expressed as a p r o p o r t i o n , and x = snow depth i n cm. The higher r-squared value f o r p a r t i a l cutovers probably r e f l e c t s the greater opportunity f o r moose to s e l e c t s h e l t e r e d s i t e s . From t h i s equation, the type of \"micro=stands\" s e l e c t e d by moose from a l a r g e area of timber can be p r e d i c t e d . The s h i e l d i n g e f f e c t of c o n i f e r s on snow depth i s i l l u s t r a t e d i n Figure 6.4. Since increased snow pack i m p l i e s increased hindrance to movements, moose would be expected to behave i n 175a Figure 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 c l o s u r e between a bed s i t e and t h e . f o r e s t stand i n which i t was loc a t e d . 176a Figure 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. 177 178 ways that would minimize encountering adverse snow. As mentioned p r e v i o u s l y , moose at Limestone Creek kept almost e x c l u s i v e l y to quamaniqs. The use of quamaniqs f o r bed s i t e s a l s o v a r i e d across the snow gra d i e n t , although the p a t t e r n v a r i e d between h a b i t a t - t y p e s (Table 6.7). In the f o r e s t , three of the 11 bed s i t e s found i n shallow snow were i n quamaniqs. The comparable data were one of two i n moderate snow pack, and 11 of 11 i n deep snow packs. The corresponding proportions f o r p a r t i a l l y - l o g g e d stands were one of one, two of 14, and seven of e i g h t . The reason f o r t h i s departure from the f o r e s t p a t t e r n i s not known. For open types, one of the 3 0 beds i n shallow and moderate snow cl a s s e s were i n quamaniqs, wh i l e one of two was i n the deep snow. Table 6.7 Location of Beds i n Quamaniqs, as A f f e c t e d by Habitat and Snow Depth Class Snow depth P r o p o r t i o n of beds i n quamaniqs (s) c l a s s (cm) f o r e s t p a r t i a l cutover open T o t a l s shallow (0-40) 3/11 1/1 1/3 5/15 moderate (41-80) 1/2 2/14 0/27 3/43 deep (> 81) 11/11 7/8 1/2 19/21 T o t a l s 15/24 10/23 2/32 27/79 Moose bedding s i t e s d i f f e r e d from ambient snow depth (Table 6.8). D i f f e r e n c e s i n the compression r a t i o (snow depth i n a bed d i v i d e d by ambient snow depth (Des Meules 179 1965) v a r i e d between h a b i t a t - t y p e s . The r a t i o of 0.41 f o r p a r t i a l l y logged types was intermediate i n value, w h i l e the value f o r open h a b i t a t s was. highest at 0.49. This trend r e f l e c t e d the trend i n hardness from g r e a t e s t i n f o r e s t to l e a s t i n open s i t e s . Since most moose i n f o r e s t s bedded i n quamaniqs, undisturbed snow adjacent to bed s i t e s were l e s s than average f o r the study s i t e (Table 6.8). The r a t i o of adjacent: away depths followed a s i m i l a r p a t t e r n to compression r a t i o s , t h a t i s , l e a s t i n f o r e s t s and g r e a t e s t i n open s i t e s (Table 6.8). Table 6.8 Comparison of Snow Depths Between Moose Bedding S i t e s and Adjacent Areas Ha b i t a t - Snow compression r a t i o s \"Away\" snow type (n) bed/away adjacent/away depth (cm) Forest (20) 0.35 0.67 45 P a r t i a l cutover (13) 0.41 0.71 79 Open (11) 0.49 0.86 43 Means (44) 0.40 0.72 55 6.4 D i s c u s s i o n No attempt was made to determine number of beds made per day but t h i s has been studied by other workers. During January and February, Quebec moose e s t a b l i s h e d 5.0 beds per day per cows and c a l v e s ; b u l l s were not studied (Des Meules 1968). Although Des Meules suggested a d i f f e r e n c e i n d a i l y 180 bedding r a t e s between cows followed by calves and unaccompanied cows, I found no s i g n i f i c a n t d i f f e r e n c e (P > 0.05) i n a s t a t i s t i c a l a n a l y s i s of h i s data. In Russia, Timofeeva (1965 and 1967, quoted by Coady 1974) found t h a t i n e a r l y winter moose rested an average of f i v e times d a i l y (50-60 cm snow), and. i n l a t e winter (January to March) an average of e i g h t times d a i l y (> 7 0 cm snow). In B r i t i s h Columbia, G e i s t ' s (1963) r e s u l t s suggest that moose were l e s s a c t i v e i n winter than summer, and t h e r e f o r e might bed l e s s f r e q u e n t l y but f o r longer periods. Most r e c e n t l y , Franzmann et a l . (197 6) estimated that. Alaskan b u l l moose bed 5.5 times per day, while cow moose bed 5v4 times per day (ranges 4-7). A l l these above s t u d i e s i n d i c a t e t h a t moose bed 5-6 times d a i l y . Whether or not there are a c t u a l d i f f e r e n c e s i n bedding r a t e s must await more d e t a i l e d study. Physiographic features of moose beds are aspect, slope p o s i t i o n , slope and elevation.. I d i d not record e l e v a t i o n since i t was unimportant i n the drumlinized t i l l p l a i n and l e v e l l a c u s t r i n e basins where most of the bed surveys were conducted. The aspect of bed s i t e s was d i f f e r e n t i n Nova S c o t i a (Prescott 1968) and Quebec (Brassard et a l . 1974). The n u l l hypothesis of no d i f f e r -ence between provinces i n proportions of bed s i t e s i n each aspect c l a s s was r e j e c t e d ( x 2 = 22.24 and 33.52, r e s p e c t i v e l y , df = 7). Aspects of Nova Scotian and 181 Quebecois beds, however, were s i m i l a r ( x 2 = 8.04, 7 d f ) . Thus eastern Canadian moose chose predominantly southern aspects f o r beds (SW, S, SE), w h i l e n o r t h - c e n t r a l B r i t i s h Columbian moose chose e a s t e r l y , northwesterly, and southerly aspects (Table 6.4B). These geographic d i f f e r e n c e s probably r e f l e c t d i f f e r e n c e s i n c l i m a t i c features such as p r e v a i l i n g winds. S h e l t e r t r e e s have been studied by other moose researchers. Nova Scotian moose beds were lo c a t e d an average of 157 cm from s h e l t e r t r e e s whose mean DBH was 23 cm (Prescott 1968). Moose beds (67%) i n Nova S c o t i a were near balsam f i r (Abies balsamea). Des Meules (1965) presented s i m i l a r r e s u l t s f o r moose i n Laurentide Park, Quebec, although he d e f i n e s s h e l t e r t r e e s as those near moose t r a c k s (and beds?). S h e l t e r trees were u s u a l l y balsam f i r , the predominant species, averaging 18 cm DBH. This compares wi t h 92 and 29 cm f o r my d a t a , . r e s p e c t i v e l y . Peek et a l . (1976) a l s o documented the importance of t h i s t r e e species f o r bedding s i t e s of moose i n northeastern Minnesota. Des Meules (1964) a l s o described changes i n mean diameter of s h e l t e r t r e e s and d i s t a n c e to these trees as snow depths increased (his Figures 45 and 46). At 30 cm of snow, the nearest coniferous stem diameter averaged 5 cm, while at 90 cm of snow, mean diameter rose to approximately 28 cm. Data f o r P r i n c e George were 21 cm and 34 cm, r e s p e c t i v e l y . I t appears that B.C. moose sel e c t e d l a r g e r 182 diameter t r e e s than Quebec moose. T h i s l i k e l y r e f l e c t e d d i f f e r e n c e s i n canopy f e a t u r e s of the s h e l t e r t r e e s p e c i e s and p o s s i b l y the g e n e r a l l y l a r g e r s i z e of B.C. t r e e s . D i s t a n c e s to the s h e l t e r t r e e f o r the same snow depth (30 cm and 9 0 cm) were approximately 210 cm and 65 cm, r e s p e c t i v e l y , i n Quebec compared wi t h 112 cm and 52 cm f o r my data. Again d i s t a n c e s were l e s s than i n Quebec, suggesting t h a t white spruce and s u b a l p i n e f i r d i d not p r o t e c t moose as w e l l as balsam f i r . Both P r e s c o t t (1968) and Des Meules (1965) r e f e r r e d to beds l o c a t e d i n quamaniqs. . Des Meules (1965) documented a s h i f t i n bed l o c a t i o n from open cover-types to quamaniqs o f f e r e d by l a r g e c o n i f e r s once snow reached approximately 76 cm. The work by Peek e t a l . (1976) a l s o i m p l i e d the importance of quamaniqs i n r e l a t i o n to a s h i f t from open h a b i t a t s to c l o s e d canopied f o r e s t stands. In other s t u d i e s , d i f f e r e n c e s between bed s i t e s and the macro-environment f o r other c l i m a t i c parameters support and extend my r e s u l t s . Des Meules (19 65) documented t h a t wind speed a t 19 bed s i t e s was 25 percent of ambient v e l o c i t i e s , a s i g n i f i c a n t d i f f e r e n c e at P < 0.01. Based on a s m a l l e r sample of 4, my data suggested 34 percent. Des Meules (1965) a l s o demonstrated a temperature d i f f e r e n t i a l of 2\u00C2\u00B0C between bed s i t e s and ambient (n = 19). He a l s o p o i n t e d out t h a t temperature i n the snow p r o f i l e can be c o n s i d e r a b l y higher than i n the atmosphere. Based on ten 183 samples, when surface snow temperature averaged -12.3\u00C2\u00B0C, temperature at 51 cm below the surface was -3.2\u00C2\u00B0C. In a r e l a t e d study on w h i t e t a i l deer, Ozoga (1968) documented s i g n i f i c a n t c l i m a t i c d i f f e r e n c e s between deer yards and the surrounding area. Thus s e l e c t i o n of bed s i t e s can reduce energy d r a i n s from c l i m a t i c i n f l u e n c e s . Exposure to wind can be reduced s i g n i f i c a n t l y , thereby c u t t i n g c o n v e c t i o n a l l o s s e s . The temperature d i f f e r e n t i a l can a l s o be decreased to spare conductive l o s s e s . R a d i a t i o n a l c o o l i n g can be minimized by s h e l t e r i n g under coniferous cover (Moen 1968). Energy can a l s o be conserved by s e l e c t i n g warmer aspects and mid-slope p o s i t i o n t h a t escape c o o l a i r i n swales and exposure on ridgetops. Moose d i d not e x p l o i t these t a c t i c s throughout a winter season. Rather, i t appeared that the options were used as the winter environment became harsher and when moose were presumably i n poorer c o n d i t i o n . Gasaway and Coady (197 4) b e l i e v e that these behavioural adaptations of moose are e f f i c i e n t enough to minimize thermogenesis. These authors quoted work by Markgren (1966) f o r moose, Moen (1968) f o r w h i t e t a i l deer and Hart et a l . (1961) f o r barren ground caribou i n support of t h e i r statement. However, c o n t r o l l e d s t u d i e s are needed to confirm t h i s . 7. SECONDARY SUCCESSION IN SUB-BOREAL FORESTS 7.1 I n t r o d u c t i o n For moose, change due to f i r e and logging i s the most s i g n i f i c a n t feature of sub-boreal f o r e s t s . As the f o r e s t develops a f t e r disturbance, i t s c a p a b i l i t y to provide food and s h e l t e r f o r moose v a r i e s considerably. E a r l y stages supply a superabundance of food but l i t t l e cover, while l a t t e r stages supply s u f f i c i e n t cover but much reduced forage. Since c u r r e n t changes i n sub-boreal f o r e s t s occur on already developed ecosystems, the main focus f o r moose i s on secondary succession. This i s the non-phenological, d i r e c t i o n a l change i n vegetation that occurs i n already e s t a b l i s h e d ecosystems (Mueller-Dombois and E l l e n b e r g 1974). With minor exceptions, vegetational\" changes i n f o r e s t s are i n i t i a t e d by f i r e , l o g g i n g , l a n d - c l e a r i n g and disease and i n s e c t s . As secondary succession o r i g i n a t e s from only p a r t i a l disturbances of an ecosystem i t i n v o l v e s a l e s s complete sequence of l i f e forms than primary succession. I t proceeds r e l a t i v e l y q u i c k l y and i s considered to be f r e e from e v o l u t i o n a r y changes i n e c o l o g i c a l p r o p e r t i e s of the species i n v o l v e d (Mueller-Dombois and E l l e n b e r g 1974). Probably only the l i g h t - i n t o l e r a n t species of moss disappear 184 185 completely i n secondary succession (Mueller-Dombois 1965), but other, s h a d e - i n t o l e r a n t species show dramatic changes. The sequence of communities developing over time i s termed a sere. The sere i s subdivided i n t o component s e r a i stages or developmental stages. Stages are u s u a l l y d i s t i n g u i s h a b l e by changes i n dominant l i f e forms, height, biomass and v e r t i c a l s t r a t i f i c a t i o n . For secondary succession i n bor e a l coniferous f o r e s t s , the four major s e r a i stages are commonly defined as f o l l o w s (Cooper 1913): 1) Herb: vegetation dominated by annual, biannual and p e r e n n i a l graminoids and for b s ; u s u a l l y a s i n g l e l a y e r . 2) Shrub: ve g e t a t i o n dominated by deciduous woody veg e t a t i o n ; u s u a l l y two l a y e r s , the upper, woody, and the lower herbaceous. 3) E a r l y (Pioneer) f o r e s t : v e g e t a t i o n dominated by immature c o n i f e r s ; a t h i r d l a y e r added - immature tr e e s and t a l l shrubs. 4) Mature f o r e s t : v e getation dominated by mature c o n i f e r s w i t h three vegetation l a y e r s w e l l developed;^epiphytes present. The term \"dominated\" i s used i n the sense of c o n t r i b u t i n g the most to the communities' phytomass. The v e r t i c a l s t r u c t u r e of the vegetation was described i n terms of the f o l l o w i n g l a y e r s or s t r a t a (adapted from Revel 1972): A) the t r e e l a y e r - a l l t r e e s t a l l e r than 6 m. 186 B) the shrub l a y e r - a l l woody p l a n t s from 0.45 to 6 m t a l l . C) the herb l a y e r - woody p l a n t s shorter than 45 cm, and a l l herbs, f e r n s , h o r s e t a i l s and clubmosses. Throughout t h i s s e c t i o n , these l a y e r s are u s u a l l y r e f e r r e d to by t h e i r a l p h a b e t i c name to minimize confusion between the s i m i l a r names used f o r l a y e r s and su c c e s s i o n a l stages. Previous p l a n t e c o l o g i c a l s t u d i e s of the sub-boreal spruce zone have emphasized mature f o r e s t s , both climax and near-climax. Probably the e a r l i e s t work, was conducted by Ku j u l a (1945, quoted i n Revel 1972). I l l i n g w o r t h and A r l i d g e (1960) defined 12 s i t e types i n lodgepole pine and white spruce-subalpine f i r f o r e s t s . These were character-i z e d by common or d i a g n o s t i c understory p l a n t species and tree growth. Subsequently, Wali (19 69) explored v e g e t a t i o n -environment i n t e r r e l a t i o n s h i p s . His co-worker Revel (19 72) pursued a s y n e c o l o g i c a l study i n which he devised an e c o l o g i c a l c l a s s i f i c a t i o n of t h i s ecosystem based on methods developed by K r a j i n a (1959, 1965). Revel (1972) r e f e r r e d to stu d i e s on bo r e a l v e g e t a t i o n by Moss (1953a, 1953b, 1955) as p a r t i c u l a r l y r e l e v a n t to the sub-boreal spruce biogeo-c l i m a t i c zone. To t h i s l i s t should a l s o be added the newly completed work by Annas (19 77) on the bor e a l black and white spruce b i o g e o c l i m a t i c zone. Extensive f i e l d work by J . van Barneveld and h i s a s s o c i a t e s , though l a r g e l y 187 unpublished, has c o n t r i b u t e d s u b s t a n t i a l l y to understanding the i n t e r r e l a t i o n s h i p s between environment and vegetation and the general patterns of succession. However, these foregoing s t u d i e s d e a l t p r i m a r i l y w i t h f o r e s t c l a s s i f i c a -t i o n and p e r i p h e r a l l y w i t h succession. E a r l y s u c c e s s i o n a l development has been v i r t u a l l y unreported. Given the dearth of inform a t i o n on secondary p l a n t succession i n sub-boreal f o r e s t s , and r e a l i z i n g the importance of succession to moose, the f o l l o w i n g o b j e c t i v e s were set f o r major s e r a i stages: 1) To describe the p l a n t species composition and abundance ( f l o r i s t i c s ) . 2) To q u a n t i f y the above-ground phytomass of the understory v e g e t a t i o n , by forage c l a s s e s . 3) To estimate the above-ground net p r o d u c t i v i t y of the understory v e g e t a t i o n , by forage c l a s s e s . 4) To describe changes i n the height of the Shrub l a y e r . 5) To compare the changes i n the height and mass of browse i n the Shrub l a y e r . 6) To describe trends i n the production of food and cover. 7) To assess the s i g n i f i c a n c e of major environmental f a c t o r s on the patterns and r a t e s of succession. These o b j e c t i v e s were a p p l i e d mainly on the Deserters environmental u n i t since i t occupied the l a r g e s t area w i t h i n the study area. Other u n i t s were t r e a t e d as time allowed. 188 7.2 Methods 7.2.1 S t r a t i f i c a t i o n Successional trends were studi e d by using side-by-side comparisons (Mueller-Dombois and E l l e n b e r g 1974). Although t h i s method i s l e s s s a t i s f a c t o r y than s t u d i e s on the same area over a time p e r i o d , i t i s the only one f e a s i b l e f o r short-term s t u d i e s . The major f a c t o r s i n f l u e n c i n g s u c c e s s i o n a l patterns were time-dependent and site-dependent. The time f a c t o r was of prime importance because the value of a p l a n t community as food- or cover-producing h a b i t a t v a r i e d according to the time since logging or w i l d f i r e . For i n v e s t i g a t i o n s of chronosequences, the main, approach was to hold a l l other e c o l o g i c a l ( s i t e ) f a c t o r s as constant as p o s s i b l e , and examine stands of d i f f e r e n t ages. Of. l e s s e r importance were s i t e - r e l a t e d f a c t o r s . Analogous to the need f o r c o n t r o l of s i t e f a c t o r s i n studying chronosequences, i s the need to hold time and non-ta r g e t s i t e f a c t o r s constant while examining f a c t o r s of i n t e r e s t . Of the many s i t e - r e l a t e d f a c t o r s t h a t could be stu d i e d , I s e l e c t e d the broad p h y s i c a l environment (climate-substrate) as the p r i n c i p a l one f o r study. Based on f a m i l i a r i t y w i t h the study area and w i t h r e l e v a n t e c o l o g i c a l s t u d i e s , t h i s f a c t o r was judged to be of f i r s t concern. The non-target s i t e f a c t o r s were p a r t l y c o n t r o l l e d . 189 Slope and aspect were c o n t r o l l e d by having as f a r as p o s s i b l e n i l slope and n i l aspect. Cause of disturbance was not completely c o n t r o l l e d since c l e a r c u t s o l d e r than approximately ten years were not a v a i l a b l e f o r study. Thus I used w i l d f i r e s i t e s as samples f o r o l d e r stages of the sere. A l s o , the v a r i a t i o n r e s u l t i n g from type and time of f i r e , and \" q u a l i t y \" of logging was not c o n t r o l l e d . These were considered important v a r i a b l e s that should be addressed i n a subsequent study. S t r a t i f i c a t i o n was the i n i t i a l step i n s e l e c t i n g p o t e n t i a l study s i t e s . The b a s i s of s t r a t i f y i n g was the framework of environmental u n i t s described i n Section 2.1. From t h i s framework, e f f o r t was.concentrated on mesic environments over t i l l and l a c u s t r i n e s u b s t r a t e s . (Reasons f o r t h i s choice were given i n Section 2.1.) General observations were made f o r the other u n i t s as time permitted, e s p e c i a l l y the r i p a r i a n u n i t s . P r e l i m i n a r y s i t e s e l e c t i o n followed s t r a t i f i c a t i o n . F orest cover-type maps (1:15840) were s t u d i e d to s e l e c t t e n t a t i v e s i t e s that represented a wide range i n stand age and i n composition of tre e species. Subsequently, a e r i a l photographs were examined to determine a c c e s s i b i l i t y and to check t h a t map data were s t i l l a p p l i c a b l e . S i t e s were f i n a l l y chosen a f t e r they were checked i n the f i e l d to v e r i f y the c l a s s i f i c a t i o n of environmental u n i t and cover-type d e s c r i p t i o n s . Approximately 75 f i e l d checks were made 190 before s e l e c t i n g the f i n a l s i t e s . 7.2.2 F i e l d Sampling Procedures Two types of f i e l d procedures were used: d e t a i l e d and synoptic. The f i r s t was to determine approximate above-ground net understory p l a n t p r o d u c t i v i t y and biomass i n the four c l a s s i c a l s u c c e s s i o n a l stages: herb, shrub, pioneer f o r e s t and mature f o r e s t (Odum 1971). For t h i s d e t a i l e d study, vegetation sampling began i n J u l y 1972 a f t e r maximum p l a n t development was a t t a i n e d . Only the t i l l s ubstrate was examined. As the s i t e s were on drumlinized t i l l , I decided to sample only c r e s t s of drumlin to minimize confounding due to slope and aspect. Four age c l a s s e s were sampled i n d u p l i c a t e : a) a one year o l d c l e a r cut that was burned i n September 1971, b) an 11 year o l d s e l e c t i v e l y logged s i t e t h a t was subsequently burned by w i l d l i f e i n August 1961, c) a 40 year o l d w i l d f i r e , and d) a 195 year o l d w i l d f i r e . At each sampling s i t e b a s i c s i t e features were noted, e.g., slope, aspect, microtopography, moose use. A l l t rees greater than 2.5 cm diameter at breast height (DBH = 1.4 m) w i t h i n a 20- x 20-m p l o t were t a l l i e d by species, diameter c l a s s and c o n d i t i o n ( a l i v e or dead). Heights and increment borings were taken from f i v e t rees i n 191 each p l o t . In the centre of t h i s p l o t , a 5- x 5-m perquadrat was l a i d out, and ten 2- x 0.5-m quadrats were located w i t h i n the perquadrat. A l l f o r b s , shrubs and grasses were c l i p p e d at ground l e v e l from these quadrats, and bagged separately by species. Shrubs remaining i n the perquadrat were then c l i p p e d , a l s o at ground l e v e l . In the l a b o r a t o r y , shrubs were separated i n t o the f o l l o w i n g components: leaves, annual (1972) twigs, o l d e r twigs. For some species such as white spruce, bearberry (Aretostaphylos uva-ursi) and P r i n c e ' s pine, complete separations were not p o s s i b l e . A l l separated shrub mate r i a l , forbs and grasses were weighed a f t e r d r y i n g f o r at l e a s t 24 h at 50\u00C2\u00B0C. The second type of f i e l d procedure, synoptic sampling, was used to extend the geographic area of coverage and to meet other o b j e c t i v e s of t h i s s e c t i o n . The above d e t a i l e d methods were r e v i s e d and s i m p l i f i e d to enable sampling a wider range of stands. With t h i s method, f i v e s t a t i o n s were sampled at each s i t e . A l l s i t e s were s i t u a t e d from 1 to 1.5 times the adjacent stand height away from an ecotone (e.g., cut boundary). On l e v e l t e r r a i n , such as on some l a c u s t r i n e s u b s t r a t e s , s t a t i o n s were u s u a l l y spaced 30.5 m apart. On g l a c i a l t i l l , the drumlin was used as the b a s i c sampling u n i t . S t a t i o n s were placed perpendicular to the drumlin's long a x i s at or near where the drumlin was widest (Figure 7.1). Spacing of the 191a Figure 7.1 The s i t e and s t a t i o n layout used to study secondary p l a n t succession. A. S I T E L A Y O U T 192 I LONGITUDINAL AXIS WIDEST PART OF DRUMLIN OF DRUMLIN B. D E T A I L S O F S A M P L E S T A T I O N L A Y O U T M O S S - L I T T E R S A M P L E 2 0 x 5 0 C M m M mi i I HERBS & SHORT SHRUB PLOTS 1 x 1 M. TALL SHRUB PLOT 2 x 2 M . -rt-QUADRATS FOR FLORISTICS 20 x 50 C M . , 1.5 M. APART 'STATION CENTER (CENTER OF WEDGE PRISM PLOT) 193 s t a t i o n s was e q u i d i s t a n t , w i t h a c t u a l distances dependent upon the width of the drumlin. Each t r a n s e c t began and f i n i s h e d j u s t above the e a s i l y defined swale or r e c e i v i n g area between drumlins. This type.of spacing was done to maximize drumlin-based v a r i a t i o n w i t h i n s i t e s and thus minimize i t s between s i t e s e f f e c t . For each s i t e , the f o l l o w i n g general i n f o r m a t i o n was recorded on a s i t e sheet: a) l o c a t i o n of s i t e on 1:15,000 cover-type maps, 1:15,000 a e r i a l photographs, and by f o r e s t management u n i t , b) e l e v a t i o n , topography and parent m a t e r i a l , c) s i t e photograph d) general notes on bearing and spacing of s t a t i o n s , type of s i t e , evidence of moose a c t i v i t y both summer and wi n t e r , and s p e c i f i c l o c a t i o n of s i t e . At each s t a t i o n , data were recorded f o r understory f l o r i s t i c s , understory phytomass and s e v e r a l features of the f o r e s t , v i z . , species composition, c o n d i t i o n , b a s a l area, crown c l o s u r e , height and age. Slope and aspect were determined by using a compass and clinometer. P l o t lay-out i s i l l u s t r a t e d i n Figure 7.1. The information used to describe f l o r a l changes i n secondary succession was p l a n t species composition and abundance. The sampling method and type of data used f o r f l o r i s t i c s d i f f e r e d f o r each of the three vegetation l a y e r s A, B and C. For the \"A\" l a y e r , species composition was based on the species recorded i n the wedge p r i s o n sampling and abundance was the percentage composition of the tre e stems i n these samples. S i m i l a r l y f o r the \"B\" l a y e r , composition and abundance were based on stem counts i n quadrats (see second paragraph below). F l o r i s t i c s of the C l a y e r were recorded using the Daubenmire (1959) method. In each of ten 20- x 50-cm quadrats, the canopy-coverage of v a s c u l a r p l a n t species (excluding i n f l o r e s c e n c e ) was rated s u b j e c t i v e l y on a 1-6 scale (Table 7.1). Canopy-coverage was assessed by es t i m a t i n g the p r o p o r t i o n of a quadrat occupied by the t o t a l area of v e r t i c a l l y p r o j e c t e d (imaginary) polygons t h a t enclosed perimeters of p l a n t s of each species. Coverage was recorded i f a p l a n t occurred i n the quadrat, even i f i t was rooted outside the frame. Quadrats were spaced 1.5 m apart along a t r a n s e c t that was at r i g h t angles to the l i n e of s t a t i o n s i n l e v e l t e r r a i n or that was a l i g n e d p a r a l l e l to the contour i n r o l l i n g t e r r a i n . This sampling provided estimates of frequency of occurrence and cover f o r ^ vegetation below 1.4 m. Vegetation t a l l e r than t h i s , almost e n t i r e l y shrubs and t r e e s , was not in c l u d e d due to the d i f f i c u l t y of a c c u r a t e l y e s t i m a t i n g canopy-coverage. Those species missed i n sampling but at the s i t e were a l s o noted. Understory biomass was estimated by a combination of two methods. For the t a l l shrub l a y e r , woody p l a n t s t a l l e r 195 Table 7.1 Scale Used to Assess Canopy-Coverage of Understory Vegetation (C Layer) ( a f t e r Daubenmire 1959), plus Domin Scale Equivalents Approximate Canopy-coverage Range of Mid-point eq u i v a l e n t i n r a t i n g coverage of c l a s s Domin s c a l e * 1 0.1 - 5. 0 2.5 +, 1, 2, 3 2 5.1 - 25. 0 15.0 4, 5, 6 3 25.1 - 50.0 37.5 7 4 50.1 - 75. 0 62.5 8 5 75.1 - 95.0 85. 0 9 6 95.1 - 100.0 97.5 10 *To a i d comparisons w i t h R e v e l 1 s (1972) data. than 45 cm, a 2 - x 2-m perquadrat was l a i d out at the samp-l i n g s t a t i o n . Diameters at 10 cm of a l l shrubs > exceeding 45 cm were recorded by species. Height and oven-dried weights were p r e d i c t e d from these diameters using r e g r e s s i o n equations (described i n s e c t i o n 7.2.3). For the \"C\" or herb l a y e r , two 1- 2-m quadrats were randomly chosen from the four p o s s i b l e i n the perquadrat. From these two quadrats a l l shrubs l e s s than 45 cm t a l l , and other vegetation was c l i p p e d to ground l e v e l and bagged according to the f o l l o w i n g forage c l a s s e s : a) forbs b) graminoids c) ferns d) h o r s e t a i l s e) clubmosses 196 f) shrubs g) coniferous s e e d l i n g s . These samples were weighed a f t e r d r y i n g f o r 24 h at 50\u00C2\u00B0C. Trees were sampled using the B i t t e r l i c h v a r i a b l e p l o t method as described by D i l w o r t h and B e l l (1971). Basal area was determined using a wedge prism held v e r t i c a l l y over the s t a t i o n center. A basal area f a c t o r (BAF) of 20 was se l e c t e d a f t e r c o n s u l t a t i o n w i t h f o r e s t e r s p r a c t i c i n g i n the Pri n c e George area. Approximately 10 trees were sampled at each s t a t i o n using t h i s BAF. The species, c o n d i t i o n (dead or a l i v e ) , and DBH of each \" i n \" t r e e was recorded, t a k i n g precautions mentioned by the above authors. In a d d i t i o n , height was determined t r i g o n o m e t r i c a l l y f o r one dominant or co-dominant t r e e per s t a t i o n , f o r a t o t a l of f i v e per s i t e . Ages of these trees were determined by counting annual r i n g s i n increments e x t r a c t e d at 1.4 m above ground l e v e l . Estimated ages were c o r r e c t e d according to species and ' s i t e c l a s s (Forest Club 1971). A l s o , at each s t a t i o n , the crown-cl o s u r e was recorded p h o t o g r a p h i c a l l y w i t h a fi s h e y e lens (17 mm) (Brown and Worley 1965). Closure was estimated by counting \" h i t s \" or canopy presence on a d o t - g r i d ( f i v e 2 2 dots/cm ) f o r the center 25 cm of each p r i n t . A summary of parameters sampled, methods used and sampling i n t e n s i t y f o r the synoptic surveys i s presented i n Table 7.2. 197 Table 7.2 Summary of Features Sampled i n the Synoptic Study of Succession Sampling i n t e n s i t y Parameters sampled Method used u n i t s / p l o t p l o t s / s i t e TREES (\"A\" l a y e r ) : spp. composition stem count (> 4.0 cm) 1 point 5 bas a l area wedge prism (BAF 20) 1 point 5 diameter diameter tape (DBH) 0-15 trees 5 age increment borer 1-2 trees 5 height clinometer and tape 1-2 trees 5 c o n d i t i o n observation a l l t rees 5 crown c l o s u r e photographic ( f i s h e y e lens) 1 photograph 5 TALL SHRUBS (\"B\" l a y e r ) : spp. composition d i r e c t count 0 o 2 one z- x z-m quadrat 5 bas a l area c a l c u l a t e d from diameter as above 5 mass p r e d i c t i o n .from diameter as above 5 height p r e d i c t i o n from diameter as above 5 dens i t y d i r e c t count as above 5 diameter micrometer c a l i p e r s as above 5 HERBS, SMALL SHRUBS (\"C\" l a y e r ) : spp. composition observation 10 quadrats 5 occurrence presence i n quadrats as above 5 canopy-coverage 1-6 s c a l e i n quadrats as above 5 phytomass c l i p p i n g to ground l e v e l two 1- x 1-m quadrats 5 SITE FEATURES: substrate s o i l maps, p i t s 1 topography observation 1 aspect compass and clinometer 1 observation 5 slope compass and clinometer as above 5 e l e v a t i o n map, a l t i m e t e r 1 h i s t o r y cover maps, observation, et c . 1 198 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 Mass and height were estimated from morphological measurements, using r e g r e s s i o n a n a l y s i s . As Newbould (1967:10) s t a t e d , the object was \"to o b t a i n c o r r e l a t i o n s between a comparatively small d e s t r u c t i v e sampling (which i s both time consuming and d e s t r u c t i v e of the h a b i t a t ) w i t h a l a r g e r non-destructive sample which i s r e p r e s e n t a t i v e of the stand. . .\" A l l o m e t r i c regressions of weight on DBH have been widely and s u c c e s s f u l l y a p p l i e d i n f o r e s t r y since at l e a s t 1944 (Kittredge 1944). The 1973 IUFRO symposium on the mensuration of f o r e s t biomass provided a u s e f u l recent summary of t h i s subject i n f o r e s t r y (see a l s o Dunn 1974). In w i l d l i f e ecology, the main a p p l i c a t i o n of dimensional a n a l y s i s has been to estimate production and u t i l i z a t i o n of browse, u s u a l l y considered only as annual twig growth. These estimates have been commonly based on diameter or length measurements ( B a s i l e and Hutchings 1966, H a l l s and Harlow 1971, Lyon 1970, Schuster 1967, Stickney 1966, T e l f e r 1969), or l e s s commonly on crown volume (Lyon 1968, Quenet 1971), canopy area and other plant-form measurements (Peek 1970). Studies on p r e d i c t i n g t o t a l above-ground biomass f o r browse species are r e l a t i v e l y uncommon. T e l f e r (19 69) presented data based on b a s a l diameter measurements f o r 2 2 eastern North American shrub species. Some f o r e s t r y and production ecology s t u d i e s such as those on puckerbrush (Young 19 71), heath species 199 (Whittaker 1962), aspen ( B e l l a 1968), b i r c h (Gregory and Haak 1965), Douglas f i r ( C r o s s l e y 1967, Kurucz 1969), and red a l d e r (Minus rubra) (Smith 1974b) are a l s o a p p l i c a b l e . More r e c e n t l y , Brown (1976a) p r e d i c t e d mass f o r 22 Rocky Mountain shrub s p e c i e s , and crown weights f o r 11 c o n i f e r s (Brown 1976b). They were s e l e c t e d on the b a s i s of t h e i r commonness i n both s u c c e s s i o n a l and mature sub-boreal f o r e s t stands w i t h i n a 60 km r a d i u s of P r i n c e George. Between 15 and 4 8 samples were c o l l e c t e d f o r each s p e c i e s f o l l o w i n g p o i n t s suggested by Demaerschalk and Kozak (19 74). Each sample was c l a s s i f i e d a c c o r d i n g to the f o l l o w i n g four v e g e t a t i o n - parent m a t e r i a l c l a s s e s : u n f o r e s t e d on t i l l , f o r e s t e d on t i l l , u n f o r e s t e d on l a c u s t r i n e and f o r e s t e d on l a c u s t r i n e , to enable t e s t i n g f o r p o s s i b l e s i t e - r e l a t e d d i f f e r e n c e s w i t h i n a s p e c i e s as found by Peek e t a l . (1971). Only unbrowsed or l i g h t l y browsed p l a n t s were taken. The upper l i m i t s of diameters sampled f o r each s p e c i e s were based on f i e l d o b s e r v a t i o n s of the maximum s i z e encountered. A l l p l a n t s were c o l l e c t e d from J u l y - September 1973. Each sample was measured f o r t o t a l h e i g h t (H), l i v i n g crown depth (CD), stem diameter at palm width (DHW), stem diameter at the lowest l i v i n g branch (DLB), and dry weight (DW). Lengths and diameters were measured to the ne a r e s t cm and 0.01 cm, r e s p e c t i v e l y . The DHW was taken at 10 cm above the base to m i n i m i z e . v a r i a t i o n due to v a r i a b l e ' s w e l l i n g at the r o o t c o l l a r . T h i s i s analogous to the 200 measurement of DBH f o r t r e e s . The DLB measurement was suggested by Newbould (1967:18). Weights were taken a f t e r drying f o r 4 8 h at 5 0\u00C2\u00B0C. Several independent v a r i a b l e s (and t h e i r n a t u r a l l o g a r i t h m i c transformations) were evaluated as p r e d i c t o r s f o r the dependent v a r i a b l e s of t o t a l dry weight and height. This choice of p r e d i c t o r s was based on an examination of the p l o t t e d data, and on suggestions i n the l i t e r a t u r e . For dry weight, the f o l l o w i n g independent v a r i a b l e s were s e l e c t e d : DHW, DLB, (DHW)2, (DLB) 2, (DLB) 2H, and (DHW)2H. For height, the p r e d i c t o r s examined were DHW, DLB and t h e i r l o g a r i t h m i c transformations. In a l l cases, the c r i t e r i a f o r e v a l u a t i o n 2 were r , the c o e f f i c i e n t of determination, and S , the ' y .x standard e r r o r of the estimate. C a l c u l a t i o n s were performed using the s t a t i s t i c a l package, SPSS (Nie e t a l . 1970). S i t e - r e l a t e d d i f f e r e n c e s had no s i g n i f i c a n t (p > 0.05) e f f e c t on the slope of the r e g r e s s i o n equations, (b^ = b^) at l e a s t f o r the s p e c i e s , s i t e types and independent v a r i a b l e s that were compared (Table 7.3). This f i n d i n g d i f f e r s from Peek et a l . (1971) who found th a t 87 of 100 comparisons of s i t e - r e g r e s s i o n c o e f f i c i e n t s i n Minnesota d i f f e r e d s i g n i f i c a n t l y from each other (P < 0.01). Possibly, the d i f f e r e n c e s between the P r i n c e George s i t e s were not as great as those i n Minnesota. A l s o , Peek et a l . (1971) examined only current year's growth from p l a n t s t h a t r e c e i v e d v a r y i n g degrees of browsing, while f o r the present Table 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 Selected Shrub Species Regression of Sample S i t e types l o g DW w i t h s i z e s Species examined compared* independent var. Ca l c u l a t e d t * * ( s i t e t y p e ) * Abies lasiocarpa (subalpine f i r ) 1 vs. 4 log DLB 0.905 12(1) log (DLB) 2H 0.584 18(4) Amelanchier alnifolia (Saskatoon) 1 vs. 4 lo g (DHW)2H 1.113 23(1) log (DLB) 2H 0.404 17(4) Loniaera involucrata (black twinberry) 1 vs. 4 log DHW 0.981 ' 15(1) log (DHW)2H 0.655 23(4) Rosa spp. (rose) 1 vs. 4 log (DLB) 2H 0.374 21(1) log (DHW)2H 0.497 8(4) Sorbus spp. (mountain ash) 2 vs. 4 log (DLB) 2H 1.711 18(2) l o g (DHW)2H 2.455 12(4) Spiraea lueida (spirea) 1 vs. 4 same as above 0.403 14(1) - 0.693 15(1) Viburnum edule (squashberry) 1 vs. 4 same as above 1.705 16(1) 2.052 26(4) Populus tremuloides (trembling aspen) 1 vs. 3 lo g (DLB) 2H 0.270 10(1) 1 vs. 4 0.208 11(3) 3 vs. 4 0.062 9(4) 1 vs. 3 lo g (DHW)2H 1.540 1 vs. 4 0.857 3 vs. 4 1.510 * S i t e types: 1 = unforested on t i l l , 2 = f o r e s t e d on t i l l , 3 = unforestec on l a c u s t r i n e , 4 = fo r e s t e d on l a c u s t r i n e . **A11 c a l c u l a t e d \" t \" ' s were not s i g n i f i c a n t at the 95 percent l e v e l . 202 study e n t i r e p l a n t s were sampled that were at most, only l i g h t l y browsed. As might be expected from such a d i v e r s e grouping of p l a n t s , no one estimator was c o n s i s t e n t l y the best p r e d i c t o r of phytomass (Table 7.4). Logarithmic transformations of 2 (DHW) H were best i n 10 i n s t a n c e s , the log transformations 2 2 of (DLB) H and the untransformed (DHW) H were best f o r each 2 of s i x species, while untransformed (DLB) H provided the best f i t f o r one species (Table 7.4). Logarithmic 2 2 transformations of both (DLB) H and (DHW) H provided a b e t t e r f i t of the data than diameter-only measurements f o r a l l species except saskatoon, and b e t t e r than the untransformed diameter x height combination f o r a l l species except saskatoon, b i r c h , black twinberry, rose, squashberry and s p i r e a . However, i t should be noted that f o r most species there i s very l i t t l e d i f f e r e n c e between the r-squared values of the d i f f e r e n t r e g r e s s i o n . Log DHW was a p r e f e r a b l e v a r i a b l e to log DLB, and to untransformed v a r i a b l e s f o r these diameters (Table 7.4). Except f o r subalpine f i r and saskatoon, equations based on 2 the former diameter u s u a l l y had the highest r values and the lowest standard e r r o r s of the estimate. The b e t t e r f i t of the DHW data may r e s u l t p a r t l y from the complex of f a c t o r s that determine when the lowest branches die and 2 break o f f . For a l l species except thimbleberry, r values exceeded 0.90, w i t h 14 exceeding 0.96. Although the slope Table 7.4 C o e f f i c i e n t s of Determination (r Values) 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 C o e f f i c i e n t s of determination f o r v a r i a b l e s Species examined* log DHW log DLB DHW2H DLB ,2H log DHW2H log DLB 2H Abies lasiocarpa (subalpine f i r ) 0. 91 0. 95 0. 87 0. 87 0. 85 0. 98** Acer glabrum (Douglas f i r ) 0. 87 0. 82 0. 36 0. 33 0. 90 0. 87 Alnus spp. (alder) 0. 97 0. 94 0. 95 0. 96 0. 97 0. ,96 Amelanohier alnifolia (Saskatoon) 0. 76 0. 79 0. 91 0. 89 0. 75 0. ,78 Betula papyrifera (paper b i r c h ) 0. 90 0. 89 0. 96 0. 90 0. 95 0. ,97 Cornus stolonifera (red-osier dogwood) 0. 92 0. 60 0. 85 0. 31 0. 97 0. ,79 Lonioera involuorata (black twinberry) 0. 90 0. 82 0. 94 0. 92 0. 93 0. ,89 Picea glauoa (white spruce) 0. 95 0. 95 0. 40 0. 40 0. 97 0. ,96 Pinus oontorta (lodgepole pine) 0. 97 0. 95 0. 48 0. 47 0. 97 0. ,97 Populus tremuloides (trembling aspen) 0. 95 0. 91 0. ,79 0. 92 0. ,97 0. ,97 P. balsamifera (black cottonwood) 0. 97 0. 94 0. ,73 0. 63 0. 97 0. ,96 Rosa spp. (rose) 0. 87 0. 76 0. ,97 0. 97 0. ,90 0. ,87 Rubus idaeus (raspberry) 0. 94 0. 95 0. ,95 0. 96 0. ,98 0. ,99 R. parviflorus (thimbleberry) 0. 27 0. 31 0. ,13 0. 18 0. ,35 0. ,42 Salix spp. (willow) 0. 95 0. 92 0. ,93 0. 43 0. ,97 0. ,95 Sorbus spp. (mountain ash) 0. 93 0. 79 0. ,98 0. 83 0. ,98 0. ,93 Spiraea luoida (spirea) 0. 98 0. 93 0. ,89 0. 83 0. ,99 0. ,97 Vaooinium spp. (vaccinia) 0. 88 0. 85 0. ,97 0. 94 0. ,92 0. .92 Vibernum edule (squashberry) 0. 76 0. 71 0. .96 0. 89 0. ,83 0. .83 *Sample s i z e s i n Table 7.6. **Underline denotes highest r value f o r that species. o 204 of the r e g r e s s i o n f o r thimbleberry was s i g n i f i c a n t l y d i f f e r e n t from zero (P < 0.01), the best f i t accounted f o r only 42 percent of the observed v a r i a t i o n . The independent v a r i a b l e of DHW p r e d i c t e d p l a n t height reasonably w e l l (Table 7.5). A l l r e g r e s s i o n equations were h i g h l y s i g n i f i c a n t (P << 0.01). C o e f f i c i e n t s of determination ranged from 0.50 to 0.94, averaging 0.76. Standard e r r o r s of the estimate, expressed as percentages of \"x\", had a mean value of 11 percent w i t h a standard d e v i a t i o n of 4 percent. For the purposes of t h i s study, DHW was s e l e c t e d as the s i n g l e most u s e f u l independent v a r i a b l e f o r p r e d i c t i n g the mass and height of woody shrubs (Tables 7.5 and 7.6). For more d e t a i l e d s t u d i e s than mine, combinations i n v o l v i n g height and diameter would be necessary. However, i n a synoptic study aimed at d e l i n e a t i n g general trends, the a d d i t i o n a l time r e q u i r e d to measure height was considered i m p r a c t i c a l and unnecessary. Thus f o r height, r e g r e s s i o n equations derived from untransformed data were used; f o r weight, equations based on l o g a r i t h m i c a l l y transformed data were used. During vegetation sampling, f i v e species were encountered i n a d d i t i o n to the 19 f o r which r e g r e s s i o n equations had been derived. For these f i v e , e x i s t i n g r e g r e s s i o n c o e f f i c i e n t s f o r species of the same genus or growth form were used i n s t e a d . Thus f o r hardhack (Spiraea Table 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 Regression c o e f f i c i e n t s \u00E2\u0080\u009E r , . , \u00E2\u0080\u0094 7 ; ^ Range of hexghts Species a y.x. r (cm) Abies lasiooarpa (subalpine f i r ) 11. 08 51. 54 ( 4. 60) 0.79 17-166 Acer glabrum (Douglas maple) 31. 40 66. 62 ( 9. 67) 0.77 10-240 Alnus spp. (alder) 25. 42 104. 47 (10. 25) 0.81 60-291 Amelanchier alnifolia (Saskatoon) 36. 65 94. 90 (14. 67) 0.56 47-248 Betula papyrifera (paper b i r c h ) 11. 58 109. 49 ( 9. 95) 0.83 38-255 Cornus stolonifera ( r e d - o s i e r dogwood) -27. 81 164. 00 (14. 20) 0.90 14-189 Loniaera involucrata (black twinberry) 23. 26 77. 84 (12. 50) 0.50 9-137 Picea glauoa (white spruce) 12. 01 62. 69 ( 5. 48) 0.82 26-294 Pinus contorta (lodgepole pine) 15. 31 61. 33 (10. 95) 0.71 21-296 Populus tremuloides (trembling aspen) 36. 36 86. 82 ( 9. 07) 0.75 27-274 P. balsamifera (black cottonwood) 19. 95 62. 48 ( 3. 74) 0.94 23-294 Rosa spp. (rose) 8. 06 111. 89 (11. 57) 0.76 15-135 Rubus idaeus (raspberry) -19. 76 179. 17 (15. 60) 0.91 6-150 P. parviflorus (thimbleberry) -22. 55 133. 01 (23. 82) 0.63 12-73 Salix spp. (willow) 18. 23 123. 11 (10. 14) 0.84 9-202 Sorbus spp. (mountain ash) 24. 62 76. 14 (11. 25) 0.62 24-210 Spiraea lucida (spirea) 3. 81 147. 37 (15. 11) 0.75 14-115 Vaecinium spp. ( v a c c i n i a ) -0. 50 88. 74 ( 7. 54) 0.88 16-110 Vibernum edule (squashberry) 25. 18 84. 19 ( 9.. 12) 0.65 23-224 *Sample s i z e s i n Table 7.6. o Table 7.6 Species 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 Oven-Dried, Above-Ground Phytomass of 19 Sub-Boreal Shrubs from Diameter*, and from Diameter Squared by Length Measurements. A l l V a r i a b l e s Based on Logarithmic Transformed Data Regression c o e f f i c i e n t s + diameter ( d i a m e t e r ) 2 l e n g t h a b a b n Range of diameters (cm) Abies lasiocarpa (subalpine f i r ) 4. .00 2. ,33 0. ,63 0. 82 36 0. 24--3. 00 Acer gldbrum (Douglas maple) 3. .19 2. , 19** -0. ,26 0. 7 4 * * 16 0. 17--3. 82 Alnus spp. (alder) 3. .49 2. ,62** -0. ,93 0. 91** 26 0. 41- -2. 91 Amelanchier alnifolia (Saskatoon) 4. ,03 2, ,11 0. ,41 0. 74 35 0. 29--1. 72 Betula papyrifera (paper b i r c h ) 3. ,60 2. ,41** -0. ,14 0. 78 26 0. 30--2. 24 Cornus stolonifera ( r e d - o s i e r dogwood) 3. ,81 2. ,96** -0. ,39 0. 86** 16 0. 25--1. 12 Lonicera involuorata (black twinberry) 3. ,87 2. ,15** 0. ,55 0. 68** 41 0. 28--1. 60 Pioea glauoa (white spruce) 4. .10 2. ,20** 0. ,86 0. 76** 30 0. 35--4. 07 Pinus oontorta (lodgepole pine) 3. ,60 2. ,40** 0. ,13 0. 82** 15 0. 28--4. 44 Populus tremuloides (trembling aspen) 3. ,55 2. , 45** -0. ,44 0. 84** 33 0. 28--3. 23 P. balsamifera (black cottonwood) 3. ,64 2. , 28** 0. ,08 0. 80** 21 0. 30--4. 67 Rosa spp. (rose) 4. ,13 2. ,77** -0. ,25 0. 90** 31 0. 15--1. 22 Rubus idaeus (raspberry) 3. ,90 3. ,07 \u00E2\u0080\u00A2 -0. ,49 0. 84 15 0. 18--0. 85 R. parviflorus (thimbleberry) 2. ,55 1. ,34 0. ,68 0. 41 20 0. 21- -0. 70 Salix spp. (willow) 3. ,71 2. , 65** -0. ,58 0. 86** 30 0. 12--1. 70 Sorbus spp.. (mountain ash) 3. ,48 2. ,52** -0. ,49 0. 86** 30 0. 32--2. 58 Spiraea luoida (spirea) 3. ,58 2. ,23** -0. ,20 0. 76 34 0. 12--0. 68 Vaccinium spp. (v a c c i n i a ) 3. ,88 2. , 47** 0. ,18 0. 84** 21 0. 15--1. 14 Vibernum edule (squashberry) 3. ,75 2. ,75** -0. ,75 0. 95 48 0. 33- -2. 53 *Diameter r e f e r s to DHW i n a l l cases. **Best equation of the two p o s s i b i l i t i e s (DHW and DLB). Best of the s i x equations te s t e d . K> + 2 \u00C2\u00B0 General form of the re g r e s s i o n equation i s y = a+b In x, where x. s..diameter, or (diameter) l e n g t h . 207 douglasii), the equations f o r s p i r e a were used; f o r Douglas f i r , white spruce was used; f o r c u r r a n t s , v a c c i n i a was used; f o r e l d e r b e r r y , black twinberry was used; and f o r s o a p a l a l l i e (Shepherdia canadensis) , a l d e r was used. 7.2.4 Data A n a l y s i s F l o r i s t i c data were analyzed as f o l l o w s . For the \"C\" l a y e r , canopy-coverage r a t i n g s were converted to t h e i r mid-point percent coverage values (see Table 7.1). Then data from a l l s i t e s that represented each stand were summed and d i v i d e d by the t o t a l number of quadrats (50 times number of s i t e s ) . Frequency of occurrence was the p r o p o r t i o n of a l l quadrats f o r a stand i n which a species was recorded, expressed as a percentage. I t i s important to remember that t h i s s t a t i s t i c i s s t r o n g l y a f f e c t e d by the area and shape of the quadrat used i n sampling (Greig-Smith 1964). However, as the 0.2 - x 0.5-m sampling frame i s commonly used i n western North America, many o p p o r t u n i t i e s f o r comparison e x i s t . For the purposes of. t h i s study, only \"major\" p l a n t species are presented. A major species was defined as one that had canopy-coverage greater than f i v e percent i n at l e a s t one of the s t a t i o n s r epresenting a stand. Complete data are on f i l e at the W i l d l i f e Research Section o f f i c e of the 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. 208 For the shrub and t r e e l a y e r s , species composition was based on stem counts. Phytomass estimates f o r the \"C\" l a y e r vegetation was based on weight data f o r each s e r a i stage. Shrub weight estimates f o r the \"B\" l a y e r were based on l o g a r i t h m i c r e g r e s s i o n equations. As B a s k e r v i l l e (1972) and other f o r e s t e r s have pointed out, these types of p r e d i c t i o n s are biased. This b i a s was not c o r r e c t e d f o r i n my r e s u l t s , but recent work by Brown (1976a)indicated t h a t i t was l i k e l y s m a l l , probably averaging f i v e percent. Height estimates were unbiased since p r e d i c t i o n equations u t i l i z e d a r i t h m e t i c rather than l o g a r i t h m i c data. Height and weight estimates were based on stem diameter measure-ments from a l l 2- x 2-m quadrats f o r each s e r a i stage. 7.3 Results f o r Mesic Upland P l a n t Communities 7.3.1 The Data Base and i t s P r e s e n t a t i o n Results were based on data c o l l e c t e d from 51 s i t e s . T h i r t y - f o u r s i t e s were i n the Deserters environmental u n i t ( m e s i c - t i l l ) , w i t h 15 i n mesic l a c u s t r i n e environments (Berman and Bednesti) and two i n h y d r i c t i l l . D i s t r i b u t i o n by age of stand was wide, ranging from 1 year a f t e r logging to 200 year o l d spruce-pine stands (Table 7.7). Except f o r e i g h t s i t e s , sampled stands had o r i g i n a t e d from w i l d f i r e or from f o r e s t s that were c l e a r c u t and burned. 209 Table 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 Pl a n t Succession Study Environment -substrate Nominal age (yr) S i t e number (age i n years i n parenthesis) M e s i c - t i l l 1 MF2 (1) , MF5(1), MF6(2), M l ( l ) , M3(l) 5 MF8(4), MF10(5) 10 MF22 (12) , Gl (11) , G4(11) 25 MF19(?)*, MF21(23) 45 MF4(39) , G2(41) , G3 (41) , SRI(45), SR3(52) 75 SR4(69) 110 MF1(116), MF9(106), SR9(119), SR11(110) 135 SR2(130).*, SR22 (137), SR23 (134) 150 MF3(167)*, SR5(148), SR13(151)* 200 MF7(195), MF1K179)*, M2 (195) , M4(195), SR6 (174), SR10 (183) H y d r i c - t i l l 110 SR16(115) 135 SR15(135) M e s i c - l a c u s t r i n e 1 MF12 (1) 5 MF15(4), MF17(4) 10 MF13(8), MF18(12) 25 MF20 (21) 45 SR12(54) 110 MF14 (101) , SR18(119) 150 MF16(154), SR8(153), SR14(140)*, SR17(151)*, SR20(142)*, SR21(149) T o t a l 51 s i t e s sampled * P a r t i a l l y logged stands. Others e i t h e r w i l d f i r e , or c l e a r c u t and burned. 210 Results are presented i n f i v e subsections. The f i r s t one describes the s e r a i stages f l o r i s t i c a l l y , t h a t i s , i n terms of p l a n t species composition and abundance. Each s e r a i stage i s described according to changes to the three vegetation l a y e r s : Tree, Shrub and Herb or A, B and C, r e s p e c t i v e l y . The second subsection documents more d e t a i l e d changes i n the Tree l a y e r from e a r l y to mature f o r e s t stages. Trends i n species composition, height, b a s a l area, and crown canopy c l o s u r e are the major aspects t r e a t e d . The t h i r d subsection deals i n d e t a i l w i t h temporal changes i n the Shrub l a y e r . P a r t i c u l a r a t t e n t i o n i s p a i d to changes i n phytomass and height. The f o u r t h subsection examines the Herb l a y e r , e s p e c i a l l y f o r changes i n phytomass. The f i n a l subsection provides data on the net primary p r o d u c t i v i t y of the v a s c u l a r understory vegetation. Results are presented f o r a l l the understory, that i s , f o r B and C l a y e r s combined. As shrubs form the major p a r t of the winter d i e t of moose, most of the data are f o r woody p l a n t s . 7.3.2 F l o r i s t i c Changes i n S e r a i Succession F l o r i s t i c changes r e l a t e s to v a r i a t i o n s i n p l a n t species composition and abundance recorded f o r seres on t i l l and l a c u s t r i n e s u b s t r a t e s . Results are presented f o r the A, B and C l a y e r s of each s e r a i stage. S e r a i p l a n t communities were i d e n t i f i e d by three-part names (Table 7.8). Each name represents the most abundant species i n the three 211 vegetation s t r a t a , beginning w i t h the \"A\" l a y e r . Only two names are provided when the tre e l a y e r was missing. Table 7.8 Pl 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 Nominal stand age P l a n t (yr) t i l l s ubstrate community name l a c u s t r i n e substrate 1 raspberry-geranium aspen-sedge 5 spi r e a - f i r e w e e d saskatoon-aster 10 wi l l o w - p i n e w i l l o w - w i l l o w 25 pin e - w i l l o w - t w i n f l o w e r a s p e n - w i l l o w - s a r s p a r i l i a 45 p i n e - r o s e - s p i r e a aspen-spirea-bunchberry 75 pine-twinberry-spruce 110 spruce-twinberry-subalpine f i r pine-subalpine f i r -bunchberry 135 s p r u c e - v a c c i n i a -subalpine f i r 150 spruce-subalpine f i r -cloudberry pine-spirea-bunchberry 200 spruce-thimbleberry-bunchberry P a r t i a l l y logged stands 135 s p r u c e - v a c c i n i a -bunchberry 150 spruce-maple-bunchberry subalpine f i r - v a c c i n i a -bunchberry 200 subalpine f i r - s q u a s h -berry-subalpine f i r 212 P l a n t succession proceeded through herb-, shrub- and tree-dominated stages. This p a t t e r n i s t y p i c a l of secondary f o r e s t succession whether i t o r i g i n a t e s from o l d f i e l d s , w i l d f i r e s , or c l e a r c u t t i n g and slashburning. The f o l l o w i n g paragraphs h i g h l i g h t important changes and d i f f e r e n c e s between s t u d i e d s e r a i stages ( F l o r i s t i c data f o r each s i t e are presented i n Appendix F) . Results are presented i n Tables 7.9 to 7.14 and s e l e c t e d s u c c e s s i o n a l stages are i l l u s t r a t e d i n Figure 7.2. T i l l - y e a r 1: Raspberry-Geranium The A and B l a y e r s were absent. The C l a y e r was ofte n patchy, depending upon the f i r e i n t e n s i t y , amount of r e s i d u a l s l a s h and s i t e disturbance. A l s o , species composition was v a r i a b l e , depending upon the foregoing f a c t o r s and the species composition and abundance of the pre-disturbance p l a n t community. However, raspberry and geranium (Geranium bioknellii) were the most common pioneer species. Other common shrub species i n the C l a y e r were rose and f l a t - t o p s p i r e a ; coniferous species were t y p i c a l l y absent or very inconspicuous. Geranium was abundant p a r t i c u l a r l y i n year 1, but decreased sharply by year 2. S a r s a p a r i l l a (Aralia nudioaulis), f ireweed (Epilobium angustifolium) , bunchberry (Cornus canadensis), sedges (Carex spp.), and bentgrass (Agrostis spp.) were common, having average cover exceeding 5 percent or frequency of occurrence Table 7.9 Percent Canopy-Coverage/Frequency of Occurrance 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 Nominal s u c c e s s i o n a l age (yr) 1 5 10 25 45 75 110 135 150 200 MAJOR GROUP Species EVERGREEN TREES AND SHRUBS: Abies lasiocarpa (subalpine f i r ) - - 1/2 - t / 1 * * t/4 15/35 21/54 14/32 7/19 Picea glauca (white spruce) - t/1 4/24 3/10 3/5 9/30 t/3 t/1 1/2 2/6 Pinus aontorta (lodgepole pine) - 1/9 25/46 - - 1/4 \u00E2\u0080\u0094 \u00E2\u0080\u0094 \u00E2\u0080\u0094 \u00E2\u0080\u0094 DECIDUOUS TREES AND SHRUBS: Acer glabrum (Douglas maple) - - - - - - - - - -Betula papyrifera (paper b i r c h ) - 1/10 - 2/14- - - - 1/2 \u00E2\u0080\u0094 \u00E2\u0080\u0094 Lonicera involucrata (black twinberry) - - - - - - - - - \u00E2\u0080\u0094 Populus tremuloides (trembling aspen) - t/2 - 1/4 - - - - - -Rosa spp. (rose) 4/21 5/16 10/50 2/14 7/36 4/18 t/4 1/9 t/2 2/10 Rubus idaeus (raspberry) 7/29 7/36 t/2 - - - - t/1 t/2 2/9 R. parviflorus (thimbleberry) 3/9 7/28 - - 3/9 2/8 2/11 5/21 5/24 4/13 Salix spp. (willow) - 2/14 9/32 - t/1 - - - - -Spiraea lucida ( f l a t - t o p spirea) 5/27 6/28 7/52 - 11/56 8/36 3/19 4/22 - 3/24 S. douglasii (hardhack) .- - - - - 2/24 1/5 - 1/12 -Vaccinium caespitoswn (dwarf blueberry) - - t/2 - - - - - - -V. membranaceum (mountain b i l b e r r y ) t/5 - 2/20 - t/3 2/12 3/19 16/57 3/22 5/20 Viburnum edule (squashberry) t/1 1/5 t/2 - 1/4 - 2/7 t/4 1/12 t/1 FORBS AND DWARF SHRUBS: Achillea millefolium (yarrow) - - 1/14 - - 1/4 - - - -Aralia nudicaulis ( s a r s a p a r i l l a ) 3/29 5/32 - - 2/20 - 5/24 7/38 16/66 7/30 Aster spp. (aster) t/3 1/7 7/36 - 1/5 2/8 t/1 - - 1/3 Clintonia uniflora (queen's cup) t/5 1/9 1/10 - 1/4 1/14 5/25 7/51 2/28 4/25 Cornus canadensis (bunchberry) 4/43 17/77 12/58 - 7/51 4/40 8/46 13/79 10/72 12/71 Epilobium spp. (fireweed) 8/43 21/79 2/18 - 2/8 t/2 - t/2 1/6 t/2 Table 7.9, Continued Nominal s u c c e s s i o n a l age (yr) MAJOR GROUP Species 1 5 10 25 45 75 110 135 150 200 Galium (bedstraw) _ 1/9 t/4 \u00E2\u0080\u0094 - t/2 1/7 1/3 1/6 t/1 Geranium bioknellii ( B i c k n e l l s geranium) 21/49 t/1 - - - - - - - -Hypochaeris radieata (cat's ear) - - 6/54 - - - - - - -Linnaea borealis (twinflower) t/4 3/5 14/52 18/56 9/44 3/18 1/11 6/28 2/18 4/25 Retasites frigida ( c o l t s f o o t ) - - - - 1/8 3/8 t/5 t / 1 - 1/6 Rubus chamaemorus (cloudberry) - - - t/2 \u00E2\u0080\u00A2 - 4/24 2/13 3/21 19/88 1/2 R. pedatus ( t r a i l i n g rubus) - \u00E2\u0080\u00A2 - - - - - 3/15 1/7 - 2/9 Streptopus amplexifolius (twisted s t a l k ) - - - - 3/16 6/40 3/14 4/24 2/6 3/10 Trifolium repens (white clover) - - - - - - - - - -GRAMMINOIDS Agrostis alba (bentgrass) 6/16 - 10/34 - - - - - - t/1 Calamagrostis spp. (reedgrass) 6/2 t/1 3/10 - 10/17 - - - - -Carex spp. (sedge) 4/26 1/7 6/20 - 4/8 - - 6/1 - -OTHER TAXA: Dryopteris austriooa (spiny wood-fern) t/1 - - - t/1 3/10 3/17 7/32 16/58 4/17 Equisetum spp. (horse t a i l ) - - - - 1/3 4/22 2/11 2/8 2/6 4/15 No. spp. recorded ..27 31 29 7 32 28 32 35 24 38 No. s i t e s sampled (quadrats) 3(150) 2(100) 3(150) 1(50) 2(100) 1(50) 3(150) 3(150) 1(50) 3(150) *Species w i t h at l e a s t one s i t e where canopy-coverage s > 5% and frequency of occurrence > 20%. * * t i s l e s s than 0.5% coverage. Table 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 Nominal s u c c e s s i o n a l age (yr) MAJOR GROUP Species 1 5 10 25 45 110 150 5/12 t/ 2 * 1/3 12/32 7/18 - - 5/17 - 9/13 1/4 2/6 1/5 t/2 1/1 - -5/27 3/6 1/1 - -1/6 . 2/14 4/18 4/10 5/19 1/6 4/21 6/28 5/15 7/14 t/4 - - 6/1 6/36 7/52 9/43 1/10 7/33 2/17 5/29 - t/4 2/15 t/2 - - -- 1/8 1/7 1/8 1/1 t/2 1/4 t/2 1/3 19/50 7/16 3/12 - t/1 1/18 1/9 3/14 4/12 5/28 7/32 12/49 2/12 9/26 1/4 - 6/33 t/1 2/4 9/60 1/12 3/16 - - 1/6 1/2 - - - t/2 7/30 12/42 6/25 \u00E2\u0080\u0094 2/10 1/4 5/28 1/2 \u00E2\u0080\u0094 1/9 1/5 5/29 t/2 1/2 t/3 - 1/8 - 18/62 3/15 1/8 3/21 4/30 11/60 5/37 - 1/6 2/16 7/33 - 1/6 1/3 2/2 - 1/12 5/37 EVERGREEN TREES AND SHRUBS: Abies lasiocarpa (subalpine f i r ) Picea glauca (white spruce) Pinus contorta (lodgepole pine) DECIDUOUS TREES AND SHRUBS: Acer glabrum (Douglas maple) Betula papyrifera (paper b i r c h ) Lonicera involucrata (black twinberry) Populus tremuloiales (trembling aspen) Rosa spp. (rose) Rubus idaeus (raspberry) R. parviflorus (thimbleberry) Salix spp. (willow) Spiraea lucida ( f l a t - t o p spirea) S. douglasii (hardhack) Vaccinium caespitosum (dwarf blueberry) V. membranaceum (mountain b i l b e r r y ) Viburnum edule (squashberry) FORBS AND DWARF SHRUBS: Achillea millefolium (yarrow) Aralia nudicaulis ( s a r s a p a r i l l a ) Aster spp. (aste r ) Clintonia uniflora (queen's cup) Table 7.10, Continued MAJOR GROUP Species Nominal s u c c e s s i o n a l age (yr) 1 5 10 25 45 110 150 Cornus oanadenis (bunchberry) 9/80 7/58 9/63 13/68 9/63 27/90 16/79 Epilobium spp. (fireweed) - 3/25 9/65 8/50 1/5 t/5 t/4 Galium spp. (bedstraw) - 3/22 7/21 1/10 1/8 1/9 2/17 Geranium bioknellii ( B i c k n e l l s geranium) \u00E2\u0080\u00A2t/2 2/17 t/1 t/2 - - -Hypochaeris radieata (cat's ear) - - 1/1 - - - -Linnaea borealis (twinflower) 1/8 t/6 t/3 t/2 7/48 7/35 3/23 Petasites frigida ( c o l t s f o o t ) 3/58 8/59 9/44 2/28 4/36 9/52 5/44 Rubus chamaemorus (cloudberry) - 6/40 3/17 t/2 3/20 2/11 1/7 R. pedatus ( t r a i l i n g rubus) 1/8 8/28 1/4 t/2 1/5 1/4 3/13 Streptopus amplexifolius (twisted s t a l k ) - - - - 1/7 - 3/5 Trifolium repens (white clover) - - 1/2 8/32 - - -GRAMMINOIDS: Agrostis alba (bentgrass) t/2 - - - - - -Calamagrostis spp. (reedgrass) 2/6 9/21 3/13 7/32 - 1/10 t/1 Carex spp. (sedge) 18/64 1/10 2/11 2/10 1/2 - -OTHER TAXA: Dryopteris austrioaa (spiny wood-fern) - - - - 1/14 1/7 11/20 Equisetum spp. (horse : t a i l ) - - - - 5/26 2/14 2/8 No. spp. recorded 20 33 36 32 33 33 39 No. s i t e s sampled (quadrats) 1(50) 2(100) 2(100) 1(50) 2(100) 2(100) 3(150) *Species w i t h at l e a s t one s i t e where canopy-coverage s > 5% and frequency of occurrence k 20%. * * t i s l e s s than 0.5% coverage. 217 Table 7.11 Percent Canopy-Coverage/Frequency of Occurrence Values f o r Major* Plant Species of the Herb (C) Layer i n P a r t i a l l y Logged Sub-Boreal F o r e s t Stands i n a Mesic Environment on T i l l and Lacustrine Substrates T i l l Lacustrine MAJOR GROUP Species 135 150 200 EVERGREEN TREES AND SHRUBS: Abies lasiocarpa (subalpine f i r ) Picea glauca (white spruce) Pinus contovta (lodgepole pine) DECIDUOUS TREES AND SHRUBS: Acer glabrum (Douglas maple) Betuhx papyrifera (paper birch) Lonicera involucrata (black twinberry) Populus tremuloides (trembling as Rosa spp. (rose) Rubus idaeus (raspberry) R. parviflorus (thimbleberry) Salix spp. (willow) Spiraea lucida (flat-top spirea) S. douglasii (hardhack) Vacoinium caespitosum (dwarf blueberry) V. membranaceum (mountain b i l b e r Viburnum edule (squashberry) FORBS AND DWARF SHRUBS: Achillea millefolium (yarrow) Aralia nudicaulis (sarsaparilla) Aster spp. (aster) Clintonia uniflora (queen's cup) Cornus canadensis (bunchberry) Epilobium spp. (fireweed) Galium spp. (bedstraw) Geranium bicknellii (Bicknell's geranium) Hypochaeris radicata (cat's ear) Linnaea borealis (twinflower) Petasites frigida (coltsfoot) Rubus chamaemorus (cloudberry) R. pedatus ( t r a i l i n g rubus) Streptopus amplexifolius (twisted stalk) Trifolium repens (white clover) CRAMMINOIDS: Agrostis alba (bentgrass) Calamagrostis spp. (reedgrass) Carex spp. (sedge) OTHER TAXA: Dryopteris austrioca (spiny woe Equisetum spp. (horse t s i l ) No. spp. recorded No. s i t e s sampled (quadrats) 16/50 10/32 16/36 1/2 1/6 1/6 150 16/34 1/3 8/22 8/15 1/1 3/5 1/4 3/8 2/9 t/21* 2/6 , 1/3 pen) - 1/3 - 1/5 2/22 2/13 3/12 1/7 E/2 - - 1/2 11/45 9/41 3/14 4/19 - t/4 -7/32 9/36 8/38 5/25 2/14 f) 26/90 9/38 - 11/37 t/2 3/11 1/2 t/3 7/52 5/28 4/32 7/40 t/1 6/20 2/8 10/70 3/20 1/12 3/38 21/92 11/70 12/58 17/81 t/5 3/14 2/18 1/6 2/10 t/5 t/6 t/1 13/47 7/32 9/38 11/35 1/4 \u00E2\u0080\u00A2- -3/20 t/1 - 2/25 1/6 1/5 1 1/6 2/15 1/5 . t/2 -- --fern) t/2 4/16 2/10 - 2/14 1/10 t/1 25 32 26 37 1(50) 2(50) 1(50) 3(150) where canopy -coverage s 1 5% and frequency of occurrence 2. 20%. * * t i s less than 0.5% coverage. Table 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 Nominal succes s s i o n a l age (yr) Species 1 5 10 25 45 75 110 135 150 200 Abies lasiocarpa (subalpine f i r ) 16 15 32 15 Acer gldbrum (Douglas maple) 11 2 11 Alnus spp. (alder) 3 13 6 Amelanchier alnifolia (Saskatoon) 1 15 26 Betula papyrifera (paper b i r c h ) 4 30 Cornus stolonifera ( r e d - o s i e r dogwood) 2 5 Lonicera involucrata (black twinberry) 14 6 32 22 Picea glauca (white spruce) 5 15 17 2 Pinus contorta (lodgepole pine) 2 15 Populus tremuloides (trembling aspen) 5 2 1 P. balsamifera (black cottonwood) Pseudotsuga menziesii (Douglas f i r ) 1 2 Ribes spp. (currant) 4 10 2 2 16 5 Rosa (rose) 17 16 5 7 37 3 5 16 9 Rubus idaeus (raspberry) 73 3 R. parviflorus (thimbleberry) 3 6 9 11 31 Salix spp. (willow) 1 56 48 Sambucus racemosa (elderberry) 5 Shepherdia canadensis ( s o a p a l a l l i e ) 11 Sorbus spp. (mountain ash) 5 11 2 2 Spiraea lucida ( f l a t - t o p spirea) 23 4 3 6 9 19 5 S. douglasii (hardhack) 28 10 5 16 Vaccinium spp. (v a c c i n i a ) 3 4 51 5 1 Viburnum edule (squashberry) 11 11 5 10 No. stems 41 192 55 61 64 31 174 47 19 81 S i t e s 3 2 1 1 3 1 4 2 1 3 Table 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 T i l l Substrates Nominal s u c c e s s i o n a l age (yr) Species 1 5 10 25 45 110 150 Abies lasiooarpa (subalpine f i r ) 5 23 2 Acer glabrum (Douglas maple) Alnus spp (alder) Amelanchier alnifolia (Saskatoon) 23 8 13 Betula papyrifera (paper b i r c h ) 6 5 Cornus stolonifera ( r e d - o s i e r dogwood) 4 2 Lonicera involucrata (black twinberry) 7 13 3 16 . 21 6 Picea glauoa (white spruce) 7 Pinus contorta (lodgepole pine) 1 Populus tremuloides (trembling aspen) 93 18 4 6 P. balsamifera (black cottonwood) 2 Pseudotsuga menziesii (Douglas f i r ) Ribes spp. (currant) 3 1 Rosa spp. (rose) 19 1 20 21 15 Rubus idaeus (raspberry) R. parviflorus (thimbleberry) 2 4 Salix spp. (willow) 4 72 83 8 1 Sambucus racemosa (elderberry) Sheperdia canadensis ( s o a p a l a l l i e ) Sorbus spp. (mountain ash) 4 Spiraea lucida ( f l a t - t o p spirea) 3 36 6 30 S. douglasii (hardhack) 15 19 18 Vaooinium spp. (va c c i n i a ) 2 Viburnum edule (squashberry) 1 2 7 No stems ( s i t e s ) 30(1) 113(2) 318(2) 64(1) 25(1) 48(2) 188(3) AO 220 Table 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 T i l l L a c u s t r i n e Species 135 150 200 150 Abies lasiocarpa (subalpine f i r ) 8 14 6 Acer glabrum (Douglas maple) 19 26 6 Alnus spp. (alder) Amelanchier alnifolia (Saskatoon) Betula papyrifera (paper b i r c h ) 1 3 3 Cornus stolonifera ( r e d - o s i e r dogwood) Lonioera involucrata (black twinberry) 3 17 6 Picea glauoa (white spruce) 3 Pinus contorta (lodgepole pine) Populus tremuloides (trembling aspen) 2 9 7 P. balsamifera (black cottonwood) 1 Pseudotsuga menziesii (Douglas f i r ) Ribes spp. (currant) 3 Rosa spp. (rose) 3 12 6 1 Rubus idaeus (raspberry) R. parviflorus (thimbleberry) 2 10 Salix spp. (willow) Sambuous raoemosa (elderberry) Shepherdia canadensis ( s o a p a l a l l i e ) Sorbus spp. (mountain ash) f 2 6 3 Spiraea lucida ( f l a t - t o p spirea) 4 9 19 S. douglasii (hardhack) 14 Vacoinium spp. ( v a c c i n i a ) 43 16 21 Viburnum edule (squashberry) 14 11 54 18 No. of stems ( s i t e s ) 37(1) 125(2) 35(1) 72(3) 220a Figure 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 s u b s t r a t e s . Page 221 shows t i l l s i t e s aged approximately 1(a), 10(b), 45(c) and 200(d) years. Page 222 shows l a c u s t r i n e s i t e s aged approximately 5 and 25 years. Page 22 3 compares t i l l and l a c u s t r i n e s i t e s at approximately 40 years (a and b) and at (c and d), r e s p e c t i v e l y . 221 223 224 greater than 25 percent or both. Grasses tended to be most common i n e a r l y stages south of Pr i n c e George, and uncommon or absent i n northern areas. Moss cover was v a r i a b l e ; the l i v e r w o r t , marchantia (Mavohantia polymorpha) occurred on the more severely burned s i t e s but was not abundant. T i l l - y e a r 5: Spirea-fireweed The Tree (A) l a y e r remained undeveloped, but the Shrub (B) l a y e r showed considerable change from year 1, w i t h respect to species d i v e r s i t y and canopy-coverage. Douglas maple, black twinberry, rose, squashberry and f l a t - t o p s p i r e a were the most common members of the Shrub l a y e r . Less common were paper b i r c h , currants raspberry, mountain ash and v a c c i n i a . The Herb (C) l a y e r was w e l l developed w i t h fireweed and bunchberry most abundant. Frequently o c c u r r i n g species were rose, raspberry, thi'mbleberry, f l a t - t o p s p i r e a and s a r s a p a r i l l a . Geranium was almost absent and remained so f o r the duration of the sere. Grasses, g r a s s l i k e species and other taxa such as ferns and h o r s e t a i l s were uncommon or absent. T i l l - y e a r 10: Willow-pine As i n e a r l i e r stages, the Tree l a y e r was absent. The Shrub l a y e r d e c l i n e d s l i g h t l y i n d i v e r s i t y and densit y . Willow dominated t h i s s e r a i stage w i t h saskatoon and pine f r e q u e n t l y o c c u r r i n g but much l e s s f r e q u e n t l y than w i l l o w . White spruce, rose and f l a t - t o p s p i r e a were l e a s t common. The Herb l a y e r was diverse and more complex than i n e a r l i e r stages. C o n i f e r s , e s p e c i a l l y lodgepole pine, were very abundant and subalpine f i r appeared f o r the f i r s t time. Other common species were rose, w i l l o w , f l a t - t o p s p i r e a , a s t e r (Aster spp.), bunchberry, twinflower {Linnaea borealis) and bentgrass. V a c c i n i a , white spruce, fireweed, c a t s ' ear {Hypoehaeris radiaata) and sedges occurred f r e q u e n t l y but ge n e r a l l y had low coverage. T i l l - y e a r 25: Pi n e - w i l l o w - twinflower By t h i s time the Tree l a y e r had developed. Pine was the major tr e e species, followed by paper b i r c h and wil l o w s i n approximately equal p r o p o r t i o n s . The Shrub l a y e r continued to show a reduction i n number of species. As w i t h year 10, w i l l o w c l e a r l y dominated t h i s l a y e r . Paper b i r c h was next i n abundance, followed by white spruce, rose and trembling aspen. Species composition of the shrub l a y e r at t h i s stage was more v a r i a b l e and diverse than i n d i c a t e d by the s i n g l e sample. The Herb l a y e r was considerably reduced i n d i v e r s i t y and species coverage. Twinflower was w e l l represented but none of the other species had coverages exceeding 5 percent. Ranked w e l l below twinflower were white spruce, paper b i r c h , black twinberry, and rose. 2 2 6 T i l l - y e a r 45: P i n e - r o s e - s p i r e a The Tree l a y e r was w e l l defined by t h i s stage, and c o n s i s t e d almost e n t i r e l y of lodgepole pine. Deciduous tre e species occurred o c c a s i o n a l l y , v i z . , paper b i r c h and w i l l o w . White spruce was uncommon. The Shrub l a y e r was dominated by rose, hardhack and white spruce. Lodgepole pine was e s s e n t i a l l y absent i n t h i s l a y e r since by t h i s age, they had grown out of the Shrub l a y e r and i n t o the Tree l a y e r . The lack of subsequent pine regeneration a l s o meant that pine d i d not reappear i n the lower l a y e r s . R e l a t i v e l y l e s s common shrub species were a l d e r , black twinberry, thimbleberry and f l a t - t o p s p i r e a . Increasing d i v e r s i t y c h a r a c t e r i z e d the Herb l a y e r at t h i s stage. In a d d i t i o n to s p i r e a , rose, bunchberry, twinflower and reedgrasses ( Cinna latifolia) were common. Shade-tolerant species appeared or became more abundant than i n previous years, v i z . , mitrewort (Mitella nuda) , c o l t s f o o t (Petasites frigidus) , t w i s t e d s t a l k [Stveptopus amplexifolius), and f e r n s . T i l l - y e a r 75: Pine- twinberry-spruce The Tree l a y e r again was almost e n t i r e l y lodgepole pine. White, spruce comprised l e s s than 10 percent of the tree stems. Deciduous species were r a r e , w i t h none encountered i n the sample p l o t s f o r t h i s s e r a i stage. The Shrub l a y e r reached i t s lowest recorded density 2 27 (Table 7-12). In a d d i t i o n to twinberry, saskatoon was v common; l e s s abundant were a l d e r , currants and hardhack. Rose and s p i r e a were uncommon. Spruce, black twinberry, f l a t - t o p s p i r e a and t w i s t e d s t a l k were most prominent members of the Herb l a y e r . A l so common but w i t h lower coverages were rose, v a c c i n i a , queen's cup [Clintonia uni flora), bunchberry, twinf lower, mitrewort, cloudberry {Rubus ohamaemorus), mountain r i c e g r a s s {Oryzopsis asperifolia) , and h o r s e t a i l . Ferns continued to become more abundant. T i l l - y e a r 110: Spruce- twinberry-subalpine f i r White spruce replaced pine as the commonest t r e e i n the Tree l a y e r , although both species were abundant. Subalpine f i r made i t s f i r s t appearance i n the tree l a y e r . Douglas f i r and paper b i r c h were o c c a s i o n a l members. In the Shrub l a y e r , species composition was more div e r s e than at year 75. Fourteen species were recorded compared to 7 f o r the e a r l i e r stand. Subalpine f i r was common as w e l l as black twinberry. Of intermediate abundance were mountain ash, f l a t - t o p s p i r e a and squashberry. A l d e r , r e d - o s i e r dogwood, rose, thimbleberry, hardhack and v a c c i n i a were l e s s common than the foregoing shrubs. The Herb l a y e r was c h a r a c t e r i z e d by s a r s a p a r i l l a , queen's cup and bunchberry. Ferns remained r e l a t i v e l y common while grasses d e c l i n e d i n d i v e r s i t y and coverage. 228 T i l l - y e a r 135: Spruce- v a c c i n i a - s u b a l p i n e f i r Spruce continued as the major tree species. Pine d e c l i n e d , p r i m a r i l y at the expense of subalpine f i r . S c a t tered paper b i r c h p e r s i s t e d i n the maturing f o r e s t . By t h i s age, the Shrub l a y e r appeared to d e c l i n e i n species d i v e r s i t y and stem d e n s i t y . While v a c c i n i a c l e a r l y dominated the stem composition, other common species were subalpine f i r and f l a t - t o p s p i r e a . In the Herb l a y e r , mountain b i l b e r r y {Vaooinium membranaceum), and bunchberry were q u i t e common. Other f r e q u e n t l y o c c u r r i n g species were thimbleberry, s p i r e a , s a r s a p a r i l l a , queen's cup, twinflower, t w i s t e d s t a l k and spiny wood-fern {Dryopteris austriaca). T i l l - y e a r 150: Spruce- subalpine f i r - c l o u d b e r r y Spruce continued to maintain i t s p o s i t i o n as the dominant tree species i n t h i s s e r a i stage. Lodgepole was al s o common, being at comparable l e v e l s to the previous stage. Paper b i r c h was found i n f r e q u e n t l y . The dominance of subalpine f i r i n the Shrub l a y e r from previous dominance of the herb l a y e r r e f l e c t e d the gradual development of t h i s species i n sub-boreal f o r e s t s . Less common species were c u r r a n t s , rose, thimbleberry and hardhack. The Herb l a y e r continued to show subalpine f i r as a very common species , too, along w i t h s a r s a p a r i l l a , bunchberry, and spiny wood-fern. Thimbleberry and currants were l e s s abundant but occurred f r e q u e n t l y . T i l l - y e a r 200: Spruce- thimbleberry-bunchberry The Tree l a y e r presented a composition g e n e r a l l y s i m i l a r to the three previous stands. White spruce was commonest, followed by lodgepole pine and subalpine f i r . Douglas f i r and b i r c h occurred s p o r a d i c a l l y . The Shrub l a y e r had s e v e r a l s i m i l a r i t i e s w i t h other stands exceeding 100 years i n age. Among the most common species were subalpine f i r and thimbleberry. The dominance of thimbleberry represented the peak of a trend that began at approximately age 45. Saskatoon, paper b i r c h , lodgepole pine, raspberry and w i l l o w were absent. S o a p a l a l l i e occurred f o r the f i r s t time, although observations elsewhere i n d i c a t e d t h a t i t was present i n other stands before t h i s stage. The Herb l a y e r , too, had common features of other, younger stands. Subalpine f i r , v a c c i n i a , s a r s a p a r i l l a , queen's cup, bunchberry, twinflower, t w i s t e d s t a l k and spiny wood-fern were g e n e r a l l y the commonest species. Grasses were commonly absent or very i n f r e q u e n t . Although l a c u s t r i n e s i t e s were sampled l e s s i n t e n s i v e l y than t i l l s i t e s , d i f f e r e n c e s i n su c c e s s i o n a l f l o r i s t i c s were obvious. The f o l l o w i n g paragraphs h i g h l i g h t 230 the s t u d i e d stages and document d i f f e r e n c e s between the substrat e s . L acustrine-year 1: Aspen-sedge As w i t h t i l l , the Tree l a y e r was absent i n e a r l y stages of development. However, i n i t i a l Shrub l a y e r development was apparent. The abundance and growth of aspen probably r e f l e c t e d the r e s p r o u t i n g response from previous aspen t r e e s . L a c u s t r i n e - y e a r 5: Saskatoon-aster Although the Tree l a y e r remained undeveloped, the Shrub l a y e r increased i n species d i v e r s i t y and density. Almost as common as saskatoon were black twinberry, trembling aspen, rose and hardhack. Absent or uncommon were the t y p i c a l t i l l species of Douglas maple, paper b i r c h , and f l a t - t o p s p i r e a . Rose and twinberry were common on both substra t e s . By year 5, the Herb l a y e r was diverse and lu s h . Many species were common w i t h coverages exceeding 5 percent and frequency of occurrence greater than 2 0 percent: rose, hardhack, bunchberry, c o l t s f o o t , cloudberry, t r a i l i n g rubus (Rubus pedatus), reedgrass and fescue {Festuaa oaaidentalis). Fireweed, raspberry, thimbleberry and s a r s a p a r i l l a were uncommon. 231 L a c u s t r i n e - y e a r 10: Willow-willow As the community name i n d i c a t e s , t h i s stage was c l e a r l y dominated by w i l l o w s . The Shrub l a y e r was 72 per-cent w i l l o w and other species each comprised 7 percent or l e s s of the stand. Stem d e n s i t i e s were the highest recorded for any stage or substrate. Lodgepole pine was very in f r e q u e n t and subalpine f i r represented 5 percent of the stems (cf. w i t h t i l l ) . In the Herb l a y e r , coniferous species were not common, e s p e c i a l l y lodgepole pine: t h i s l a t t e r species was the most common one i n the herb l a y e r on t i l l s u b s t r a t e s . In a d d i t i o n to w i l l o w , other common species i n the herb l a y e r were paper b i r c h , rose, yarrow {Achillea millefolium), a s t e r , bunchberry, fireweed, bedstraw {Galium spp.), c o l t s f o o t and dandelion {Taraxacum officinale) . While a few of these species were a l s o common i n t i l l communities, the c o n s t e l l a t i o n of common species was c l e a r l y d i f f e r e n t . L a c u s t r i n e - y e a r 25: Aspen- w i l l o w - s a r s a p a r i l i a The appearance of a Tree l a y e r re-emphasizes the prolonged deciduous phase on l a c u s t r i n e as compared wi t h t i l l s u b s t r a t e s . Trembling aspen was v i r t u a l l y the only tree species. On both t i l l and l a c u s t r i n e m a t e r i a l s , w i l l o w s dominated the Shrub l a y e r of t h i s s e r a i stage. Willow 232 comprised 83 percent of the shrub stand, w i t h the remaining species of paper b i r c h , aspen, cottonwood and squashberry forming the balance. Paper b i r c h and white spruce were common i n the t i l l but not i n the l a c u s t r i n e s o i l s . U nlike the Herb l a y e r on the t i l l , complexity of the l a c u s t r i n e Herb l a y e r d i d not d e c l i n e from the previous s e r a i stage. In a d d i t i o n to s a r s a p a r i l l a were the common, fr e q u e n t l y o c c u r r i n g squashberry, bunchberry, fireweed, wintergreen (Pyrola spp.), dandelion, white c l o v e r (Trifolium . re-pens), and reedgrass. This compared markedly w i t h the impoverished f l o r a on t i l l where only twinflower was common. Coniferous seedlings were uncommon, s i m i l a r to the t i l l stands. Lacustrine-year 45: Aspen-spirea-bunchberry Aspen continued to dominate the Tree l a y e r , making up almost 70 percent of the stems. Emergence of the coniferous species was revealed by the presence of lodgepole pine and white spruce. Subalpine f i r occurred too, but only i n f r e q u e n t l y . The Shrub l a y e r was l e s s d i verse than i t s t i l l c ounterpart. In a d d i t i o n to s p i r e a , the only other common shrubs were black twinberry and rose compared w i t h white spruce, rose and hardhack on t i l l . Stem d e n s i t i e s were comparable i n both substrates. As i n the age 25 stand, common herb species of the 233 two substrates overlapped only s l i g h t l y . The common l a c u s t r i n e herb species were black twinberry, rose, s p i r e a , hardhack, mountain b i l b e r r y , t winflower, c o l t s f o o t and h o r s e t a i l . Of these, only the f i r s t , t h i r d and l a s t were e q u a l l y as common on t i l l . L a c u s trine-year 110: Pine- subalpine f i r - b u n c h b e r r y By t h i s stage, t i l l and l a c u s t r i n e p l a n t communities demonstrated some convergence. In the Tree l a y e r , pine remained the commonest species, while trembling aspen became almost i n s i g n i f i c a n t . White spruce was almost as common as pine, subalpine f i r was o c c a s i o n a l and Douglas f i r , i n f r e q u e n t . Paper b i r c h was r a r e l y encountered. In the Shrub l a y e r , subalpine f i r , b lack twinberry, rose, mountain ash, v a c c i n i a and the spi r e a s were species i n common on both substrates. Abundance of a l d e r , r e d - o s i e r dogwood, saskatoon and thimbleberry d i f f e r e d between l a c u s t r i n e and t i l l u n i t s . In the Herb l a y e r , common species were subalpine f i r , f l a t - t o p s p i r e a , mountain b i l b e r r y , bunchberry, queen's cup, twinflower and c o l t s f o o t . Many of these were a l s o common i n the s e r a i stage on t i l l . Less abundant but commonly o c c u r r i n g species were saskatoon, Canada blueberry {Vaccinium myrtillus), s a r s a p a r i l l a , bedstraw and h o r s e t a i l . 234 L a c u s t r i n e - y e a r 150: Pine- spirea-bunchberry The Tree l a y e r was b a s i c a l l y c o n i f e r o u s , c o n s i s t i n g of lodgepole pine and white spruce. Subalpine f i r represented about 5 percent of the stem or approximately one-half that i n comparable stands on t i l l . Aspen and paper b i r c h remained uncommon. In the Shrub l a y e r , the number of species continued to increase from a low of f i v e at year 45 to 12 at year 150. Stem d e n s i t i e s showed a s i m i l a r trend. Besides s p i r e a , other common species were saskatoon, rose and hardhack. Most species c h a r a c t e r i s t i c of e a r l y s e r a i stages were absent, v i z . , paper b i r c h , lodgepole pine, trembling aspen and w i l l o w . Species common on t i l l but absent or infrequent on l a c u s t r i n e substrates were Douglas maple, saskatoon, thimbleberry, and s o a p a l a l l i e . The Herb l a y e r was g e n e r a l l y s i m i l a r to the year 110 s e r a i stage w i t h respect to species composition and abundance. Major increases were shown by black twinberry, s a r s a p a r i l l a , a s t e r , queen's cup, wood reedgrass and wood-fe r n . A l s o , the communities of both t i l l and l a c u s t r i n e had many species i n common, e.g. those above plus subalpine f i r , r ose, s p i r e a , mountain b i l b e r r y , bunchberry, twinflower and h o r s e t a i l s . In summary, s u c c e s s i o n a l d i f f e r e n c e s i n f l o r i s t i c s between t i l l and l a c u s t r i n e substrates were: 235 1. A g e n e r a l l y more d i v e r s e and l u s h herb and shrub l a y e r on the l a c u s t r i n e s o i l s . 2. A prolonged deciduous shrub and t r e e phase on l a c u s t r i n e s u b s t r a t e s . 3. Lodgepole pine was comparatively l e s s s i g n i f i c a n t on l a c u s t r i n e s o i l s than, on t i l l s o i l s . 4. An i n c r e a s i n g convergence i n s p e c i e s composition and abundance between the two s u b s t r a t e s as the seres developed. 5. A pronounced r e d u c t i o n i n understory d i v e r s i t y around year 25 on t i l l but not on l a c u s t r i n e parent m a t e r i a l s . The r e l a t i v e l y s m a l l sample s i z e p r e c l u d e d a f u l l d e s c r i p -t i o n of f l o r i s t i c s i m i l a r i t i e s and d i f f e r e n c e s , but i t was s u f f i c i e n t to demonstrate g e n e r a l a t t r i b u t e s . P a r t i a l l y logged stands were a l s o d e s c r i b e d (Tables 7-11 and 7-14). On t i l l s u b s t r a t e s , the t r e e l a y e r c o n s i s t e d of one-half or more of white spruce except f o r one s e l e c t i v e l y logged, 200 year o l d stand where subalpine f i r was as abundant. Lodgepole pine was g e n e r a l l y a minor member of t h i s l a y e r . Paper b i r c h was g e n e r a l l y more common than p i n e , making up to 19 per c e n t of the stems. No oth e r s p e c i e s were recorded. The shrub l a y e r s i n s e l e c t i v e l y logged f o r e s t s on t i l l were s i m i l a r to un d i s t u r b e d stands w i t h r e s p e c t to the abundance of subalpine f i r , b l a c k twinberry, rose, s p i r e a and v a c c i n i a . Douglas maple and squashberry were commonest 236 i n the logged stands. Herb l a y e r i n both d i s t u r b e d and undisturbed stands were g e n e r a l l y a l i k e . The commonest species were subalpine f i r , Douglas maple, thimbleberry, f l a t - t o p s p i r e a , mountain b i l b e r r y , s a r s a p a r i l l a , queen's cup, bunchberry and twinflower. Ferns and h o r s e t a i l s were n o t i c e a b l y i n f r e q u e n t i n the logged s i t e s . Three p a r t i a l l y logged stands were sampled on the l a c u s t r i n e s u b s t r a t e , a l l 150 year o l d stands (Tables 7-11 and 7-14). In the s i t e s sampled, subalpine f i r was commonest i n the tree l a y e r . White spruce and lodgepole pine were s l i g h t l y l e s s abundant. Douglas f i r and aspen were uncommon i n the overstory. The Shrub l a y e r was more dense and more diverse than t h e i r unlogged counterparts. Twinberry, aspen, s p i r e a , v a c c i n i a and squashberry were most abundant i n the logged stand, w h i l e subalpine f i r , c u r r a n t s , rose, and hardhack were the commonest species i n the corresponding 150 year o l d f o r e s t . The Herb l a y e r a l s o showed some marked d i f f e r e n c e s from the f l o r i s t i c s of unlogged areas (Table 7-11). Subalpine f i r , thimbleberry, s a r s a p a r i l l a and twinflower were most abundant i n logged stands. Less common species i n these d i s t u r b e d stands were spiny wood-fern, wood reed grass, c o l t s f o o t , rose and a s t e r . 237 7.3.3 Temporal Dynamics of the Tree Layer The t r e e l a y e r developed through two or three main stages. F i r s t , a t r a n s i t i o n occurred from p r i m a r i l y a deciduous f o r e s t to a coniferous one (Table 7.15). On t i l l s u b s t r a t e s , the deciduous phase was b r i e f , l a s t i n g only f o r the f i r s t 25 years of succession; a f t e r 25 years, evergreens were more numerous than deciduous t r e e s . On l a c u s t r i n e s u b s t r a t e s , the deciduous phase was more obvious and longer-l a s t i n g than on t i l l . C o n i f e r s appeared more than 25 years a f t e r i n i t i a t i o n of the sere, and d i d not form the m a j o r i t y of stems u n t i l a f t e r 45 years (Table 7.15). On both t i l l and l a c u s t r i n e s u b s t r a t e s , a few deciduous t r e e s p e r s i s t e d i n mature f o r e s t s . The second main stage was the s h i f t from lodgepole pine-dominated stands to stands dominated by white spruce (Table 7.15). Although considerable v a r i a t i o n i n the timi n g of t h i s change can be expected, the data suggest t h a t on t i l l , white spruce became more abundant than pine a f t e r 100-120 years. On l a c u s t r i n e s u b s t r a t e s , t h i s s h i f t was not so pronounced since both pine and spruce d i d not become common u n t i l a f t e r approximately 100 years. On these f i n e r t e xtured s o i l s , the pine stage was not always present and some deciduous stands went d i r e c t l y to white spruce. A t h i r d major change was the l a t e and gradual emergence of subalpine f i r as a member of the t r e e l a y e r . Table 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*'\u00E2\u0080\u00A2' Species Composition (%) Nominal Pinus Picea Abies Pseudotsuga stand oontovta glauea lasiocarpa menziesii Sample s i z e age Deciduous (lodgepole (white (subalpine (Douglas Substrate (yr.) spp.** pine) spruce) f i r ) f i r ) stems s i t e s TILL 10 100 3 3 25 30 63 7 40 2 45 3 97 428 5 75 94 6 54 2 110 5 40 43 10 2 456 10 135 4 32 44 14 154 4 150 33 57 8 1 72 3 200 42 44 12 1 148 4 LACUSTRINE 10 100 1 2 25 100 9 1 45 68 19 12 1 136 7 110 2 48 43 6 1 123 3 150 2 66 34 4 1 134 3 *Includes reconnaissance p l o t s . **Includes paper b i r c h , trembling aspen, black cottonwood and w i l l o w . * * * \" t \" i s l e s s than 1%. ho 00 Several workers p o s t u l a t e t h a t subalpine f i r would form the c l i m a t i c climax f o r the sub-boreal zone (Revel 19 72, I l l i n g w o r t h and A r l i d g e 1960, Rowe 1973). Swannell's (1913) comments a l s o suggest a climax dominated by subalpine f i r . However, since extensive w i l d f i r e s and logging have truncated the f u l l s u c c e s s i o n a l p a t t e r n , I was unable to loc a t e a subalpine fir-dominated stand w i t h i n the study area. Reasons f o r the coniferous species change were r e f l e c t e d i n the nature of t r e e regeneration (Table 7.16). On t i l l , lodgepole pine seedlings were abundant p r i m a r i l y i n the f i r s t 10-45 years. The occurrence of seedlings i n 45 year o l d stands was unusual and may r e f l e c t a h i s t o r y of ground f i r e s (van Barneveld, unpubl.). This type of f i r e can open the serotinous pine cones and provide s u f f i c i e n t s i t e disturbance f o r a s u i t a b l e seed bed. However, lodge-pole seedlings can a l s o e s t a b l i s h under an overstory of pine without f i r e ( J . Peek, pers. comm.). Whatever the reason f o r these s e e d l i n g s , i t i s obvious t h a t lodgepole g e n e r a l l y f a i l s to reproduce i t s e l f under a maturing f o r e s t canopy. Compared w i t h pine, white spruce seedlings occurred i n a l l age c l a s s e s except f o r one year (Table 7.16). In ol d e r stands, spruce regeneration drops sharply a f t e r peaking around 25 - 75 years. However, almost 10 percent of seedlings i n 200 year o l d stands were white spruce and thus t h i s species can maintain i t s e l f i n mature f o r e s t s . Table 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 SUBSTRATE Species Composition (%) ominal Pinus. Picea Abies Pseudotsuga Sample stand oontorta gtauca lasioaarpa menziesii age (lodgepole (white (subalpine (Douglas No. of No. of (yr.) pine) spruce) f i r ) f i r ) p l a n t s s i t e s 1 6 5 73 19 2 3 63 4 10 57 39 4 23 3 25 76 24 29 2 45 34 42 22 2 59 9 75 80 12 10 2 110 12 88 169 10 135 69 31 36 5 150 45 55 11 3 200 9 91 11 3 1 1 5 37 63 8 3 10 1 73 26 1 884 4 25 1 45 89 11 37 9 110 10 90 60 4 150 20 80 5 2 200 100 4 1 TILL LACUSTRINE K3 \u00E2\u0080\u00A2o 241 The eventual dominance of subalpine f i r i s shown by abundant regeneration i n most age c l a s s e s (Table 7.16). In p a r t i c u l a r , i t represented approximately 9 0 percent of the coniferous seedlings i n the o l d e s t stands on t i l l . The change from pine to spruce to subalpine f i r i s a n t i c i p a t e d and c l e a r l y supported by the species' regeneration data. On l a c u s t r i n e s u b s t r a t e s , lodgepole pine e s t a b l i s h e d very few s e e d l i n g s , while spruce and subalpine f i r were c l e a r l y much more s u c c e s s f u l . Pine plays a l e s s important r o l e on these substrates than on t i l l . The r e l a t i v e d i f f e r e n c e s between spruce and subalpine f i r regeneration p a r a l l e l l e d t h a t on t i l l s u b s t r a t e s : seedlings of spruce were most numerous between 10 - 45 years, and those of subalpine f i r were commonest i n l a t t e r stages (100 - 200 years) . Changing dominance of tree species was a l s o demonstrated by the species composition of dead trees (Table 7.17). Lodgepole pine was the most abundant dead species i n a l l age c l a s s e s sampled on both s u b s t r a t e s . The exception of the 150 year o l d stand on t i l l was probably due to sampling v a r i a t i o n . Dead spruce t r e e s occurred l a t e r i n succession on t i l l than on l a c u s t r i n e substrates because the species was uncommon i n e a r l y t i l l stage. S i m i l a r l y , dead subalpine f i r t r e e s were recorded for t i l l but not f o r l a c u s t r i n e s o i l s . The comparative lack of dead deciduous trees was notable on both s u b s t r a t e s . Table 7.17 Temporal Changes i n Composition and Propo r t i o n s of Dead Trees i n Mesic Sub-Boreal Forest Stands on T i l l and L a c u s t r i n e Substrates Substrate Nominal stand age (yr.) Deciduous spp. Species composition (%) Pinus contovta (lodgepole pine) Picea glauca (white spruce) Abies Zasiocavpa (subalpine f i r ) No. tre e s i n sample dead only l i v e and dead TILL 25 0 ( 0 % ) * 15 45 1 99 151 (28%) 549 75 100 1 (3%) 31 110 9 41 32 18 22 (11%) 203 135 5 62 29 5 21 (16%) 129 150 75 25 4 (15%) 27 200 3 92 5 37 (27%) 138 LACUSTRINE 25 100 1 (5%) 22 45 86 14 7 (20%) 35 110 63 37 19 (16%) 117 150 60 40 15 (9%) 159 Mean prop. (%) of dead tr e e s : t i l l 23% 1,092 l a c u s t r i n e 13% 333 ^Parentheses enclose p r o p o r t i o n of dead trees of t o t a l sample. P o s s i b l y deciduous species f e l l s h o r t l y a f t e r dying w h i l e c o n i f e r s remained standing f o r longer periods. This p o s s i b i l i t y would be true i f deciduous species died p r i m a r i l y from disease (e.g., root rot) and became s t r u c t u r a l l y unsound, and c o n i f e r s died p r i m a r i l y from l i g h t competition but remained sound. An i n t e r e s t i n g s t a t i s t i c was th a t the average p r o p o r t i o n of dead trees i n t i l l stands was almost double that i n l a c u s t r i n e stands (23% vs. 13%, r e s p e c t i v e l y ) . Stand parameters of b a s a l area, canopy c l o s u r e and height of dominant trees a l s o v a r i e d during succession as w e l l as changes i n species composition (Table 7.18). By age 2 25 years, stand b a s a l area was approximately 14 m /ha on 2 t i l l and 19 m /ha on l a c u s t r i n e s u b s t r a t e s . Stem area continued to increase r a p i d l y u n t i l about age 100-110, a f t e r which i t increased more slowly. The g e n e r a l l y b e t t e r growing c o n d i t i o n s on l a c u s t r i n e s o i l s was i l l u s t r a t e d by d i f f e r e n c e s i n bas a l area. At age 110, b a s a l area f o r 2 stands on t i l l averaged 32 m /ha while f o r l a c u s t r i n e 2 stands, the mean was 46 m /ha. At age 200 years, comparable 2 2 f i g u r e s were 37 m /ha and 44 m /ha. The s u c c e s s i o n a l trend i s i l l u s t r a t e d i n Figure 7.3. Height of the tre e l a y e r a l s o increased q u i c k l y up to approximately 100 years, although i n i t i a l development was slow. As i n d i c a t e d by heights of dominant t r e e s , the tre e l a y e r more than doubled i n height between age 25 and 45 Table 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 Canopy closure (%) Basal area (m2/ha) Nominal Ht (m) of stand c o n i f e r s a l l a l l prop. dominant Substrate age (yr) only trees trees c o n i f e r s tree (n) TILL 25 ( D * 14 20% 6.2 + 1.5** (5) 45 (5) 56 \u00C2\u00B1 8 56 \u00C2\u00B1 8** 19 \u00C2\u00B1 10** 99% 14.5 + 8.3 (9) 75 (1) 27 100% 20.0 + 5.9 (5) 110 (4) 67 \u00C2\u00B1 11 67 \u00C2\u00B1 13 32 \u00C2\u00B1 6 93% 27.9 + 4.4 (24) 135 (3) 33 \u00C2\u00B1 12 93% 26.3 + 3.5 (10) 150 (1) 35 100% 30.5 + 5.8 (5) 200 (5) 65 \u00C2\u00B1 8 65 \u00C2\u00B1 8 37 \u00C2\u00B1 4 99% 27.2 + 4.1 (25) LACUSTRINE 25 (1) 0 54 \u00C2\u00B1 36 19 0% 7.6 + 2.2 (2) 45 (1) 10 100% 14.4 + 2.7 (4) 110 (2) 75 \u00C2\u00B1 8 75 \u00C2\u00B1 8 46 \u00C2\u00B1 10 96% 24.1 + 1.6 (10) 150 (3) 66 \u00C2\u00B1 6 68 \u00C2\u00B1 8 44 \u00C2\u00B1 1 99% 26.8 + 3.6 (14) PARTIALLY LOGGED STANDS: TILL 135 (1) 17 84% 31.0 + 8.6 (4) 150 (2) 34 \u00C2\u00B1 15 44 \u00C2\u00B1 14 18 \u00C2\u00B1 2 94% 22.3 + 5.6 (9) 200 (1) 49 \u00C2\u00B1 9 54 \u00C2\u00B1 10 18 84% 16.3 + 8.7 (5) LACUSTRINE 150 (3) 15 I 4 88% 24.0 + 9.5 (15) *No. of s i t e s . **Mean \u00C2\u00B1 SD. to 244a Figure 7.3 Some t r e n d s . i n the f o r e s t stand features of b a s a l area, dominant tr e e height, and crown canopy closure 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. 245 246 years, and doubled again by age 110 years. Beyond t h i s time, height increases were minimal. On l a c u s t r i n e s u b s t r a t e s , the t r e e l a y e r demonstrated a s i m i l a r trend (Figure 7.3). P a r t i a l l y logged stands had lower tree l a y e r s and lower b a s a l areas than comparable unlogged f o r e s t s (Table 7.18). Gen e r a l l y , b a s a l areas were approximately one-half that of undisturbed s i t e s , and t r e e heights two to ten m l e s s . Of course, a more i n t e n s i v e sampling would r e v e a l wide v a r i a t i o n s i n the d i f f e r e n c e s between p a r t i a l l y logged and unlogged stands. P a r t i a l logging i s a broad d e s c r i p t i v e term that encompasses va r y i n g degrees of logging - the f u l l extent depending upon type of s e l e c t i o n , the stand's species composition, and d i s t r i b u t i o n of stem diameters. 7.3.4 Phytomass, Height and Basal Area of the Shrub Layer i n S e r a i Communities 7.3.4.1 For the combination of species. Phytomass of the Shrub (B) l a y e r v a r i e d markedly over the 200 year sere (Table 7.19). On t i l l s u b s t r a t e s , the phytomass of 2 shrubs > 45 cm t a l l was only 13 g/m a f t e r the f i r s t growing season. This increased n i n e - f o l d by the f i f t h season, and by 278 times by the t w e n t y - f i f t h season. Greatest phytomass 2 was a t t a i n e d at year 25 w i t h an estimated 3,616 g/m . By year 45, above-ground p l a n t m a t e r i a l had d e c l i n e d sharply to 247 Table 7.19 S t a t i s t i c s (mean \u00C2\u00B1 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 Shrub parameter (mean \u00C2\u00B1 sd) Substrate BA stem den. mean d i a . Nominal age wt (g/m2) ht (cm) (cm 2/m 2)** (no/m2) (cm) T i l l 1(15 ,10 ) * 13\u00C2\u00B1 36 73\u00C2\u00B113 0. 2\u00C2\u00B1 4 0. 7 \u00C2\u00B11 . 5 \u00E2\u0080\u00A2 0.5310.09 5(10) 120\u00C2\u00B1 29 85\u00C2\u00B129 2. 1\u00C2\u00B1 2. 0 4. 8 \u00C2\u00B1 4 . 1 0.7210.30 10 (5 , 1 ) 160\u00C2\u00B1 233 112\u00C2\u00B1 9 2. 6\u00C2\u00B1 3. 5 2. 7+2.6 0.8210.23 25(5) 361611767 361\u00C2\u00B148 29. 9\u00C2\u00B112 . 7 3. 1\u00C2\u00B11 . 2 3.1410.44 45(15 , 5 ) 102\u00C2\u00B1 205 78\u00C2\u00B141 1. 0\u00C2\u00B1 2. 0 1. 1 \u00C2\u00B1 1 . 3 0.7910.70 75 (5 ,1 ) 77\u00C2\u00B1 145 77125 0. \u00E2\u0080\u00A25\u00C2\u00B1 0. 7 1. 5 \u00C2\u00B1 2 . 0 0.5510.21 110 (20 ,1 ) 194\u00C2\u00B1 217 84131 2. 2\u00C2\u00B1 2. 1 2. 2 \u00C2\u00B11 . 8 1.0310.56 135 (10,1) 56\u00C2\u00B1 86 74\u00C2\u00B134 0. 6\u00C2\u00B1 0. 7 1. 1\u00C2\u00B10 . 7 0 .9810.77 150(5) 226\u00C2\u00B1 285 90\u00C2\u00B153 1. 9\u00C2\u00B1 2. 5 0. 9 \u00C2\u00B10 .7 1.3711.16 200(15 ,3 ) 340\u00C2\u00B1 825 86\u00C2\u00B151 3. 1\u00C2\u00B1 7. 1 1. 3\u00C2\u00B11 .6 0.9811.03 L a c u s t r i n e 1 ( 5 , 1 ) 24\u00C2\u00B1 24 89\u00C2\u00B1 9 0. 6\u00C2\u00B1 0. 6 1. 5\u00C2\u00B11 . 2 0.6110.11 5 (10) 88\u00C2\u00B1 104 82118 1. 1\u00C2\u00B1 1. 4 2. 8 \u00C2\u00B1 2 . 0 0.60+0.19 10 (10 , 1 ) 348\u00C2\u00B1 316 119146 5. 8\u00C2\u00B1 5. 1 7. 9 \u00C2\u00B1 8 . 0 0 .9810.49 25(5) 2461+2295 252184 16. 8 \u00C2\u00B1 1 1 . 9 3. 2 \u00C2\u00B1 1 . 1 1.9810.63 45(5 ) 83\u00C2\u00B1 170 93184 0. 8\u00C2\u00B1 1. 4 1. 1 \u00C2\u00B1 0 . 5 0.7010.66 110(10) 41\u00C2\u00B1 53 56112 0. 4\u00C2\u00B1 0. 4 1. 2 \u00C2\u00B10 . 9 0.52\u00C2\u00B10.23 150(15 ,1) 92\u00C2\u00B1 119 76\u00C2\u00B116 0. 8+ 0. 8 3. 1\u00C2\u00B12 .7 0.54+0.12 P a r t i a l l y logged - t i l l 135(5) 46\u00C2\u00B1 63 65\u00C2\u00B1 9 0. 7\u00C2\u00B1 0. 8 1. 9\u00C2\u00B11 . 9 0.6810.14 150(10) 276\u00C2\u00B1 369 91\u00C2\u00B120 3. 5\u00C2\u00B1 2. 9 3. L+1.6 1.0210.44 200(5) 302\u00C2\u00B1 509 118136 3. 0\u00C2\u00B1 4. 0 1. 7\u00C2\u00B11 .6 1.3610.90 P a r t i a l l y logged - : l a c u s t r i n e 150(15,2) 197\u00C2\u00B1 391 104155 2. 2\u00C2\u00B1 3. 7 1. 2\u00C2\u00B10 .7 1.1110.84 *(No. of per quadrats i n sample, no. of these w i t h zero v a l u e s ) . Height and stem diameters were based on non-zero per quadrats. **Derived from diameters measured at 10 cm above the root crown (DHW). 248 2 only 102 g/m or approximately 3 percent of the peak y i e l d . Phytomass remained r e l a t i v e l y l e v e l from year 45 to about 2 year 135 at an average of 10 7 g/m . In the l a t e r stages of \u00E2\u0080\u00A2the sere, standing crop g r a d u a l l y increased, reaching 2 340 g/m by the year 200. This represented t h r e e - f o l d increase over the middle stage of the sere. Thus the o v e r a l l p a t t e r n i n phytomass was an abrupt peak and d e c l i n e w i t h i n the f i r s t quarter of the sere, followed by an intermediate p l a t e a u and then a gradual increase. A s i m i l a r p a t t e r n occurred i n the Shrub l a y e r on l a c u s t r i n e substrates (Table 7.19). A f t e r one growing 2 season, phytomass was 2 4 g/m . I t increased r a p i d l y to a 2 peak of 2,461 g/m a f t e r 25 seasons, and then d e c l i n e d to 2 83 g/m by year 45, approximately 3 percent of the peak phytomass. Probably p l a n t m a t e r i a l increased g r a d u a l l y i n l a t e r stages of the sere, although no 200 year o l d stands were sampled on t h i s s u bstrate. Phytomass was u s u a l l y greater on t i l l than on l a c u s t r i n e s ubstrates. Except f o r year 1 and year 10, the former substrate had 123 percent to 473 percent higher phytomass than the l a t t e r . Changes i n mean height of the Shrub l a y e r followed changes i n phytomass except t h a t no increase occurred i n the l a t e r s e r a i stages (Table 7.19). A f t e r one growing season on t i l l s u b s t r a t e s , the Shrub l a y e r was 73 \u00C2\u00B1 13 cm i n height. This increased almost f i v e f o l d by year 25 to 361\u00C2\u00B148 cm. A f t e r 45 growing seasons, the Shrub l a y e r had returned to a l e v e l s i m i l a r to the e a r l i e s t stages of succession, 81\u00C2\u00B16 cm (n=6). V i r t u a l l y the same p a t t e r n and same mean heights occurred on l a c u s t r i n e substrates except that maximum height at year 25 was only 70 percent that on t i l l . The f a c t t h a t height of the Shrub l a y e r on both substrates was s i m i l a r but phytomass was d i s s i m i l a r suggested t h a t e i t h e r stem d e n s i t i e s or stem diameters were l e a s t on l a c u s t r i n e m a t e r i a l s . Comparison of these parameters i n d i c a t e d t h a t stems on the f i n e textured s o i l s tended to be smaller than those on the t i l l s o i l s (Table 7.19). Except f o r years 1 and 10, mean stem diameter of shrubs on t i l l s ubstrates exceeded those growing on l a c u s t r i n e substrates by 13 percent to 154 percent. Trends i n bas a l area on both l a c u s t r i n e and t i l l s u bstrates c l o s e l y p a r a l l e l l e d changes i n phytomass (Table 7.19). On t i l l , b a s a l area increased by 150 times from 2 2 0.2 cm /m a f t e r 25 growing seasons. S i m i l a r to the phytomass p a t t e r n , b a s a l area dropped to 3 percent of t h i s peak by year 45, and showed a gradual increase i n l a t e r s u c c e s s i o n a l stages. On l a c u s t r i n e substrates a s i m i l a r , though l e s s marked p a t t e r n occurred. Greatest b a s a l area occurred at year 25 when i t was 28 times that of year 1 2 2 2 2 (16.8 cm /m vs 0.6 cm /m ), and bas a l area d e c l i n e d to 5 percent of t h i s peak f i g u r e . 250 Stem d e n s i t i e s d i d not f o l l o w the above patterns shown by phytomass, height and b a s a l area (Table 7.19). On 2 t i l l s u b s t r a t e s , stem d e n s i t y increased from about 1/m 2 a f t e r one growing season to about 5/m a f t e r f i v e seasons. 2 Density d e c l i n e d to about 1/m by year 45 and then remained at approximately t h i s l e v e l f o r the duration of the sere. On l a c u s t r i n e s u b s t r a t e s , stem d e n s i t i e s were g e n e r a l l y 2 higher than on t i l l . The number of stems/m increased from 1.5 at year 1 to almost e i g h t by year 10, d e c l i n e d to one by year 45, and then p o s s i b l y increased i n the l a t e r s u c c e s s i o n a l stages. A l l shrub measures f o r the p a r t i a l l y logged stands were g e n e r a l l y greater than t h e i r unlogged counterparts (Table 7.19). On t i l l , stem d e n s i t i e s of p a r t i a l l y logged stands averaged 216% of the unlogged stands while phytomass, height and b a s a l area had analogous values of 97 percent, 108 percent and 133 percent, r e s p e c t i v e l y . Shrubs growing i n s e l e c t i v e l y logged stands probably have d i f f e r e n t growth forms than i n unlogged f o r e s t s since increases i n b a s a l area and stem d e n s i t y were not p a r a l l e l l e d by the phytomass data. Greater d i f f e r e n c e s were noted i n the p a r t i a l l y logged stands on l a c u s t r i n e s u b s t r a t e s . Here, increases f o r phytomass, height, b a s a l area and stem density were 214 per-cent, 137 percent, 275 percent and 387 percent, r e s p e c t i v e l y , of t h e i r unlogged counterparts. The foregoing data are presented by i n d i v i d u a l 251 species i n Appendix Tables F-9 to F-20. 7.3.4.2 Trends i n phytomass and height i n food species. S e r a i stages v a r i e d i n the proportions of food-producing species. These species were defined according to the food h a b i t s data of s e c t i o n 4. On t i l l , the p r o p o r t i o n of food species showed two J-shaped inc r e a s e s . The f i r s t extended from year 1 to year 25, when proportions went from 7 percent to 89 percent. The second extended from year 45 to years 150-200, when percentages went from 0 percent to 89 percent. Although the p r o p o r t i o n a l trends were s i m i l a r , absolute phytomass at peak's were considerably d i f f e r e n t : the 25 year o l d stage supported more than 10 times the p l a n t m a t e r i a l as the o l d e s t f o r e s t stage. P l a n t communities on l a c u s t r i n e substrates contained on average, a greater p r o p o r t i o n of food species phytomass than those on t i l l . This was because aspen, a food species, was a major p l a n t of e a r l y succession on l a c u s t r i n e s u b s t r a t e s , while e a r l y s e r a i species on t i l l s ubstrates were not forage species. In l a t e r stage of succession, food species c o n t r i b u t e d d e c l i n i n g amounts to the shrub l a y e r (Figure 7.4), u n l i k e the t i l l f o r e s t stands. P a r t i a l logging apparently increased food production on l a c u s t r i n e substrates but had l i t t l e e f f e c t on t i l l s u bstrates (Figure 7.4). In the unlogged stand on the former m a t e r i a l s , food species comprised 24 percent of the 251a Figure 7.4 Trends i n height and mass of browse and non-browse shrub 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 s u b s t r a t e s . eg CO CO < 225 H 150 75H m pi / r + 4\u00E2\u0080\u0094PL VALUE FOR PARTIALLY LOGGED STANDS \u00C2\u00AB BROWSE AND SHRUB MASS ARE EQUAL PL-\u00C2\u00BB P L - a 71 0 1 5 10 45 75 + + + ! + + \u00E2\u0080\u00A2 It + + + \u00C2\u00AB-PL + + + + + + + + + + + + + + + + h300 225 150 75 135 150 T I M E S I N C E D I S T U R B A N C E ( Y R ) 200 253 shrub biomass while on p a r t i a l l y \" logged stands, the corresponding p r o p o r t i o n was 96 percent. S i m i l a r data f o r the comparable t i l l stands were 89 percent and 91 percent, r e s p e c t i v e l y . The r e l a t i v e amounts of deciduous and coniferous browse a l s o v a r i e d over time and between substrates. On t i l l , c oniferous browse (subalpine f i r ) was v i r t u a l l y absent up to at l e a s t the 25 year stage and then made up approximately 9 6 percent of the browse i n l a t e r stages. On l a c u s t r i n e s o i l s , a s i m i l a r though l e s s pronounced p a t t e r n occurred. P a r t i a l logged reduced the pr o p o r t i o n of coniferous browse i n the shrub l a y e r . This r e s u l t e d more from an increase i n biomass of deciduous shrubs rather than i n biomass of subalpine f i r . Forage shrubs g e n e r a l l y were t a l l e r than mean heights f o r s e r a i stages. This feature was more evident i n the t i l l p l a n t communities than i n the l a c u s t r i n e ones. In the former ones, food species ranged from 101 percent to 172 percent of the mean height of the community, while f o r the l a t t e r , the range was from 73 percent to 360 percent. As a r u l e , the deciduous forage species were shade-i n t o l e r a n t and were able to su r v i v e by maintaining greater height than t h e i r competitors. Many of the npn-forage shrubs were shade-tolerant and were able to surv i v e these deciduous forage p l a n t s and p e r s i s t i n t o the f o r e s t communities. 254 7.3,5 Phytomass of the Herb Layer i n S e r a i P l a n t Communities In the Herb l a y e r , four major forage c l a s s e s were defined; graminoids, f o r b s , shrubs and other taxa. Graminoids i n c l u d e d grasses, sedges and rushes but not other monocots such as members of L i l i a c e a e and Iridaceae. Forbs i n c l u d e d non-graminoid monocots and a l l non-woody (herbaceous) d i c o t s . Shrubs i n c l u d e d woody species of both angiosperms and gymnosperms that were l e s s than 4 5 cm t a l l . P l a n t s grouped as \"others\" were f e r n s , h o r s e t a i l s and clubmosses. On t i l l , phytomass developed along t y p i c a l l i n e s (Figure 7.5). Forbs and graminoids dominated the f i r s t few years of succession, comprising up to approximately 60 per-cent of the estimated phytomass. However, proportions changed q u i c k l y so that by year 5, shrubs c o n s t i t u t e d the same percentage of the biomass while herbs made up only 44 percent. A f t e r the i n i t i a l year of the sere, graminoids remained a minor component of the Herb l a y e r , making up at most 3 percent of i t s mass (Table 7.20). By age 10, the Herb l a y e r had d e c l i n e d to i t s lowest phytomass. During the remaining p a r t of the sere, phytomass increased s l i g h t l y to 2 81 g/m at age 135 and then d e c l i n e d s l i g h t l y to about 2 62 g/m . During t h i s time (45 - 200 y e a r s ) , the p r o p o r t i o n of forbs was 38+9 percent and f o r shrubs 51+9 percent. The \"others\" c l a s s increased from v i r t u a l l y zero i n the e a r l i e s t 2 stages to approximately 16 g/m i n l a t t e r stages. F l o r i s t i c 254a Figure 7.5 Percentage composition, by forage c l a s s , of the phytomass i n the \"C\" or Herb l a y e r at four s u c c e s s i o n a l stages of the sub-bor 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 s u b s t r a t e s . 255 ^ GRAMINOIDS Q FORBS f g | SHRUBS f g j OTHER TAXA PRESENT IN TRACE AMOUNTS ( DC < I DC CL h-UJ Li_ o o h-DC o Q. 8 CL PRESENT IN TRACE AMOUNTS (o ; o o\u00C2\u00B0 \u00C2\u00B0 \u00C2\u00AB o o o ' oo VACCINIA SPIREA TWINFLOWER M A T U R E F O R E S T 263 thimbleberry. At the Shrub stage, while a l d e r s t i l l produced s i m i l a r amounts as before, i t s c o n t r i b u t i o n d e c l i n e d from 17 percent to 6 percent; s p i r e a was reduced to very low amounts. Willow c o n t r i b u t e d 87 percent of the y i e l d at the shrub stage, w i t h b i r c h p r o v i d i n g 5 percent. Over the next 30 years, w i l l o w e s s e n t i a l l y disappeared as a s i g n i f i c a n t species f o r browse production. I t s place was taken by a l d e r , rose, s p i r e a and twinflower. In the mature f o r e s t , these species were a l s o present except f o r a l d e r . S o a p a l a l l i e and v a c c i n i a were added and became the major shrub producers. Major shrubs were a l s o examined wi t h respect to changes i n proportions of leaves, c u r r e n t annual twigs and o l d e r twigs (Table 7.22 and Appendix Table F ) . Two broad temporal patterns were apparent. One group, c o n s i s t i n g of a l d e r , rose, s p i r e a and v a c c i n i a showed d e c l i n i n g p r o d u c t i v i t y over time. This decrease r e s u l t e d p r i m a r i l y from fewer leaves being produced. The second group, c o n s i s t i n g of black twinberry and thimbleberry, maintained a r e l a t i v e l y constant l e v e l of p r o d u c t i v i t y . A l s o , annual production represented a high p r o p o r t i o n of these species' phytomass, approximately 70 percent. Other shrub species were separated s i m i l a r l y but u s u a l l y only f o r one s u c c e s s i o n a l stage. These are summarized below: Table 7.22 Changes i n Proportions of P l a n t Components of Selected Shrub Species at Four Successional Stages on T i l l Substrates P r o p o r t i o n (%) of t o t a l phytomass Species Component Herb Shrub Immature Forest Forest Alnus spp. (alder) ann. leaves 38(3)* 10(8) 19(4) _ ann. twigs 3 2 5 -Loniceva involucrata (black twinberry) ann. leaves - 36(1) 47(1) -ann. twigs - 32 24 -Rosa spp. (rose) ann. leaves 66(3) 33(1) 35(6) 36(5) ann. twigs 10 25 33 11 Rubus parviftorus (thimbleberry) ann. leaves 42(1) 58(2) 57(3) -ann. twigs 29 18 15 -Spirea luoida (spirea) ann. leaves 74(8) 32(1) 43(9) 30(17) ann. twigs 5 36 5 8 Vacoinium spp. (v a c c i n i a ) ann. leaves 45(9) - 35(1) 15(20) ann. twigs 28 - 31 4 *No. of samples. 265 Douglas maple (Forest stage): 10% leaves, 1% current twigs s o a p a l a l l i e (Forest stage): 22% leaves, 9% current twigs squashberry (Immature F o r e s t ) : 23% leaves, 8% c u r r e n t twigs paper b i r c h (Shrub stage): 23% leaves, 3% current twigs trembling aspen (Shrub stage): 19% leaves, 4% current twigs w i l l o w (Shrub stage): 24% leaves, 3% current twigs These p l a n t s were a l i k e i n t h a t leaves formed approximately 20 percent of the p l a n t s , and current twigs formed about 5 percent. Thus some of the main winter food species of moose, e.g., w i l l o w and paper b i r c h , incremented woody t i s s u e by very small amounts. I f annual twig growth i s considered as a v a i l a b l e browse, then i t forms a small p r o p o r t i o n of the t o t a l amount present. 7.4 Results f o r R i p a r i a n P l a n t Communities Patterns of f o r e s t succession on a l l u v i a l , r i p a r i a n areas has been described p r e v i o u s l y i n t h i s province f o r the F o r t Nelson River (Waring 1970), and f o r the F i n l a y and Parsnip Rivers (Sumanik 1968). My f i e l d observations suggested t h a t these patterns a l s o occurred on s i m i l a r s i t e s i n the P r i n c e George study area, i . e . , on the McGregor and S t e l l a k o s o i l a s s o c i a t i o n s (cf. Table 2.2). As I d i d not study these r i p a r i a n communities i n d e t a i l , the f o l l o w i n g r e s u l t s present only major features of the s u c c e s s i o n a l sequence. The information draws h e a v i l y upon the two s t u d i e s mentioned above. 26.6 Succession proceeds through s i x stages (Table 7.23). I n i t i a l l y , w i l l o w s (Salix interior S.. maokenziana and others) e s t a b l i s h on the s i l t s deposited by the slower-moving cu r r e n t s . On the g r a v e l l y and u s u a l l y d r i e r s u b s t r a t e s , black cottonwood i s t y p i c a l l y the pioneer woody species. Shrubs of these two genera grow r a p i d l y i n height and form dense t h i c k e t s w i t h i n 10 years. By age 20, three l a y e r s have developed. The cottonwoods are up t o 12 m t a l l and 8 cm DBH; they form the developing Tree or A l a y e r . The will o w s g e n e r a l l y are approximately h a l f t h i s height and are the major occupant of the Shrub or B l a y e r . The Herb l a y e r i s u s u a l l y sparse and c o n s i s t s mostly of h o r s e t a i l s . At approximately age 60, a w e l l - d e f i n e d three layered f o r e s t e x i s t s . Cottonwood i s s t i l l the only species i n the Tree l a y e r , reaching 30-35 m i n height and 30-35 cm DBH. The Shrub l a y e r has few wil l o w s but rose and red-o s i e r dogwood are common. The Herb l a y e r has developed i n d i v e r s i t y w i t h h o r s e t a i l s , various Rubus spp., and bentgrass i s common, while spruce occurs at d e n s i t i e s of approximately 4-10/ha and at heights of l e s s than 1.5 m. (see Figure 7.8). A f t e r 150-200 years, the community begins to lose i t s deciduous character. While spruce becomes a minor member (10 percent to 40 percent) of the Tree l a y e r . Cottonwood i s s t i l l the most abundant tr e e species but i s u n t h r i f t y w i t h broken tops and r o t t e n cores. The Shrub l a y e r contains more shade t o l e r a n t species i n a d d i t i o n to Table 7.23 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 abitats (adapted from Sumanik (1968) and Waring (1970)) S e r a i stage Stand age ( y r ) * Major p l a n t species by l a y e r A (Tree) B (Shrub) C (Herb) sandbar w i l l o w cottonwood - w i l l o w - h o r s e t a i l cottonwood - rose - h o r s e t a i l cottonwood - dogwood - h o r s e t a i l cottonwood - a l d e r - h o r s e t a i l spruce-rose (dogwood) - h o r s e t a i l 1-5 6-20 50 150 200 >200 cottonwood ( 6 ) * * cottonwood (35) cottonwood (45) cottonwood (40), spruce (33) spruce (35) w i l l o w rose, dogwood, currant dogwood, a l d e r , squashberry, rose a l d e r , dogwood, rose, squash-berry rose, dogwood, alder h o r s e t a i l h o r s e t a i l h o r s e t a i l , Rubus spp.,, bentgrass h o r s e t a i l , s a r s a p a r i l l a h o r s e t a i l , wintergreen h o r s e t a i l , wintergreen * Approximate ages only. * * T y p i c a l t r e e heights i n m. ON \u00E2\u0080\u00A2=~4 267a Figure 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 ) s ubstrates. 269 those found i n the previous stage. These include squashberry, mountain a l d e r , black twinberry, currants and gooseberries. Only the f i r s t species i s an important winter food f o r moose. I f the s i t e i s not destroyed or modified through e r o s i o n , the f i n a l s u c c e s s i o n a l stage i s a mature spruce f o r e s t (Table 7.23). The Tree l a y e r contains only a few, p e r s i s t e n t , decaying cottonwood t r e e s . White spruce i s 35-40 m high and 50 cm DBH. A l s o , i t forms an intermediate t r e e l a y e r of from 3 - 12 m, which i t shares w i t h mountain a l d e r . The Shrub l a y e r c o n s i s t s mainly of rose, red-o s i e r dogwood, and mountain a l d e r , w i t h l e s s e r amounts of black twinberry, currants and gooseberries. In the Herb l a y e r , h o r s e t a i l s and wintergreen are the commonest species. Mosses form a w e l l developed carpet. 7.5 D i s c u s s i o n 7.5.1 P r e d i c t i n g Successional Development P r e d i c t a b i l i t y of s u c c e s s i o n a l patterns r e q u i r e s an understanding of the b a s i c f a c t o r s causing development of p l a n t communities. This i s an exceedingly d i f f i c u l t task due to the many environmental f a c t o r s i n v o l v e d , and t h e i r i n t e r a c t i o n s (Cormack 1953). Mueller-Dombois and E l l e n b e r g (1974) attempted to untangle t h i s complexity by proposing a model f o r p l a n t community formation: 270 p l a n t community = -f ( f , a, e, h, t , ) , where f = f l o r a a = a c c e s s i b i l i t y e = e c o l o g i c a l p l a n t p r o p e r t i e s h = h a b i t a t t = time As these authors noted, t h i s model d i f f e r s only i n emphasis from the one proposed by Major (1951). He subdivided h a b i t a t i n t o the three components of c l i m a t e , parent s o i l m a t e r i a l and r e l i e f , and grouped the above f a c t o r s of f, a, and e as \"organisms\". (Major's proposal was modified from the e a r l i e r s i t e f a c t o r approach of Jenny (1941)). Neither of these two models i s completely appropriate w i t h respect to moose and f o r e s t succession since they have not i d e n t i f i e d \"disturbance\" as a major f a c t o r . Mueller-Dombois and E l l e n b e r g (1974) in c l u d e i t as a component under t h e i r h a b i t a t f a c t o r . However, since moose are adapted to ecosystems where abrupt changes are the r u l e (Geist 19 71), disturbance must be i d e n t i f i e d as a major f a c t o r . The two above models.can be modified s l i g h t l y and expanded to i n c l u d e those f a c t o r s that appear meaningful f o r the purposes of moose h a b i t a t management. The hypothesized model i s as f o l l o w s : 271 major f a c t o r major components p l a n t community = h a b i t a t climate substrate r e l i e f or topography organisms f l o r a a c c e s s i b i l i t y f a c t o r e c o l o g i c a l p l a n t p r o p e r t i e s disturbance type (e.g. f i r e , logging) s e v e r i t y ( i n annual c y c l e and i n r e l a t i o n to major seed years) time rate of change community age I t should be noted t h a t \" h a b i t a t \" i s eq u i v a l e n t to the term \"environmental u n i t \" used i n t h i s t h e s i s . I f the model i s complete, than as the e f f e c t s of these f a c t o r s and t h e i r components on p l a n t community development are d e l i n e a t e d , then the p r e d i c t a b i l i t y of s u c c e s s i o n a l patterns becomes more l i k e l y . Hewlette (1976) noted that the f o l l o w i n g four f a c t o r s i n f l u e n c e d production and composition of understory vegetation a f t e r f o r e s t c u t t i n g i n eastern American hardwood stands: s i t e , stand type, stand s t r u c t u r e , and disturbance. These correspond g e n e r a l l y to the f i r s t three f a c t o r s i n the above model. development i n sub-boreal f o r e s t s , I examined the e f f e c t s of time and sub s t r a t e . These f a c t o r s were considered as most important i n the study area f o r moose h a b i t a t . Thus the p o t e n t i a l e f f e c t due to r e l i e f was pooled through the design As a f i r s t s t e p . i n t e s t i n g the model f o r community of the sampling system. The e f f e c t s of uncommon su b s t r a t e s , and the h y d r i c and x e r i c environments (climates) were not sampled. A l s o not s t u d i e d were e f f e c t s due to type of disturbance and d i f f e r e n c e s due to standards of logging methods. These subjects r e q u i r e f u r t h e r study. An e s s e n t i a l companion study would be a r a d i o - t e l e m e t r y study of moose h a b i t a t s e l e c t i o n . Information from t h i s l a t t e r type of study w i l l define the environmental tex t u r e to which moose r e l a t e . For example, e f f e c t s of aspect on drumlins may not be important to study i f moose do not d i s c r i m i n a t e between those p l a n t communities whose d i f f e r e n c e s are due to aspect. The e f f e c t of two major substrates on the sequence of p l a n t communities was examined through major s e r a i stages of sub-boreal f o r e s t s . These \"substrate\" e f f e c t s were presented i n s e c t i o n 7.3,2. In b r i e f , the s u c c e s s i o n a l p a t t e r n on mesic l a c u s t r i n e substrates was from aspen and w i l l o w , to aspen, and then to pi n e , spruce and subalpine f i r . I observed some cases where pine was omitted from the sere, so t h a t the sequence was from aspen to white spruce. The presence or absence of pine l i k e l y depends upon the s e v e r i t y of the disturbance (whether or not aspen i s killed.) , and the a v a i l a b i l i t y of seed. On the mesic t i l l s u b s t r a t e s , the sequence was from raspberry and s p i r e a to w i l l o w , and then to pin e , white spruce and e v e n t u a l l y subalpine f i r . I t appeared that mesic l a c u s t r i n e communities produced a more 273 diverse and prolonged deciduous phase than mesic t i l l communities. Thus the former.substrate i s more valuable i n terms of food production than the l a t t e r . Since l a c u s t r i n e m a t e r i a l s mostly occur at low e l e v a t i o n i n n o r t h - c e n t r a l B r i t i s h Columbia, they must be viewed as important components of moose winter ranges. The e f f e c t of a t h i r d s u b s t r a t e , recent a l l u v i u m , was b r i e f l y considered. These f l o o d p l a i n s support c r i t i c a l moose winter h a b i t a t s . Moisture regimes were not d e l i n e a t e d w i t h i n t h i s substrate since water supply was determined mainly by ground sources r a t h e r than by r a i n and snow. Successional development appeared g e n e r a l l y s i m i l a r to e a r l i e r work by Sumanik (1968) and Waring (1970). This type of succession has a l s o been examined on the Peace River ( J e f f r e y 1961), i n Alaska (Viereck 1970), and i n Wyoming (Houston 1968). Flood p l a i n vegetation i s dynamic due to the combined e f f e c t s of f l o o d i n g and e r o s i o n . Odum (1971) r e f e r r e d to these ecosystems as p u l s e - s t a b i l i z e d . The vegetation i s a l s o heterogeneous w i t h cover-producing patches of white spruce o c c u r r i n g next to s e r a i shrub stages w i t h abundant food. Peek (19 74a) r e l a t e d the importance of these and other r i p a r i a n h a b i t a t s to Shir a s moose and provided other references to these types of communities. On t h i s s u b s t r a t e , the type of v e g e t a t i o n a l development i s s t r o n g l y i n f l u e n c e d by the long periods that the s o i l s are water saturated. This environment has favored those p l a n t species t h a t have high water r e q u i r e -ments, or t h a t are adapted to wet c o n d i t i o n s . Thus, cotton-wood develops moreso than aspen, and white spruce moreso than lodgepole pine. Because of the d i f f e r i n g requirements of these two bree species, succession has a long deciduous phase on alluvium. The e f f e c t of a f o u r t h s u b s t r a t e , coarse outwash, can be seen i n the r e s u l t s of C o t i c et a l . (1974). In mesic environments, t h i s substrate i n c l u d e s the Giscome, Peta and Saxton s o i l a s s o c i a t i o n s . (Table 2.2). These s o i l s are coarse t e x t u r e d and as such, are q u i t e permeable and d r a i n r a p i d l y . These features create a \"surface d r o u g h t i -ness,\" the main c h a r a c t e r i s t i c of these s o i l s ( Cotic e t a l . 19 74). The moisture shortage has the e f f e c t on vegetation of f a v o r i n g development of lodgepole pine f o r e s t s that have a sparse understory (cf Revel 1972). Thus substrate i s a s i g n i f i c a n t f a c t o r that i n f l u e n c e s s u c c e s s i o n a l patterns i n sub-boreal f o r e s t s . The i d e n t i f i c a t i o n and separation of environmental u n i t s was r e a l i s t i c . While the e f f e c t i s most n o t i c e a b l e f o r the Tree l a y e r , i t a l s o has an important e f f e c t on the patterns i n the understory vegetation. Time was the second major f a c t o r examined. The two main components of the time f a c t o r are the i n t e r r e l a t e d community age and rate of change. Kemper (19 71) noted that determining the age of a stand i s not always a simple 275 procedure. Sometimes the disturbance that leads to the' establishment of a f o r e s t stand occurs many years previous to the beginning of the stand. In these s i t u a t i o n s i t would seem best to d i s t i n g u i s h s u c c e s s i o n a l or community age from the age of a tre e stand. D e f i n i n g community age becomes d i f f i c u l t when s e v e r a l events only p a r t l y d i s t u r b the stand. A good example of t h i s i s the p a r t i a l logging of sub-boreal f o r e s t s . In these cases, i t i s probably most info r m a t i v e to provide ages f o r communities developing a f t e r each disturbance. The importance of time i s obvious. In the Pr i n c e George study, i t appeared that s u c c e s s i o n a l communities were most d i f f e r e n t i n e a r l y stages, but tended t o converge i n l a t e r f o r e s t stages. These e a r l y stages were of great importance f o r food production, but of l i t t l e consequence f o r cover production. A l s o , the rat e of change from deciduous t o coniferous stands d i f f e r e d between substr a t e s . Much necessary and u s e f u l research remains to be done to t e s t the hypothesized model. For the \" h a b i t a t \" f a c t o r , the i n f l u e n c e of climate and r e l i e f r e q u i r e c h a r a c t e r i z a t i o n . The \"organisms\" f a c t o r has been t r e a t e d only l i g h t l y . Such r e l a t i o n s h i p s as t h a t between aspen stem d e n s i t i e s before and a f t e r logging are obvious subjects f o r study. The elegant and w i t t y papers by Horn (1974, 1975a, 19 75b) on secondary succession provide valuable i n s i g h t s and suggestions regarding the i n f l u e n c e of cu r r e n t stand 276 composition on succeeding s u c c e s s i o n a l stages. F i n a l l y , the \"disturbance\" f a c t o r should be st u d i e d , p r i m a r i l y i n r e l a t i o n to logging methods and standards of harve s t i n g . N a t u r a l disturbances should not be overlooked, however, as they are experimental treatments t h a t w i l l not be otherwise conducted. A recent paper by Nordin and G r i g a l (1976) described r e l a t i o n s h i p s between v e g e t a t i o n , landscape f e a t u r e s , and s o i l w i t h i n a Minnesotan w i l d f i r e . This type of research provides u s e f u l i n s i g h t i n t o f a c t o r s l e a d i n g to formation of p l a n t communities. 7.5.2 Trends i n Production of Food and Cover Before d i s c u s s i n g the r e l a t i o n s between food and cover, and f o r e s t succession, s e v e r a l major p o i n t s should be considered. F i r s t , b o r e a l coniferous f o r e s t s have the l e a s t phytomass of the world f o r e s t s . Based on var i o u s sources, Whittaker (19 70) estimated that mean phytomass f o r these 2 f o r e s t s was 20,000 g/m , w i t h a normal range of 6,000 -2 40,000 g/m . This compares w i t h mean estimates f o r 2 1 temperate and t r o p i c a l f o r e s t s of 30,000 g/m and 45,000 g/m'' r e s p e c t i v e l y . As Satoo (1970) noted, mass i s a f u n c t i o n of age since stem phytomass, which i s accumulated annually, c o n s t i t u t e s an i n c r e a s i n g l y greater p a r t of the t o t a l community mass. Thus most estimates of f o r e s t biomass are probably s l i g h t l y conservative since few b o r e a l f o r e s t s sampled would be at a stage of s t a b l e phytomass. Nonethe-277 l e s s , the r e l a t i v e rank of b o r e a l f o r e s t s remains low. A second general p o i n t i s t h a t the understory vegetation t y p i c a l l y comprises a very small f r a c t i o n of the f o r e s t phytomass, at l e a s t i n mature stands. In four types of 50-60 year o l d deciduous f o r e s t stands i n Minnesota, Zavitowski (19 76) found t h a t understory phytomass 2 c o n s t i t u t e d l e s s than one percent (38-117 g/m ) of the t o t a l f o r e s t mass. In a near-climax mixed-wood stand i n Nova S c o t i a , understory v a s c u l a r p l a n t s comprised an estimated 2 238 g/m or 2.4% of the stand's t o t a l above ground phytomass ( T e l f e r 19 72). T e l f e r added that h i s estimated understory phytomass was greater than most of the 52 s t u d i e s summarized by Ovington (1962, Table 1). In the Pri n c e George area, understory phytomass f o r f o r e s t stands aged 110 to 200 years 2 on t i l l and l a c u s t r i n e s u b s t r a t e s , averaged 228 g/m (Tables 7.19 and 7.20). This estimate agrees w e l l w i t h T e l f e r ' s (19 72), and suggests that sub-boreal understory vegetation a l s o c o n t r i b u t e s l i t t l e to the stand's phytomass. The f i n a l general p o i n t i s that understory phytomass and production increase and decrease sharply i n the e a r l y stages of f o r e s t succession. This temporal aspect has received comparatively l i t t l e study i n sub-boreal and b o r e a l f o r e s t s . In the Pri n c e George f o r e s t s , net production was 97, 133, 18, and 27 g/m2 i n 1, 11, 39, and 195 year o l d f o r e s t s , r e s p e c t i v e l y . In a white spruce f o r e s t sere i n c e n t r a l A l b e r t a , t o t a l shrub production was 11, 97, 161, 191 278 2 and 59 g/m i n s c a r i f i e d cutovers that were 1, 5, 9, 17 years o l d , and i n mature stands, r e s p e c t i v e l y ( S t e l f o x e t a l . 1976). Since herbage was not measured, these l a t t e r data underestimate production, e s p e c i a l l y i n the f i r s t three ages. As w i t h phytomass, net production of the understory makes up a small p r o p o r t i o n of the t o t a l stand above ground production. Whittaker (19 70) estimated t o t a l production i n 2 b o r e a l f o r e s t s at 800 g/m with.a normal range of 400-2 2000 g/m / y r . Thus understory production c o n t r i b u t e s about 3 percent and 7 percent of the t o t a l f o r P r i n c e George and A l b e r t a f o r e s t s , r e s p e c t i v e l y . The s u c c e s s i o n a l changes i n production of food f o l l o w the general p a t t e r n shown by the understory. However, the absolute amounts w i l l be l e s s since not a l l of the understory can be considered as food. A l s o , not a l l of the food can be considered as a v a i l a b l e . The a v a i l a b i l i t y of food changes i n response to f a c t o r s that vary both w i t h i n and between years. The i n f l u e n c e s of many of these f a c t o r s are s t i l l unclear. Thus a dynamic a p p r e c i a t i o n of food a v a i l a b i l i t y i s u s u a l l y not p o s s i b l e from data i n most w i l d l i f e - s u c c e s s i o n s t u d i e s . Approaches t o g a i n i n g a b e t t e r understanding of food a v a i l a b i l i t y are described by Moen (1973). One important aspect i s to describe forage (= browse f o r moose i n winter) i n terms of components, by height i n t e r v a l s . For example, how does the phytomass of twigs l e s s than one cm i n diameter 279 and between 0 and 0.5 m above the ground vary during a f o r e s t sere. Information of t h i s type i s e s s e n t i a l l y non-e x i s t e n t i n the w i l d l i f e l i t e r a t u r e . Relevant s t u d i e s can be found i n s t u d i e s on f o r e s t p r o d u c t i v i t y , n u t r i e n t c y c l i n g and f u e l d e s c r i p t i o n s f o r f i r e c o n t r o l . For example, Sando and Wick (19 72) describe a method f o r e v a l u a t i n g crown f u e l s i n f o r e s t stands by height increments. D i s t r i b u t i o n of mass of t r e e components has been examined f o r lodgepole pine by Johnstone (1967), Gary (1976) and others. Fuel s t u d i e s on lodgepole pine include those by K i i l (1968) and Muraro (1971). Other u s e f u l s t u d i e s i n c l u d e those on aspen by B e l l a (1968), and on Douglas f i r by Kurucz (1969). The r e l a t i o n s h i p s between parameters of overstory density and understory phytomass have been examined by many researchers (see s e c t i o n 10.2.4 f o r d e t a i l s and r e f e r e n c e s ) . I explored s e v e r a l of these w i t h the Pri n c e George data. C o r r e l a t i o n between b a s a l area or canopy closure and Shrub la y e r phytomass or Shrub and Herb l a y e r phytomass were g e n e r a l l y i n s i g n i f i c a n t . The one exception was the r e l a -t i o n s h i p between Shrub l a y e r phytomass on t i l l s ubstrates and the overstory b a s a l area. The equation was: y = 9642 - 2733 l o g x, where n = 7, r 2 = 0.58, S = 934 r ' y. x and F = 6.77. (The tabu l a t e d value of F at the 95 percent p r o b a b i l i t y l e v e l f o r 1 and 5 df was 5.99). This agrees w i t h the c o r r e l a t i o n described by T e l f e r (1972). 280 Trends i n production of cover are more d i f f i c u l t t o p r e d i c t because the term \"cover\" has s e v e r a l , d i f f e r e n t meanings. For example, moose need escape cover from predators, s h e l t e r cover from d i f f e r e n t c l i m a t i c elements, and p o s s i b l y h i d i n g cover from other moose ( i . e . , space). The types of vegetation that can meet these needs probably v a r i e s w i t h the type of cover r e q u i r e d by moose. Vegetation probably begins to provide h i d i n g and escape cover once i t exceeds the standing height of a moose, assuming an adequate stand d e n s i t y . Thus the lower l i m i t of height of vegetation can be p r e d i c t e d to be between two and three m, since the mean shoulder height of an a d u l t b u l l moose i s approximately 1.8 t o 2.0 m ( B a n f i e l d 19 74, Cowan and Guiget 1973). The stage at which vegetation provides s h e l t e r cover probably v a r i e s w i t h the elements of c l i m a t e . For example, vegetation below 3.0 m can e f f e c t i v e l y reduce wind but have l i t t l e e f f e c t on i n t e r c e p t i n g snow or on reducing radia.tional\"heat l o s s . In the P r i n c e George study area, snow depth was assumed to be the most c r i t i c a l element. Thus tree crown canopy c l o s u r e r a t h e r than height (Peek e t a l . 1976) i s the most important stand feature to\"examine. Discussion of t h i s aspect i s deferred to S e ction 9. The type of vegetation f o r escape and s e c u r i t y probably d i f f e r s between summer and w i n t e r . Deciduous species of s u i t a b l e height are probably adequate i n summer 281 but may be inadequate i n wi n t e r a f t e r l e a f f a l l , e s p e c i a l l y i f stem d e n s i t i e s are low. Assuming that a 1.5 m t h r e s h o l d a p p l i e s to the P r i n c e George area i n summer, t i l l s ubstrates probably provide summer \" s e c u r i t y \" cover between 5 and 10 years a f t e r logging. At age 5, w i l l o w s were 1.8 m and s e v e r a l other deciduous species exceeded 1.0 m. At age 10, lodgepole pine was 1.4 m and w i l l o w 1.2 m. S i m i l a r data a p p l i e d f o r l a c u s t r i n e substrates so i t appears that these two parent m a t e r i a l s do not s u b s t a n t i a l l y i n f l u e n c e age at which summer cover becomes adequate. The l i m i t e d data i n Waring (19 70) suggest that on a l l u v i a l s u b s t r a t e s , adequate height i s a t t a i n e d by age 5. S t e l f o x e t a l . (1976) a l s o provided some data on summer cover. Five years a f t e r c l e a r c u t t i n g a white spruce f o r e s t , apparently l i t t l e summer (s e c u r i t y ) cover was provided w i t h poplar and w i l l o w at stem d e n s i t i e s of 13,000/ha and a mean height of 0.7 m. A f t e r 9 years, when these species averaged 14,00 0 stems/ha and 1.5 m i n height, cover appeared adequate. S i m i l a r l y , a f t e r 17 years, when 'the height remained almost the same but stem d e n s i t i e s were approximately 10,00 0/ha, moose used the cutover i n summer. (During t h i s 12 year p e r i o d , however, winter d e n s i t i e s were zero except when mature timber was nearby). Winter s e c u r i t y cover i s a f f e c t e d by s e v e r a l major f a c t o r s . T i l l substrates provided w i n t e r s e c u r i t y cover sooner than l a c u s t r i n e substrates since c o n i f e r s t y p i c a l l y 282 e s t a b l i s h e d more r e a d i l y on the former parent m a t e r i a l . Height growth a l s o v a r i e s between substrates. E i s (1966) presented height growth increments of three and four year o l d white spruce w i l d l i n g s in situ. He e s t a b l i s h e d t h a t growth v a r i e d between the three substrates of sand, loam and c l a y . Two years of height increment were 8.9, 7.0 and 5.4 cm, r e s p e c t i v e l y , f o r these subst r a t e s . In a subsequent r e p o r t , E i s (19 67) presented f u r t h e r data that allowed determination of when the \"best\" open grown white spruce reached 1.5 m. Ages at which 1.5 m were reached f o r the above substrates were 13, 17 and 21 years, r e s p e c t i v e l y . Pogue (1949) examined age-height r e l a t i o n s h i p s f o r white spruce and subalpine f i r i n r e s i d u a l stands at the Aleza Lake P r o v i n c i a l Forest. Spruce achieved 3 m at 20-21 years while subalpine f i r r e q u i r e d approximately 30 years. Stem diameters at breast height were between 2.5 and 5 cm. In an 18 year o l d burn, Pogue (1949) recorded t h a t white spruce were 3 m t a l l . My data were s i m i l a r . On t i l l , the shrub l a y e r reached 3 m at an estimated 20-21 years but on l a c u s t r i n e substrates reached only 2.5 m at 25 years. Stanek (1966) explored the i n f l u e n c e of various f a c t o r s on height growth of lodgepole pine and Engelmann spruce. He showed that i n i t i a l growth of spruce i s l e s s than that of pine, and t h a t height growth was a f f e c t e d by the p l a n t a s s o c i a t i o n , s i t e and competitive p o s i t i o n of the tr e e . For the dominant and co-dominant t r e e s , the case most 283 l i k e regeneration a f t e r c l e a r c u t t i n g or w i l d f i r e , Stanek determined t h a t pine reaches 3 m at 8, 13, 17 and 35 years on good, medium, poor, and low s i t e s . S i m i l a r ages f o r spruce were 12, 21, 27 and 50, r e s p e c t i v e l y (Stanek 1966:99). Ages to 3 m were a l s o provided on an a s s o c i a t i o n or f o r e s t type b a s i s . Results from Stanek's and my s t u d i e s can be used to determine at what ages regenerating cutovers begin to provide adequate escape, s e c u r i t y and s h e l t e r cover (except f o r snow i n t e r c e p t i o n ) . The important d e c i s i o n t h a t f i r s t must be made i s what k i n d of cover i s the cutover to provide. Once t h i s i s decided, the appropriate height can be determined and the time p e r i o d estimated when a regenerating stand begins to provide the d e s i r e d type of cover. Height browth has been given f o r other moose h a b i t a t s . P l a n t e d and volunteer coniferous species i n northeastern Minnesota a t t a i n e d 3 m at 9-11 years f o r red pine (Pinus resinosa) , jack pine (P. banksiana) and black spruce, and 21 years f o r white spruce (Peek e t a l . 1976). In the same study, average height of the understory ranged from 0.6 to 1.7 m i n coniferous and deciduous stands ranging from 1 t o 80 years. Height growth was t h e r e f o r e l e s s than I documented f o r P r i n c e George ranges. In northwestern A l b e r t a , Corns and La Roi (1976) recorded the mean heights of lodgepole i n 6- and 12-year-old stands at 0.21 and 284 0.91 m, r e s p e c t i v e l y ; corresponding values f o r aspen were 0.97 and 1.73 m. 8. NUTRITIVE ASPECTS OF MOOSE FORAGES 8.1 I n t r o d u c t i o n The purposes f o r examining n u t r i t i v e parameters of moose forages were as f o l l o w s ( p a r t l y derived from Oldenmeyer 19 74): 1) to provide a b a s i c n u t r i t i o n a l d e s c r i p t i o n of range forage, i n c l u d i n g both species and seasonal d i f f e r e n c e s , 2) to determine how n u t r i t i v e l e v e l s v a r i e d w i t h succes-s i o n a l stage, 3) to help determine why the d i e t s and h a b i t a t u t i l i z a t i o n of moose vary over time and space, 4) to determine the e f f e c t s on forage n u t r i e n t s of some major environmental f a c t o r s , p a r t i c u l a r l y o v e r s t o r y , s u b s t r a t e , and c l i m a t e , 5) to evaluate e f f e c t s of land use p r a c t i c e s and h a b i t a t manipulation on forage n u t r i e n t s . Many types of a n a l y t i c a l procedures e x i s t to determine the n u t r i t i o n a l value of forages f o r animals (e.g., H a r r i s 1970). The choice of techniques depends upon the o b j e c t i v e s of the study, and the time and money a v a i l a b l e f o r conducting the analyses. For the purposes of t h i s study, I s e l e c t e d crude p r o t e i n and l i g n i n . I b e l i e v e d that these parameters would provide s a t i s f a c t o r y data to 285 286 meet the f i v e o b j e c t i v e s l i s t e d above. The importance of p r o t e i n f o r ruminants such as moose was summarized s u c c i n c t l y as f o l l o w s by S i n c l a i r (1974:292): For ruminants the q u a l i t y of the food i n terms of p r o t e i n i s of p a r t i c u l a r importance. The r a t e of passage of food through the rumen i s determined by the s i z e of the food p a r t i c l e s . These p a r t i c l e s must reach a small enough s i z e before continuing through the gut. The lower the p r o t e i n value of the food, the higher the f i b r e content (Glover, Duthie & French, 1957) and the longer i s the time needed to reduce the food by m a s t i c a t i o n to the r i g h t s i z e . This was confirmed by e m p i r i c a l observations of c a t t l e (Hancock, 1954: Campling, Freer & Balch, 1961). Because of t h i s , there i s a l e v e l of p r o t e i n content i n the food below which i t becomes uneconomical to d i g e s t ; the energy expended i s too high, and the r a t e of n i t r o g e n intake i s l e s s than the r a t e of n i t r o g e n e x c r e t i o n . The d e f i c i t cannot be compensated f o r by i n c r e a s i n g the q u a n t i t y of food eaten because t h i s i s l i m i t e d by the r a t e at which the animal i s p h y s i c a l l y capable of m a s t i c a t i n g the food to the r i g h t s i z e ( B e l l , 1969). The process i s complicated f u r t h e r by the f a c t t h a t the a c t i o n of the rumen micro-organisms i s i n h i b i t e d at low p r o t e i n l e v e l s (Chalmers, 1961), and t h i s causes low energy a s s i m i l a t i o n ( M i l f o r d & Minson, 1966.) Hence there i s a minimum q u a l i t y of food below which the animal loses weight as i t u t i l i z e s i t s own f a t s u p p l i e s . The minimum maintenance requirement, then, i s the lowest food q u a l i t y at which the body weight of the animal remains s t a t i o n a r y . I f the ingested food q u a l i t y f a l l s below t h i s minimum maintenance l e v e l , then there i s evidence of food shortage. I f the q u a l i t y of the food i s above t h i s l e v e l then measurements on the q u a n t i t y of the food are necessary to obtain evidence of food l i m i t a t i o n . Crude p r o t e i n i n forages can be used to i n d i c a t e d i g e s t i b l e amounts of p r o t e i n i n the d i e t (Robbins 19 73:141-142). L i g n i n was used to i n d i c a t e forage d i g e s t i b i l i t y . D i g e s t i b i l i t y i s a fundamental f a c t o r i n n u t r i t i o n since i t 287 defines the p r o p o r t i o n of ingested m a t e r i a l t h a t the body can u t i l i z e (Maynard and L o o s l i 1969). L i g n i f i c a t i o n of maturing forage i s considered the major reason f o r d e c l i n i n g d i g e s t i b i l i t y (Morrison 19 72a). Using the a c e t y l bromide technique f o r d i s s o l v i n g l i g n i n y i e l d s data w e l l c o r r e l a t e d w i t h in vitro d i g e s t i b i l i t i e s of grasses and legumes (Morrison 1972a, b). The assumption t h a t l i g n i n values f o r deciduous and coniferous species were s i m i l a r l y r e l a t e d to d i g e s t i b i l i t y was not t e s t e d . However, despite the chemical d i s t i n c t n e s s of grass and wood l i g n i n s , when the same a n a l y t i c a l procedure i s used, then the u l t r a v i o l e t spectra of the a c e t y l bromide r e a c t i o n products are s i m i l a r w i t h respect to the maximum absorption peak (Morrison 19 72a). Ten p l a n t species were s e l e c t e d f o r study, c o n s i s t i n g of one c o n i f e r , e i g h t deciduous shrubs and one l i c h e n . T h e i r s e l e c t i o n was based on t h e i r abundance (importance) i n the d i e t of moose i n the study area, e s p e c i a l l y i n w i n t e r . I attempted to analyze both crude p r o t e i n and l i g n i n f o r 12 consecutive months f o r most species from most c o l l e c t i n g s i t e s . Thus, p r o t e i n l e v e l s were determined f o r a l l ten species and l i g n i n l e v e l s f o r s i x species: 288 Species Crude P r o t e i n L i g n i n Subalpine f i r Yes Yes Saskatoon Yes Paper b i r c h Yes Yes Red-osier dogwood Yes Yes Lungwort Yes Trembling aspen Yes Yes Black cottonwood Yes Willow Yes Yes Mountain ash Yes Yes Mountain b i l b e r r y Yes The sampling p e r i o d extended from A p r i l 19 72 through to A p r i l 1973. A t o t a l of f i f t e e n c o l l e c t i n g s i t e s were e s t a b l i s h e d (Table 8.1). A l l r e s u l t s of the p r o t e i n and l i g n i n analyses are given i n Appendix G, Tables G-l and G-2. 8.2 Methods Samples were c o l l e c t e d i n a c o n s i s t e n t fashion at mid-monthly i n t e r v a l s . Each sample c o n s i s t e d of approxi-mately 200-250 g wet-weight of p l a n t m a t e r i a l c l i p p e d from at l e a s t ten p l a n t s per species. This procedure minimized p o s s i b l e d i f f e r e n c e s between p l a n t s i n l e v e l s of p r o t e i n and l i g n i n . Only current annual growth was c o l l e c t e d , except f o r the l i c h e n , lungwort, which d i d not e x h i b i t d i s t i n g u i s h a b l e annual production. For t h i s species, m a t e r i a l was t o r n from branches t a k i n g care not to include the substrate t r e e ' s bark nor other l i c h e n species. Most samples were then oven-dried at 50\u00C2\u00B0C f o r 24 h. 289 Table 8.1 Location, Habitat, Substrate, and S p e c i e s C o l l e c t e d f o r Crude P r o t e i n and Lignin Analyses STUDY AREAS Site Habitat Species collected for Substrate chemical analysis EAGLE (750 \u00C2\u00B1 75 m) E l AO yr old birch forest lacustrine E2 40 yr old shrub sere lacustrine E3 100 yr old spruce- lacustrine Gubalpine f i r forest E4 250 yr old spruce- lacustrine subalpine f i r forest GROVE (750 \u00C2\u00B1 75 m) B l 12 yr old shrub sere t i l l B2 50 yr old lodgepole t i l l pine forest B3 50 yr old lodgepole t i l l pine forest Gl 12 yr old shrub sere t i l l G2 12 y r old shrub sere lacustrine G3 120 yr old lodgepole \" t i l l pine forest G4 200 yr old spruce- lacustrine subalpine f i r forest SALMON (750 \u00C2\u00B1 75 m) 51 spruce-black recent cottonwood forest alluvium 52 mixed wood forest beach deposits 53 trembling aspen beach forest deposits S5 120 yr o l d lodgepole t i l l pine forest saskatoon, paper birch, red-osier dogwood, willow, mountain ash, vaccinia saskatoon, paper birch, red-osier dogwood, trembling aspen, willow, mountain ash subalpine forest, lungwort subalpine f i r paper b i r c h , trembling aspen, black cottonwood, v/illow willow paper bi r c h , willow paper birch, black cottonwood, willow paper birch, trembling aspen, willow subalpine f i r , black cottonwood subalpine f i r , red-osier dogwood, lungwort red-osier dogwood paper birch, willow willow subalpine f i r , willow 290 Drying at higher temperatures than t h i s may increase l i g n i n a r t i f i c i a l l y (Goering and Van Soest 1967). For s e l e c t e d speci e s , some monthly samples were f i r s t separated i n t o l e a f and stem components before d r y i n g . Dried components were weighed so th a t whole sample estimates of p r o t e i n and l i g n i n were determinable. For example, the percent n i t r o g e n f o r a sample of stem and leaves combined was estimated as f o l l o w s : (% N i n leaves) (wt of leaves i n g) x ^ QQ$. (% N i n stems) (wt of stems i n g) A l l d r i e d p l a n t samples were ground i n i t i a l l y using a hammer m i l l , and then a Wiley m i l l equipped w i t h a 1 mm mesh s t a i n l e s s s t e e l screen. Subsequently, t h i s c o a r s e l y ground m a t e r i a l was mixed thoroughly. A sub-sample was withdrawn, ground i n a Wiley m i l l w i t h a 4 0 mesh screen and stored i n a l a b e l l e d p l a s t i c bag u n t i l a n a l y s i s . D i g e s t i b i l i t y determinations were based on procedures described by Johnson e t a l . (1964) and Morrison (1972a, b). The former authors o u t l i n e l i g n i n determina-t i o n s f o r woody t i s s u e s , while the l a t t e r one r e l a t e d l i g n i n determination to d i g e s t i b i l i t i e s f o r grasses and legumes. I assumed a s i m i l a r l y r e l i a b l e c o r r e l a t i o n between l i g n i n content i n wood samples and t h e i r d i g e s t i b i l i t y by moose. L i g n i n l e v e l s are i n v e r s e l y r e l a t e d to d i g e s t i b i l i t y (Maynard and L o o s l i 1969). Crude p r o t e i n was estimated using the conventional conversion of 6.25 times percent n i t r o g e n . Nitrogen was 291 determined using a semi-micro K j e l d a h l a n a l y s i s . To maintain a check on v a l i d i t y of analyses, approximately every tenth sample was analyzed i n d u p l i c a t e . These d u p l i c a t e d data a l s o provided an estimate of experimental e r r o r . The mean d i f f e r e n c e between d u p l i c a t e determinations was 0.25 \u00C2\u00B1 0.2 3 percent crude p r o t e i n (n = 28) (Table 8.2). Table 8.2 Estimates of Experimental E r r o r i n P r o t e i n Analyses f o r Sel e c t e d Species Species No. of .. Estimated e r r o r t e s t e d d u p l i c a t e s mean \u00C2\u00B1 sd (%) Abies lasiooarpa (subalpine f i r ) 4 0. 35 + 0. 30 Cornus stolonifera ( r e d - o s i e r dogwood) 4 0. 20 + 0. 14 Populus tremuloides (trembling aspen) 6 0. 27 + 0. 26 P. balsamifera (black cottonwood) 3 0. 37 + 0. 40 Salix spp. (willow) 10 0. 20 + 0. 19 Sorbus spp. (mountain ash) 1 0. 6 Tot a l s 28 0. 25 + 0. 23 8.3 Crude P r o t e i n Levels For a l l the woody species the average crude p r o t e i n l e v e l was 6.7 percent, based on a f u l l year from May 1972 to A p r i l 1973 (Table 8.3). Annual means v a r i e d between species. Paper b i r c h and trembling aspen had the highest mean annual l e v e l s of 7.4 percent and 7.5 percent crude p r o t e i n , r e s p e c t i v e l y , followed by r e d - o s i e r dogwood, subalpine f i r and mountain ash at about 7.0 percent. Willow 292 Table 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) . Mean percent content (\u00C2\u00B1 sd) Species crude p r o t e i n l i g n i n Abies lasiocarpa (subalpine f i r ) 7.1 \u00C2\u00B1 1. 70 (49) 9. 4 + 1.5 (22) Amelanohiev alnifolia (Saskatoon). 8.1 \u00C2\u00B1 3.05 (4) Betula papyvifera (Paper b i r c h ) -7.4 \u00C2\u00B1 3.24 (36) 10. 0 + 1.3 (50) Cornus stotonifera ( r e d - o s i e r dogwood) 6.9 \u00C2\u00B1 3.20 (34) 8. 5 + 2.3 (19) Lobavia pulmonavia (Lungwort) \u00E2\u0080\u00A211.9 \u00C2\u00B1 1.40 (9) Populus tremuloides (trembling aspen) \u00E2\u0080\u00A2 7.5 \u00C2\u00B1 2.34 (20) 8. 8 + 0.7 (9) P. bdlsamifeva (black cottonwood) 6.5 \u00C2\u00B1 2.7 (19) Salix spp. (willow) \u00E2\u0080\u00A2 6.7 + 2. 3 (90) 10. 5 + 2.1 (44) Sovbus spp. (mountain ash) 6.9 \u00C2\u00B1 2.5 (13) 8. 6 + 2.0 (4) Vaocinium merrbvanacewn (mountain b i l b e r r y ) 8.3 1 4.1 (2) Weighted o v e r a l l mean* 6.7 (276) 9. 8 (148) *Lungwort excluded. A l l species mean = 7.1. ranked next highest w i t h 6.7 percent crude p r o t e i n \u00E2\u0080\u00A2 Black cottonwood ranked lowest i n value w i t h 6. 5 percent Thus the main food species of moose--paper b i r c h , r e d - o s i e r dogwood, subalpine f i r , trembling aspen and willow--had among the highest mean l e v e l s of crude p r o t e i n . A l l browse species showed s i m i l a r annual patterns f o r crude p r o t e i n (Figure 8 . 1 ) . Beginning i n A p r i l , p r o t e i n l e v e l s jumped sharply to peak l e v e l s i n May or June, and then d e c l i n e d g r a d u a l l y over the r e s t of the summer. By 292a Figure 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 . 293 294 October, a f t e r l e a f f a l l , a l l species reached low crude p r o t e i n l e v e l s which remained e s s e n t i a l l y unchanged throughout the winter. Minor f l u c t u a t i o n s recorded between October and A p r i l more l i k e l y r e f l e c t e d v a r i a t i o n s i n experimental e r r o r than a c t u a l changes i n the p l a n t s . R e l a t i v e ranking of browse species changed between seasons, using seasonal d i v i s i o n s as defined i n the food h a b i t s s e c t i o n . One obvious trend was the i n c r e a s i n g rank of subalpine f i r as the winter progressed. This r e s u l t e d from the f a c t t h a t subalpine f i r maintained comparatively higher l e v e l s of p r o t e i n over the winter (6.5 percent) than the other species. Aspen, paper b i r c h and w i l l o w were a l s o among the high ranking species during the two w i n t e r seasons (Nov.-Jan. and Feb.-Apr.). Peak p r o t e i n l e v e l s v a r i e d between species and were reached i n d i f f e r e n t months (Figure 8.1). Paper b i r c h and r e d - o s i e r dogwood a t t a i n e d the highest recorded l e v e l s of 15.3 percent and 16.6 percent crude p r o t e i n i n May. Other browse species had g e n e r a l l y lower peaks, ranging from 11.1 percent to 12.7 percent, t h a t were reached i n June. These d i f f e r e n c e s probably r e f l e c t e d d i f f e r e n c e s i n phenology. Increased p r o t e i n content i n summer browse was due to high l e v e l s i n both the leaves and the stems (Table 8.4). Combing a l l a v a i l a b l e data f o r w i l l o w s (which tends to provide a conservative p i c t u r e ) revealed the f o l l o w i n g Table 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 Selected Browse Species Species sampled (sample s i z e s ) Stems Leaves June J u l y Aug. Sept. June J u l y Aug. Sept. CRUDE PROTEIN Abies lasiooarpa (subalpine f i r ) (5) Betula papyrifera (paper b i r c h ) (1-4) Cornus stolonifera ( r e d - o s i e r dogwood) (1-4) Populus tremuloides (trembling aspen) (T-2) P. balsamifera (black cottonwood) (2) Salix spp. (willow) (1-9) LIGNIN Abies lasiooarpa (subalpine f i r ) (3) Betula papyrifera (paper b i r c h ) (2-4) Cornus stolonifera ( r e d - o s i e r dogwood) (1-2) Populus tremuloides (trembling aspen) (1) Salix spp. (willow) (1-7) 10.1 9.8 7.6 11.5 5.2 8.6 9.0 L I 7.8 4.3 4.0 5.7 4.1 9.7 10.5 8.1 13.0 10.2 5.6 4.7 4.4 5.9 3.9 5.0 8.9 10.2 10.2 9.4 10.0 16.1 16.9 12.6 7.6 4.0 6. 7 14.6 14.0 11.8 13.4 11.9 13.7 11.8 7.9 7.8 5.5 5.6 6.0 7.8 10.3 8.3 9.5 5.5 8.3 10. 7 7.1 4.1 8.6 7.1 ho AO 296 p a t t e r n f o r twigs: crude p r o t e i n increased from approximated 5.3 percent i n l a t e w i n t e r to almost 8 percent by mid-July, a d i s t i n c t d e c l i n e followed i n August to 4 percent, and then a r e t u r n i n the e a r l y f a l l to w i n t er l e v e l s . P r o t e i n l e v e l s i n leaves were u s u a l l y 1.5 to 3 times t h e i r corresponding twig l e v e l s : content appeared to be high i n e a r l y l e a f development r i s i n g to a peak probably i n J u l y , and then tapered o f f u n t i l l e a f f a l l i n September or e a r l y October. Paper b i r c h , r e d - o s i e r dogwood and trembling aspen showed s i m i l a r p a t t e r n s . \u00E2\u0080\u00A2 P r o t e i n l e v e l s i n leaves were p o s i t i v e l y a s s o c i a t e d w i t h those i n stems. The r e g r e s s i o n equation was: y = 4.71x + 1.12, where y = percent crude p r o t e i n i n l e a f samples, and x = percent crude p r o t e i n i n stem sample 2 (n = 16, r =0.56, and standard e r r o r of estimate = 2.21). This r e l a t i o n s h i p i s s i g n i f i c a n t at P < 0.01 (F r a t i o = 17.84). I t can be used to reduce the number of analyses required when sampling shrubs i f .both stem and l e a f p r o t e i n contents are d e s i r e d , by p r e d i c t i n g l e a f from stem values. The e f f e c t of t h i s p r o t e i n d i f f e r e n t i a l should be considered w i t h the p r o p o r t i o n of stems and leaves i n current annual growth. Stems u s u a l l y make up l e s s than 5 0 percent of annual growth but proportions ranged from 29 percent 'for black cottonwood to 70 percent f o r o v a l -l e a f e d blueberry (Table 8.5). For example, i n a 100 g sample of trembling aspen annual growth, approximately 5.5 g 297 Table 8.5 Proport i o n s of Stem and Leaf Tissue i n Current Annual Growth of Selected Browse Species C o l l e c t e d i n September 1972* P r o p o r t i o n of annual Species sampled growth as stem R a t i o of (sample s i z e ) (mean \u00C2\u00B1 sd) stem: l e a f Abies lasiooarpa (subalpine f i r ) (5) 0. 33 + 0. 06 0. 5 : : 1. 0 Amelanchier alnifolia (Saskatoon) (4) 0. 58 + 0. 12 \u00E2\u0080\u00A2 1. 4 : : 1. 0 Betula glandulosa (bog b i r c h ) (1) 0. 45 0. 8 : : 1. 0 B. papyrifera (paper b i r c h ) (7) 0. 49 \u00E2\u0080\u00A2+ 0. 04 1. 0 : : 1. 0 Cornus stolonifera ( r e d - o s i e r dogwood) (4) 0. 44 + 0. 05 0. 8 : : 1. 0 Lonioera involucrata (black twinberry) (1) 0. 53 1. 1 : : 1. 0 Populus tremuloides (trembling aspen) (2) 0. 42 + 0. 08 0. 7 : : 1. 0 P. balsamifera (black cottonwood) (2) 0. 29 + 0. 01 o. 4 : : 1. 0 Salix spp. (willow) (8) 0. 43 + 0. 08 0. 8 : : 1. 0 Sorbus spp. (mountain ash) (2) 0. 73 + 0. 14 2. 7 : : 1. 0 Vaocinium ovalifolium ( o v a l - l e a f e d blueberry) (2) 0. 70 + 0. 04 2. 3 : : 1. 0 *A11 samples c o l l e c t e d between 0.5 and 2 m above the ground. of crude p r o t e i n was a v a i l a b l e In leaves but only 2.5 g i n stems. In a species such as paper b i r c h , the corresponding p r o t e i n content would be 5.3 g and 2.3 g, r e s p e c t i v e l y . P r o t e i n l e v e l s i n lungwort d i f f e r e d i n a l l respects from those of the foregoing woody species. Average annual crude p r o t e i n content was 11.9 percent, higher than a l l other species (Table 8.3). Lungwort showed no i n d i c a t i o n of the annual p a t t e r n t y p i c a l f o r browse species. S i m i l a r l y , the 298 l i c h e n kleotovia savmentosa does not show an annual p a t t e r n (J . Rochelle, pers. comm.). Except f o r the months of May, June and p o s s i b l y J u l y , lungwort had the highest crude p r o t e i n of a l l the species sampled. Results presented so f a r demonstrate gross d i f f e r e n c e s between species. They do not r e v e a l species d i f f e r e n c e s when growing on the same.site. A l s o , i t i s w e l l known that n u t r i e n t l e v e l s between p l a n t s of the same species are a f f e c t e d by many f a c t o r s . Some of these f a c t o r s are s u b s t r a t e , age of stand, browsing h i s t o r y , overstory crown c l o s u r e , and p o s s i b l y p l a n t density. Moreover, i n d i v i d u a l species react d i f f e r e n t l y to these f a c t o r s . Thus while broad d i f f e r e n c e s between species are u s e f u l , more d e t a i l e d information i s needed f o r understanding food s e l e c t i o n and h a b i t a t use, and f o r s e l e c t i n g techniques of vegetation management. The r e s t of t h i s s e c t i o n examines i n f l u e n c e s on crude p r o t e i n of the f o l l o w i n g f a c t o r s : a) species d i f f e r e n c e s , same s i t e b) substrate c) h a b i t a t - t y p e d) age of stand e) c l i m a t e f) s p a t i a l patterns of shrubs (density) P r o t e i n d i f f e r e n c e s r e l a t e d to substrate occurred at l e a s t f o r paper b i r c h (Table 8 . 6 ) . Comparison of monthly samples of t h i s species from the t i l l and l a c u s t r i n e Table 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 Habitat at the Grove Study Area Willow Paper b i r c h P r o t e i n L i g n i n P r o t e i n L i g n i n Month T i l l L a c u s t r i n e T i l l L a c u s t r i n e T i l l L a c u s t r i n e T i l l L a c u s t r i n e May 12.1 6.0 13.1 June 11.1 10.1 August 8.4 7.6 September 6.1 7.2 November 5.4 5.7 11. 4 10.5 5.0 5.9 10. 2 10.9 December 5.4 6.0 5.0 10. 9 11.5 January 5.0 5.8 11.1 13.1 5.2 5.7 11. 0 10.6 February 4.9 4.6 5.2 6.3 8.9 10. 7 March 5.6 5.3 11. 8 10. 2 6.5 10.5 A p r i l 5.0 5.0 12. 4 11. 6 4.6 6.6 8.9 10. 7 Mean* 6.9 6.3 11.7 11. 3 5.0 6.1 + + 10. 0 10.9_ SD 2.7 1.6 0.6 1.3 0.3 0.4 1.0 0.4 *Based only on those months with p a i r e d data. * * S i g n i f i c a n t l y d i f f e r e n t at the 95 percent and 9 0 percent p r o b a b i l i t y l e v e l s , r e s p e c t i v e l y . sD 300 substrates at the Grove Burn revealed a general p a t t e r n of higher p r o t e i n and higher l i g n i n on the l a c u s t r i n e substrate. The e f f e c t of substrate was not s i g n i f i c a n t f o r w i l l o w . This suggests that the e f f e c t of substrate may vary on a species b a s i s . The p o s s i b i l i t y of d i f f e r e n c e s w i t h i n a species being r e l a t e d to the age of a stand were explored (Figure 8.2). Resolution of t h i s p o s s i b i l i t y was l i m i t e d by the f a c t t h a t not a l l species occurred i n a l l stages of succession. For example, w i l l o w could be e a s i l y c o l l e c t e d i n open areas but was d i f f i c u l t to c o l l e c t i n f o r e s t s . The e f f e c t of d i f f e r e n t i a l browsing was probably s l i g h t since browsing appears to have l i t t l e a f f e c t on p r o t e i n content (Oldenmeyer 1974). A l s o other f a c t o r s such as f o r e s t canopy cl o s u r e tend to be as important determinants of browse q u a l i t y as age, thus confounding i n t e r p r e t a t i o n of r e s u l t s . The e v a l u a t i o n of the importance of these and other f a c t o r s can be b e t t e r determined by p r o p e r l y designed f i e l d experiments, r a t h e r than using n a t u r a l s i t u a t i o n s (e.g., Welch e t a l . 1977). One s o l u t i o n to the dilemma was to sample a species that was g e n e r a l l y unbrowsed yet present i n many stages. Subalpine f i r n e a r l y conformed to t h i s except that small t r e e s i n f o r e s t may be of widely d i f f e r i n g ages due to varying growth suppression. Notwithstanding t h i s d i f f i c u l t y , subalpine f i r growing i n young stands (ca. 100 yr) had 300a Figure 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. 3 0 2 g e n e r a l l y l o w e r p r o t e i n l e v e l s t h a n t h o s e i n o l d e r s t a n d s (> 2 0 0 y r ) ( F i g u r e 8 . 2 ) . A g a i n , t h i s i s p a r t l y c o n f o u n d e d b y t h e f a c t t h a t o l d s t a n d s h a v e m o r e o p e n c a n o p i e s t h a n y o u n g e r o n e s . H a b i t a t d i d n o t a f f e c t p r o t e i n c o n t e n t f o r t h e t y p e s a n d s p e c i e s t e s t e d . F o r e x a m p l e , o n t h e s a m e s u b s t r a t e , a t t h e s a m e e l e v a t i o n a n d e x p o s u r e , a n d ' o r i g i n a t i n g f r o m t h e s a m e b u r n , s h r u b s i n d i f f e r e n t h a b i t a t s s h o w e d s i m i l a r c r u d e p r o t e i n l e v e l s ( T a b l e 8 . 7 ) . A t E a g l e , s a s k a t o o n , r e d - o s i e r d o g w o o d , w i l l o w a n d m o u n t a i n a s h g r o w i n g u n d e r a b i r c h f o r e s t h a d g e n e r a l l y s i m i l a r p r o t e i n l e v e l s a s t h e s a m e s p e c i e s g r o w i n g i n t h e o p e n ( T a b l e 8 . 7 ) . A t S a l m o n , w i l l o w g r o w i n g i n a s t a n d o f t r e m b l i n g a s p e n a n d l o d g e p o l e p i n e h a d s i m i l a r p r o t e i n l e v e l s t o t h e s a m e s p e c i e s g r o w i n g i n a n a s p e n s t a n d . D i f f e r e n t s p e c i e s g r o w i n g a t t h e s a m e s i t e h a d d i f f e r e n t c r u d e p r o t e i n l e v e l s ( F i g u r e 8 . 3 ) . T h e s e d i f f e r e n c e s w e r e a s s e s s e d b y c o m p a r i n g a n n u a l c y c l e s o f t w o o r m o r e s p e c i e s c o l l e c t e d f r o m t h e s a m e l o c a l e . D i f f e r e n c e s b e t w e e n s p e c i e s w e r e n o t c o n s t a n t t h r o u g h o u t a n a n n u a l c y c l e ( F i g u r e 8 . 3 ) . A t G r o v e 4 , a s p r u c e - s u b a l p i n e f i r s t a n d , r e d - o s i e r d o g w o o d h a d h i g h e r l e v e l s o f p r o t e i n i n s u m m e r t h a n s u b a l p i n e f i r , b u t t h e s e p o s i t i o n s r e v e r s e d b y a u t u m n . A s i m i l a r p a t t e r n w a s r e c o r d e d f o r c o t t o n w o o d a n d w i l l o w i n t h e G r o v e b u r n . I n m o s t i n s t a n c e s , i n t e r - s p e c i f i c d i f f e r e n c e s i n s u m m e r p r o t e i n l e v e l s w e r e l e s s t h a n t h o s e o f Table 8.7 E f f e c t of Habitat on Crude P r o t e i n Levels i n Sel e c t e d Browse Species at the Eagle and Salmon Study Areas STUDY AREA Species \" t \" value Habitat tab. at P = 0.05 c a l c u l a t e d EAGLE: shrub-type birch-type Arnelanohier alnifolia (Saskatoon) (2)* 8.3 \u00C2\u00B1 4.0 7.9 \u00C2\u00B1 3.4 12. 71 0. 78 Cornus - stolonifera. (red-o s i e r dogwood) (10) 5.9 \u00C2\u00B1 3.0 6.0 + 2.5 2.26 0. 36 Salix spp. (willow) (9) 6.5 \u00C2\u00B1 2.7 6.8 \u00C2\u00B1 3.1 2. 31 0.90 Sorbus spp. ash) (5) (mountain 7.1 \u00C2\u00B1 3.3 6.5 \u00C2\u00B1 2.1 2.78 0. 85 SALMON: aspen-type aspen-pine type Salix spp. (willow) (8) 6.1 \u00C2\u00B1 1.8 6.5 \u00C2\u00B1 2.3 2.37 2. 28 *No. of p a i r s i n sample. o 303a Figure 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 . \u00E2\u0080\u00941 -1 I 1 1 \u00E2\u0080\u0094I 1 1 1 1- 1 1 T-APR JUNE AUG OCT DEC FEB APR 305 w i n t e r . For example, at Grove 4, summer d i f f e r e n c e s between subalpine f i r and r e d - o s i e r dogwood ranged from 0 - 1.6 percent while winter d i f f e r e n c e ranged from 1.5 per-cent to 2.6 percent (Figure 8.3). Li m i t e d data suggested that d i f f e r e n c e s between some species were c o n s i s t e n t , whatever the h a b i t a t . At a l l three s i t e s where w i l l o w and aspen were con c u r r e n t l y sampled, winter l e v e l s of p r o t e i n were almost always greater i n aspen than i n w i l l o w (Figure 8.4). The magnitude of t h i s d i f f e r e n c e v a r i e d between the s i t e s but the p a t t e r n remained constant. Year-to-year v a r i a t i o n s i n climate a l s o a f f e c t e d crude p r o t e i n content f o r a given calendar date. Climate i n f l u e n c e s p h e n o l o g i c a l development, f o r example, by r e t a r d i n g l e a f i n g out and so de l a y i n g v e r n a l p r o t e i n increases. Some i n d i c a t i o n of the d i f f e r e n c e s experienced i n the study area were revealed by examining crude p r o t e i n l e v e l s i n A p r i l of 1972 and 1973. The average d i f f e r e n c e between years was 0.9 percent (Table 8.8). Levels i n 1972 exceeded those those f o r 19 73 except w i t h trembling aspen at s i t e G2 where the opposite p a t t e r n occurred. D i f f e r e n c e s were l e a s t f o r r e d - o s i e r dogwood (0.45 percent) and approximately one percent f o r the other species. These d i f f e r e n c e s suggested t h a t the 1972 growing season began e a r l i e r than the 19 7 3 one. Based on mean monthly temperature data from the Prin c e George weather 305a i Figure 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 . Table 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 Crude P r o t e i n (%) L i g n i n (%) Species sampled S i t e A p r i l 1972 A p r i l 19 73 A p r i l 19 72 A p r i l 1973 Betula papyrifera (paper birch) B l B3 E2 7.0 6.0 9.7 12. 7 9.4 11.2 Cornus stolonifera (red-o s i e r dogwood) E l 4.9 4.2 E2 4.5 4.3 7.9 8.9 Populus tremuloides (trembling aspen) E2 G2 7.1 5.6 6.0 6.6 8.5 8.1 Salix spp. (willow) B3 E l 5.3 4.2 9.8 11. 8 E2 5.3 4.3 11.1 14.1 Mean d i f f e r e n c e s 0.9 1.4 o - j 308 o f f i c e (Environment Canada 1972-1973), the growing season began on approximately A p r i l 6 o r 7 i n 19 72 and on A p r i l 11 i n 1973. Growing season was assumed to begin when mean d a i l y temperatures reached 5.6\u00C2\u00B0C (Conrad 1950, quoted i n Harper 1969). Moisture a l s o a f f e c t s growth and p r o t e i n content. R a i n f a l l was 17 mm f o r A p r i l , 1972 but o n l y 10 mm i n A p r i l , 1973. Thus f o r the years compared, crude p r o t e i n content r e f l e c t e d moisture d i f f e r e n c e s more c l o s e l y than temperature d i f f e r e n c e s . The i n t e r a c t i o n between c l i m a t e and n u t r i t i v e content of forages i s c l e a r l y more complex than i n d i c a t e d here. The s i g n i f i c a n t p o i n t i s t h a t annual v a r i a t i o n s do occur. These may be c r i t i c a l f o r moose coming out of a w i n t e r on sub-maintenance d i e t s . 8.4 L i g n i n L e v e l s L i g n i n content averaged 9.8 p e r c e n t based on a l l months f o r the f i v e woody s p e c i e s which had s u f f i c i e n t data (Table 8.3). S i m i l a r to crude p r o t e i n l e v e l s , annual means v a r i e d between s p e c i e s . Red-osier dogwood had the lowest l i g n i n v a l u e s . L e v e l s i n mountain ash were a l s o comparatively low w i t h a mean annual l i g n i n value of 8.6 per-cent, although more samples are needed to c o n f i r m t h i s . Trembling aspen ranked next w i t h an annual mean of 8.8 per-cent, while w i l l o w and paper b i r c h were l i k e l y the l e a s t d i g e s t i b l e of the f i v e s p e c i e s , w i t h l i g n i n contents of 10.5 p e r c e n t and 10.0 percent, r e s p e c t i v e l y . 309 Two types of annual patterns of l i g n i n content were i l l u s t r a t e d by the f i v e species (Figure 8.5). One p a t t e r n , showed by subalpine f i r , was f o r r e l a t i v e l y constant l i g n i n l e v e l s throughout the year, except f o r low l e v e l s i n e a r l y s p r i n g t i s s u e s . The second p a t t e r n was demonstrated by paper b i r c h , w i l l o w , and r e d - o s i e r dogwood. Beginning i n A p r i l , l i g n i n decreased to a mid-summer minimum. This low probably c o i n c i d e d w i t h maximum l e a f development and minimal l i g n i f i c a t i o n of stems. L i g n i n content increased a f t e r August u n t i l approximately October and then remained / comparatively constant throughout the winter. Trembling aspen probably followed a s i m i l a r p a t t e r n although f u r t h e r data are needed f o r the summer p e r i o d . The lowest l i g n i n l e v e l s v a r i e d between specie s , and they were a t t a i n e d i n d i f f e r e n t months (Figure 8.5). Red-o s i e r dogwood had the lowest recorded l i g n i n value of 5.1 percent i n June. Willow had i t s l e a s t l i g n i n value of 7.9 percent i n August, w h i l e paper b i r c h had 8.8 percent and 8.9 percent i n August and May, r e s p e c t i v e l y . Subalpine f i r had r e l a t i v e l y high l i g n i n values i n summer (approximately 10 percent) and the lowest l e v e l of 6.7 percent i n A p r i l . L i g n i n content v a r i e d between stem and l e a f components and so a f f e c t e d whole-sample l e v e l s during summer. In a l l cases examined, l i g n i n was lower i n leaves than i n stems over the June - September p e r i o d (Table 8.4). Leaf l i g n i n was l e a s t i n r e d - o s i e r dogwood and highest i n 309a Figure 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 . 310 311 paper b i r c h . Stem samples showed a corresponding p a t t e r n . Unlike crude p r o t e i n , stem l i g n i n was not s i g n i f i c a n t l y 2 r e l a t e d to l e a f l i g n i n (n = 13, r = 0.04, F r a t i o = 0.44). Tabulated F values at the 95 percent p r o b a b i l i t y l e v e l i s 4. 75. S i m i l a r to crude p r o t e i n , l i g n i n content was examined to determine i f and how various s i t e f a c t o r s a f f e c t e d i t . Since l i g n i n data were l e s s comprehensive than p r o t e i n data, the e v a l u a t i o n of these f a c t o r s i s l e s s complete. Nonetheless s i g n i f i c a n t and c o n s i s t e n t e f f e c t s were n o t i c e a b l e . Species growing at the same s i t e had d i s s i m i l a r l i g n i n l e v e l s (Table 8.9 and Figure 8.6). For example, i n the shrub type on the Eagle study area, r e d - o s i e r dogwood had c o n s i s t e n t l y lower percent l i g n i n than paper b i r c h (Figure 8.6a). However, i n c o n s i s t e n c y i n d i f f e r e n c e s was the r u l e as at the mature spruce-subalpine f i r f o r e s t s i t e at Grove (Figure 8.6c). Here, r e d - o s i e r dogwood had lower l i g n i n i n summer than subalpine f i r regeneration, but higher l e v e l s i n winter. The p a t t e r n of d i f f e r e n c e s between species appeared to h o l d , whatever the s i t e . The best data to i l l u s t r a t e t h i s p o i n t were f o r paper b i r c h and w i l l o w on beach deposits at Salmon, l a c u s t r i n e substrates at Eagle and t i l l at Grove (Figure 8.6a, b, d). At these three s i t e s , w i l l o w had lowest l i g n i n l e v e l s i n August, while f o r paper b i r c h i t was Table 8.9 Difference 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 Mean % l i g n i n content* Substrate Habitat ( s i t e ) Subalpine f i r Paper b i r c h Red-osier dogwood Trembling aspen Willow T i l l : 12 y r o l d burn ( B l , Gl) 50 yr o l d pine f o r e s t (B3) 120 y r o l d pine f o r e s t (S5) L a c u s t r i n e : 12 y r o l d burn (G2) 40 y r o l d burn (E2) 200 y r o l d s p r u c e - f i r f o r e s t (G4) 250 y r o l d s p r u c e - f i r f o r e s t (E4) Beach deposit: aspen f o r e s t (S2, S3) Recent a l l u v i u m : spruce-cottonwood f o r e s t (Si) 9.2 9.2 8.9 9.5 10. 0 9.6 9.4 7. 8 7.4 11.9 9.0 8.3 10.5 10. 0 9.9 11.0 12.4 10.2 *matched samples used i n a l l cases, N 4, 5, 4, 3, and 3, f o r species as l i s t e d above. 312a Figure 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. 14 12 10 8 2 <2 4 313 [a. SPRUCE-SUBALPINE FIR FOREST (G4) \u00E2\u0080\u00A2 SUBALPINE FIR A RED OSIER DOGWOOD 12 14 1 i \u00C2\u00BB i i i i i i A M J J A S O N D J F M A !b. MIXED WOOD (S2/S3) \u00E2\u0080\u00A2 BIRCH A WILLOW \u00E2\u0080\u00A2 BIRCH A WILLOW A RED OSIER DOGWOOD c. 40 YEAR OLD SHRUB TYPE (E2) 12 KH 12 ' 1 I I I I I I | 1 | -m A M J J A S O N D J F M A d. 12 YEAR OLD SHRUB TYPE (G1) \u00E2\u0080\u00A2 BIRCH A WILLOW 10 84 A M J J A S O N D J F M A ^ A M J J A S O N D J F M A 314 September. Late winter l i g n i n i n w i l l o w increased but d e c l i n e d i n paper b i r c h . Substrate a l s o a f f e c t e d l i g n i n content, at l e a s t f o r paper b i r c h (Table 8.9, Figure 8.6). L i g n i n i n paper b i r c h growing on a t i l l substrate was g e n e r a l l y lower than that growing on a l a c u s t r i n e substrate. Willow showed no s i g n i f i c a n t d i f f e r e n c e s . These two s i t e s were l o c a t e d on the Grove study area at two nearby s i t e s t h a t d i f f e r e d only i n nature of the substrate. Age of a sere appeared to modify l i g n i n values (Table 8.9). The percent l i g n i n g e n e r a l l y d e c l i n e d w i t h stand age f o r w i l l o w , paper b i r c h and trembling aspen on t i l l and l a c u s t r i n e s u b s t r a t e s . Unlike crude p r o t e i n l e v e l s , l i g n i n contents i n consecutive A p r i l s (1972 and 1973) were not s i g n i f i c a n t l y 2 r e l a t e d (r = 0.44, c a l c u l a t e d F r a t i o = 3.18, t a b u l a t e d F r a t i o = 6.61 at 95 percent p r o b a b i l i t y l e v e l (Table 8.7). A p o s s i b l e reason f o r t h i s d i f f e r e n c e i s t h a t p r o t e i n has a mobile component t h a t could be t r a n s l o c a t e d from o l d to new t i s s u e . However, l i g n i n forms p a r t of a twig's s t r u c t u r e and i s t h e r e f o r e unable to be t r a n s l o c a t e d i n a s i m i l a r way. 8.5 Discussion 8.5.1 Crude P r o t e i n Levels i n Moose Forages Moose forages i n the P r i n c e George area had crude p r o t e i n l e v e l s s i m i l a r to those reported f o r other moose 315 ranges (Table 8.10), at l e a s t i n winter (November - March). This time p e r i o d was s e l e c t e d f o r comparative purposes since mid-winter l e v e l s were most s t a b l e ; v a r i a t i o n s i n time of sampling would th e r e f o r e have minimal e f f e c t on crude p r o t e i n comparisons. For subalpine f i r , w i n t e r crude p r o t e i n was 6.5 percent i n the P r i n c e George area, 6.4 per-cent i n the Quesnel area (120 km S. of P r i n c e George) (Cowan et a l . 1950), and 5.6 percent i n Wyoming (Houston 1968). For paper b i r c h , crude p r o t e i n was 5.3 percent, 7.0 percent, 7.5 percent and 6.8 percent i n P r i n c e George (Table 8.10), Quesnel (Cowan e t a l . 1950), Alaska (Oldenmeyer 1974), and i n Sweden (Ahlen 1975, species not i d e n t i f i e d ) . For red-o s i e r dogwood, Cowan e t a l . (1950) reported the same crude p r o t e i n l e v e l of 4.8 percent. Data f o r aspen, cottonwood, w i l l o w , and ash were al s o s i m i l a r over t h i s wide geographic range (Table 8.10). A notable exception i s the r e s u l t s of S i l v e r (1976) from northeastern B r i t i s h Columbia. His l e v e l s of crude p r o t e i n were approximately one-half of the average l e v e l s summarized i n Table 8.10. Other species he sampled were a l s o notably lower than those reported i n the l i t e r a t u r e . A f t e r re-checking to r u l e out major sources of e r r o r , S i l v e r (19 76) concluded that these low values were genuine. However, i t i s d i f f i c u l t to b e l i e v e that these low estimates were genuine, given the general agreement i n a l l other st u d i e s f o r higher l e v e l s . Furthermore, the moose i n h i s Table 8.10 Comparisons of Crude P r o t e i n Values f o r Current Annual Growth of Common Winter Foods of Moose (November-March) Crude p r o t e i n value i n w i n t e r (%) Species 1* 2 3. 4 5 6 7 8 9 mean 1 sd** Abies lasiooarpa (subalpine f i r ) 6.5 6.4 5.6 6.2 + 0.5 Amelanohier alnifolia (Saskatoon) 5.5** * 7.0 6.1 6.2 + 0.8 Betula papyrifera (paper b i r c h ) 5.3 7.0 7.5 2.9 7.9 6.9 + 1.1 Cornus stolonifera (dogwood) 4.8 4.8 6.4 6.5 + 1.3 Lobaria pulmonaria (lungwort) 11.9 12.1*** 12.0 + 0.1 Populus tremuloides (aspen) 6.0 7.1 6.9 6.5 6.8 3.1 8.7 6.0 8.5 7.1 + 1.0 P. balsamifera (cottonwood) 4.7 6.1 8.0 6.3 + 1. 7 Salix spp. (willow) 5.4 6.0 6.5 3.1 8.2 6.7 7.0 6.6 + 1.0 Sorbus spp. (mountain ash) 5.5 5.5 Vaooinium spp. ( v a c c i n i a ) 5.4 8.1*** 6.7 + 1.9 *Sources of data: I - - . t h i s study, 5 - Oldenmeyer 2,- Cowan (1974), 6 -et a l . - S i l v e r (1950), 3 -' (1976), 7 Houston (1968), 4 - Dietz - Peek et a l . (1976), (1972), 8 - J . Rochelle (pers. comm.), f o r Lobaria and Vaooinium, Kubota et a l . (1970)for the others, 9 - Stewart et a l . (1977). **Excluding data from source 8 ( S i l v e r 1976). ***A. florida, L. oreganum (?), V. parvifolium. 317 study area were healthy, productive animals t h a t wintered i n t y p i c a l l y high d e n s i t i e s . I f crude p r o t e i n values were as low as he reported, low p r o d u c t i v i t y and low d e n s i t i e s would be expected. As S i l v e r (1976) recommends, re-sampling i s necessary to s u b s t a n t i a t e h i s i n i t i a l data. A d i e t a r y l e v e l of 7 percent crude proten i s judged to be minimal f o r maintenance of moose (Gasaway and Coady 1974), w h i t e t a i l deer (French et a l . 1956), mule deer (Dietz 1965), reindeer (Luick et a l . 1971), and domestic ruminants (Swenson 1970, quoted by LeResche and Davis (1973)).. French et a l . (1956) suggested that approximately 13 percent crude p r o t e i n l e v e l s were needed i n summer f o r w h i t e t a i l deer. That many herbivores can s e l e c t d i e t s higher i n crude p r o t e i n and other n u t r i e n t s than determined from hand picked samples has been demonstrated, e.g., B i s s e l l (.1959,). I t has a l s o been shown than species mixes can have higher d i g e s t i b i l i t i e s than those p r e d i c t e d from s i n g l e species determinations. These same phenomena probably apply to moose. A l s o , the P r i n c e George moose t y p i c a l l y ate only the t e r m i n a l p o r t i o n s of annual growth. These t e r m i n a l p o r t i o n s have higher p r o t e i n l e v e l s than would be estimated from sampling a l l annual growth (see s e c t i o n 8.5.4). Thus the d i e t a r y crude p r o t e i n l e v e l f o r P r i n c e George moose was greater than the l e v e l judged necessary f o r maintenance. P r o t e i n content of lungwort was almost twice t h a t of the browse species (11.9 percent vs. 6.7 percent). On 318 northern Vancouver I s l a n d , Rochelle (pers. comm.) found s i m i l a r values f o r another species of Lobavia (probably oveganum). Crude p r o t e i n averaged 12.1 percent f o r two samples picked from trees i n December.and May. This and other l i c h e n species, made a v a i l a b l e through l i t t e r f a l l , form an important p a r t of b l a c k t a i l deer w i n t e r d i e t i n t h i s study area. The r o l e of l i t t e r f a l l as food f o r ungulates was summarized by Rochelle (1976?). High crude p r o t e i n values have been recorded f o r Peltigera spp., another f o l i o s e l i c h e n , at 17.5 percent and 19.8 percent by S c o t t e r (1965) and LeResche and Davis (1973), r e s p e c t i v e l y . Lichens were not eaten by moose i n greater amounts probably due to f a c t o r s such as a v a i l a b i l i t y , p a l a t a b i l i t y , d i f f i c u l t y of securing i t and low d i g e s t i b i l i t y (Rochelle, pers. comm.). I t should be noted that many other l i c h e n s have comparative-l y low p r o t e i n l e v e l s of approximately 2-4 percent. 8.5.2 Assessing the N u t r i t i v e Values of Forages The r e l a t i o n s h i p of forage p r o t e i n and d i g e s t i b i l i t y to moose n u t r i t i o n assumes i n c r e a s i n g importance w i t h i n t e n s i f i c a t i o n of moose management. The n u t r i t i o n a l parameters t h a t l i m i t moose p r o d u c t i v i t y r e q u i r e c l e a r d e f i n i t i o n , e s p e c i a l l y since food q u a l i t y i s one of the major determinants of c a r r y i n g c a p a c i t y . Food c o n s i s t s of energy, minerals, vitamins or t h e i r p r e c u r s o r s , and complex organic compounds such as p r o t e i n s , c e l l u l o s e s , l i g n i n , 319 f a t s , e t c . An optimum food supply contains the necessary c o n s t i t u e n t s i n c o r r e c t balance to meet.changing n u t r i t i o n a l needs of the animal. As no s i n g l e p l a n t species i s a \" p e r f e c t \" food, animals attempt to meet t h e i r needs by feeding on a v a r i e t y of s p e c i e s , and by securing some mineral supplements by gnawing bones and a n t l e r s , and by e a t i n g s o i l at mineral l i c k s . Moose calves may l e a r n to eat c o r r e c t combinations of food from t h e i r mothers (Edwards 1976) . Developing methods to assess the n u t r i t i o n a l adequacy of f o o d s t u f f s i s a focus of considerable past and present research. The most v a l i d method i s by feeding t r i a l s , but these become very demanding of resources i f a l l f o o d s t u f f s and combinations are assessed. The general approach has been to evaluate major food species through feeding t r i a l s both s i n g l y and i n l i m i t e d combinations, and e x t r a p o l a t e from chemical analyses f o r others. An i d e a l measure of forage q u a l i t y would be one that enables \"the e s t i m a t i o n of d i g e s t e d end products absorbed per u n i t of time, and should be meaningful to a l l kinds of forage-consuming animals under a l l environmental c o n d i t i o n s \" (Lucas 1962, quoted by Morris and Kovner (1970)). Although development of such c r i t e r i a . , are u n l i k e l y , considerable e f f o r t has been devoted to discovery of analyses that are b i o l o g i c a l l y meaningful, that i s , analyses whose r e s u l t s c o r r e l a t e w e l l w i t h standards of production, and yet are r e l a t i v e l y economic, r o u t i n e and repr o d u c i b l e . N u t r i t i v e values of forages depend upon both chemical composition of the forage, and the d i g e s t i b i l i t y and nature of digested products. Many chemical methods of assessing both aspects have been developed. U n t i l q u i t e r e c e n t l y , the most widely used one was the proximate or Weedne system of analyses. I t separates p l a n t s i n t o s i x components, v i z . , dry matter, ether e x t r a c t , ash, crude p r o t e i n , crude f i b e r and n i t r o g e n free e x t r a c t (NFE). The l a t t e r two attempt t o separate i n d i g e s t i b l e and d i g e s t i b l e components of carbohydrates, r e s p e c t i v e l y , but t h i s separation i s imperfect. D i g e s t i b i l i t y of the crude f i b e r f r a c t i o n v a r i e s according to i t s composition; and NFE contain some i n d i g e s t i b l e and p r a c t i c a l l y i n d i g e s t i b l e components (Ha r r i s 19 70). Since NFE i s determined by s u b t r a c t i o n , i t a l s o \"pools\" e r r o r s made during the other analyses. Proximate a n a l y s i s r e s u l t s a l s o have other d i f f i c u l t i e s . I n t e r p r e t a t i o n of the ether e x t r a c t f r a c t i o n i s made d i f f i c u l t since the true f a t s may form l e s s than one-half of the ether e x t r a c t ( S u l l i v a n 1962), and sin c e some of the e x t r a c t e d e s s e n t i a l o i l s i n h i b i t ruminal f u n c t i o n (Nagy et a l . 1964; Oh et a l . 1967, 1968 and 1970). The shortcomings of proximate analyses have been w e l l described by Van Soest (1965, quoted i n Robbins et a l . 1975). The l e a s t problematic component i s probably p r o t e i n : 321 Maynard and L o o s l i (19 69) suggested that the d i s t i n c t i o n between crude and true p r o t e i n i s not worthwhile since non-p r o t e i n n i t r o g e n compounds such as amides and ammonium s a l t s r a r e l y form a s i g n i f i c a n t p r o p o r t i o n of the t o t a l . Improvements upon the proximate a n a l y s i s system have been e i t h e r chemical or b i o l o g i c a l , and focused mainly upon o b t a i n i n g r e a l i s t i c estimates of d i g e s t i b i l i t y . The detergent system proposed by Van Soest (1967) has been widely adopted, i n c l u d i n g browse species, e.g., Robbins et a l . (1975). The T i l l e y and Terry (1963) system and i t s v a r i a n t s have a l s o been widely used as in vitro approaches to e s t i m a t i n g d i g e s t i b i l i t i e s . Source of i n o c u l a and processing of residues can a f f e c t estimates of d i g e s t i b i l i t y (Robbins e t a l . 1975). Useful references to assess forage values f o r w i l d ruminants are Van Dyne (1968). and H a r r i s (1970) . Two of the more recent methods t e s t e d f o r determining d i g e s t i b i l i t y are the a c e t y l bromide determina-t i o n of l i g n i n used i n t h i s study, and the use of i n f r a r e d r e f l e c t a n c e spectroscopy (Norris et a l . 1976). The l a t t e r technique holds much promise due to high c o r r e l a t i o n s w i t h crude p r o t e i n (0.99), a c i d and n e u t r a l detergent f i b e r (0.96 and 0.98) , and in vitro and in vivo r e s u l t s . While both the above methods re q u i r e v a l i d a t i o n w i t h browse species and w i l d ungulates, they o f f e r r a p i d means of assessing important forage parameters. 322 8.5.3 Factors A f f e c t i n g N u t r i e n t Levels That n u t r i e n t l e v e l s vary i n p l a n t s i s a truism. A n a l y t i c a l demonstrations of t h i s f o r n a t i v e forages, i n c l u d i n g shrubs, date back i n North America to at l e a s t 1911 (Knight et a l . 1911, c i t e d i n Dietz 1972). Comprehen-s i v e s t u d i e s e x i s t f o r many geographic areas, many of which are given i n Dietz (1972). Fortescue and Marten (1970) reviewed m i c r o n u t r i e n t v a r i a t i o n s i n f o r e s t s . S i m i l a r s t u d i e s r e l e v a n t to moose forages i n c l u d e those by Cowan et a l . (1950), Kubota (1974), Kubota e t a l . (1970), Peek e t a l . (1976), and S i l v e r (1976). Many f a c t o r s have been i d e n t i f i e d and examined as to t h e i r e f f e c t on forage n u t r i e n t s , yet there are few coherent explanations. W i l d l i f e - o r i e n t e d reviews by Laycock and P r i c e (1970), Dietz (1972)., Oelberg (1956) and Short et a l . (19 7 2) found t h a t many f a c t o r s have equ i v o c a l e f f e c t s on n u t r i e n t l e v e l s . Probably the s i n g l e most important problem c o n f r o n t i n g f a c t o r i d e n t i f i c a t i o n and e v a l u a t i o n was defined by Laycock and P r i c e (1970:44): In most of the s t u d i e s reported, e f f e c t s of i n d i v i -dual environmental f a c t o r s were confounded w i t h those of other i n f l u e n c e s or w i t h stages of p l a n t development. C a r e f u l l y c o n t r o l l e d s t u d i e s are needed to define the e f f e c t s of these f a c t o r s , both alone and i n combinations,, on the chemical composition of forage p l a n t s . Another problem i s t h a t most s t u d i e s assume a l l species respond s i m i l a r l y to a given f a c t o r . However, species vary 323 markedly i n t h e i r adaptations to environmental f a c t o r s such as l i g h t , s o i l n u t r i e n t s and competition. These d i f f e r i n g adaptations could be r e f l e c t e d i n p l a n t n u t r i e n t l e v e l s as w e l l . For example, Wali and K r a j i n a (1973) demonstrated that many sub-boreal s p e c i e s , such as bearberry, foam flower and white peavine, vary i n t h e i r abundance along gradients of l i g h t and n u t r i e n t s . Thus i t may be that at a given l e v e l of l i g h t or other f a c t o r s , the species best adapted to i t would have the highest l e v e l of a p a r t i c u l a r n u t r i e n t . Another major d i f f i c u l t y f a c i n g f a c t o r e v a l u a t i o n i s unstandardized sample c o l l e c t i o n . Some workers c o l l e c t a l l current annual growth while others c o l l e c t only the t e r m i n a l few cm; some separate leaves from twigs while others do not. Given the marked gradients of n u t r i e n t s i n p l a n t s , comparisons among r e s u l t s are questionable. Despite the foregoing problems, continued s t u d i e s on the t o p i c of f a c t o r s . a f f e c t i n g n u t r i e n t l e v e l s i n d i c a t e i t s importance to w i l d l i f e b i o l o g i s t s . V a r i a t i o n s and explanation of n u t r i e n t l e v e l s can a i d i n t e r p r e t a t i o n of h a b i t a t preferences by moose and other herbivores. Changes i n feeding behaviour and d i e t a l s o are more e x p l a i n a b l e i f adequate n u t r i e n t data are a v a i l a b l e . Avoidance of some seemingly acceptable species or p l a n t s has been explained by stu d i e s on a n t i - q u a l i t y f a c t o r s . The e v a l u a t i o n and s e l e c t i o n of h a b i t a t management p r a c t i c e s o f t e n depends upon 324 knowing how they w i l l a f f e c t forage n u t r i e n t s as w e l l as f o r how long (Peek 19 74). N u t r i e n t v a r i a t i o n s a l s o provide important inputs to n u t r i t i o n a l s t u d i e s and models (Gasaway and Coady 1974, Robbins 1973). F i n a l l y , from a procedural p e r s p e c t i v e , an awareness of e f f e c t s of p r i n c i p a l f a c t o r s i s a p r e r e q u i s i t e f o r proper experimental design and sample c o l l e c t i o n . Factors governing forage n u t r i e n t l e v e l s can be c l a s s i f i e d i n t o the f o l l o w i n g f i v e l e v e l s : 1) v a r i a t i o n s between taxonomic or s t r u c t u r a l p l a n t c l a s s e s 2) v a r i a t i o n s between species w i t h i n a c l a s s 3) v a r i a t i o n s between p l a n t s w i t h i n a species (genetic d i v e r s i t y , etc.) 4) v a r i a t i o n s between p l a n t p a r t s w i t h i n a p l a n t 5) v a r i a t i o n s w i t h i n p l a n t p a r t s ( n u t r i e n t gradients) The importance of these l e v e l s v a r i e s according to the season and to the k i n d of p l a n t . For example, p h y s i o l o g i c a l stage of growth may be most important i n summer but not i n w i n t e r , when species d i f f e r e n c e s may be most marked. Crude p r o t e i n and l i g n i n l e v e l s v a r i e d between the three c l a s s e s of p l a n t s sampled, v i z . , l i c h e n s , c o n i f e r s and deciduous d i c o t s . The l i c h e n , lungwort, had comparatively high crude p r o t e i n values t h a t showed l i t t l e evidence of annual v a r i a t i o n . However, l i c h e n s show considerable v a r i a t i o n i n t h e i r annual means; some f o l i o s e l i c h e n s such as Lobaria oveganum (?) and Peltigera spp. have high p r o t e i n 325 l e v e l s , but other species and growth forms have values as low as 2 - 3 percent p r o t e i n ( K e l s a l l 1968, J . Rochelle, pers. comm., S i l v e r 1976). L i g n i n or d i g e s t i b i l i t y was not determined f o r lungwort. However, i n a r e l a t e d species, Lobavia (oreganum?), Rochelle (pers. comm.) found low in vitro d i g e s t i b i l i t i e s f o r samples c o l l e c t e d i n December (16.5 percent, n = 4) and i n May (10.5 percent, n = 2). For the arbore a l l i c h e n , Alectoria spp. , S i l v e r (19 76:82) found an average of 10.1 percent l i g n i n i n samples, c o l l e c t e d i n May, September and January. For the t e r r e s t r i a l f o l i o s e l i c h e n , Peltigera spp., LeResche and Davis (1973) found 25.3 percent crude f i b e r i n samples c o l l e c t e d i n May. These r e s u l t s suggest t h a t l i c h e n s have low d i g e s t i b i l i t y . B o real and sub-boreal deciduous shrubs t y p i c a l l y show a d e f i n i t e seasonal p a t t e r n i n crude p r o t e i n . Peak values develop rather q u i c k l y i n e a r l y summer and then d e c l i n e sharply to a low, steady value during f a l l and w i n t e r (e.g. G r i g a l et a l . 1976, S i l v e r 1976, Tew 1970, Oldenmeyer 19 74). Browse species from the P r i n c e George area showed a s i m i l a r p a t t e r n (Figure 8.1). Presumably, t h i s p a t t e r n r e f l e c t s the u s u a l l y abrupt, b r i e f growing seasons of northern l a t i t u d e s . Changes i n p r o t e i n l e v e l s of both l e a f and twig t i s s u e s are responsible f o r the dramatic changes i n seasonal crude p r o t e i n . Current leaves had higher crude 326 p r o t e i n l e v e l s than twigs and, even though twig crude p r o t e i n l e v e l s double over the summer, t h e i r values were ofte n only 50-60% those i n leaves. Moreover, the m a j o r i t y of species had higher proportions of leaves than twigs i n t h e i r c u r r e n t annual growth. The high crude p r o t e i n l e v e l s of leaves may e x p l a i n the common feeding behaviour of summer moose s t r i p p i n g f o l i a g e from woody shoots. However, i n South Dakota, Dietz (19 72) found that annual twigs of s e v e r a l shrub species showed i n c r e a s i n g p r o t e i n between f a l l and winter. Since a d d i t i o n a l data on date of c o l l e c t i o n , sampling methods were not provided, i t i s d i f f i c u l t to assess whether h i s r e s u l t s r e f l e c t s p e c i e s - s p e c i f i c d i f f e r e n c e s or sampling d i f f e r e n c e s . Crude p r o t e i n a n d . l i g n i n l e v e l s i n the s i n g l e c o n i f e r s t u d i e d , subalpine f i r , were g e n e r a l l y s i m i l a r to those described f o r the deciduous shrubs. However, the r e l a t i v e ranking of t h i s species increased over the f a l l and winter so t h a t by l a t e w i n t e r , i t had the highest p r o t e i n values r e l a t i v e to other browse species. N u t r i e n t l e v e l s vary among p l a n t s of a s i n g l e species. This v a r i a b i l i t y has both s i t e (environmental) and genetic components. The i n f l u e n c e of s i t e f a c t o r s appears to be a most undecided i s s u e . The complex and v a r i a b l e i n f l u e n c e of these f a c t o r s l e d Oldenmeyer (1974:220) to recommend that \" I f exact values are needed f o r n u t r i e n t content, they should be obtained from the area where the 327 study i s t a k i n g place and not e x t r a p o l a t e d f o r other areas.\" Results from the present study i n d i c a t e t h a t substrate has v a r i a b l e e f f e c t s . The l i m i t e d data obtained f o r w i l l o w and b i r c h suggested t h a t species and substrate i n t e r a c t . Substrate a f f e c t e d paper b i r c h but not w i l l o w f o r l a c u s t r i n e and t i l l s o i l s . In a study of three species browsed by w h i t e t a i l deer i n V i r g i n i a , Hundley (1959) found t h a t crude p r o t e i n l e v e l s were s i g n i f i c a n t l y a f f e c t e d by s o i l type. Bergerud and Manuel (19 68) found up to 2 2 per-cent d i f f e r e n c e s i n crude p r o t e i n l e v e l s i n current growth of balsam f i r . However, other f a c t o r s such as moisture, overstory, e t c . were not c o n t r o l l e d . Conversely L i n d l o f e t a l . (19 74) b e l i e v e d that many species are able to maintain chemical composition w i t h i n l i m i t s , i r r e s p e c t i v e of s o i l p r o p e r t i e s . D a n iel and Harper (1934) (quoted i n Laycock and P r i c e 1970) agreed w i t h t h i s . The general opinion suggests t h a t p l a n t s of the same species growing i n d i f f e r e n t s o i l s often d i f f e r i n chemical composition and, consequently, i n p a l a t a b i l i t y (Heady 1964 quoted i n Laycock and P r i c e 1970). Peek e t a l . (1976) attempted to r e l a t e twig n u t r i e n t l e v e l s to s e v e r a l s i t e f a c t o r s : stand age, stand canopy c l o s u r e and s o i l l e v e l s of P, K, Ca, Fe and Mg. Simple c o r r e l a t i o n s between the f i r s t two f a c t o r s were g e n e r a l l y weak f o r beaked h a z e l , aspen and w i l l o w . Twig l e v e l s of N and P showed no s i g n i f i c a n t ' r e l a t i o n s h i p ; potassium and calcium l e v e l s g e n e r a l l y increased w i t h i n c r e a s i n g stand age and 328 canopy c l o s u r e , at l e a s t f o r h a z e l and aspen. However, m u l t i p l e r e g r e s s i o n analyses i n v o l v i n g Ca, N, P and K i n haz e l and aspen were s i g n i f i c a n t l y r e l a t e d to stand age and canopy c l o s u r e . C o r r e l a t i o n s w i t h s o i l n u t r i e n t l e v e l s were not s i g n i f i c a n t except a c o e f f i c i e n t of 0.84 and 0.95 f o r calcium i n ha z e l and phosphorus i n aspen, r e s p e c t i v e l y . However, i t i s probably most c o r r e c t to st a t e that e f f e c t s of s o i l ( n u t r i e n t , moisture, t e x t u r e , etc.) vary w i t h the spec i e s , i . e . , whether they -are stenotrophic or e u r y t r o p h i c . Thus some p l a n t s are u s e f u l i n d i c a t o r s of minerals or concentrate elements, e.g. selenium i n Astragalus spp., while others do not. A u s e f u l l i n e of research would be to t e s t the consistency w i t h which a given species responds to d i f f e r e n t s o i l s . G e n e t i c a l l y i d e n t i c a l stock would be e s s e n t i a l since adjacent p l a n t s of the same species can vary i n t h e i r l e v e l s of crude p r o t e i n and other n u t r i e n t s , e.g., Bergerud and Manuel (1968), Flower - E l l i s (1971). Overstory has ofte n been given as a f a c t o r t h a t i n f l u e n c e s forage n u t r i e n t s . Development of overstory modifies temperatures, reduces l i g h t and t h r o u g h - f a l l p r e c i p i t a t i o n , a l t e r s competitive r e l a t i o n s h i p s , and changes the status of s o i l moisture and n u t r i e n t a v a i l a b i l i t y . A l l these f a c t o r s i n f l u e n c e forage n u t r i e n t s so t h a t overstory -n u t r i e n t i n t e r - r e l a t i o n s h i p s are complex and r e p l e t e w i t h i n t e r a c t i o n s . Not s u r p r i s i n g l y , r e s u l t s of s p e c i f i c f a c t o r s t u d i e s are often confounded. Laycock and P r i c e (1970:39) 329 e x p l a i n the higher l e v e l s of p r o t e i n i n shade by retarded phenology, higher s o i l moisture ( t h i s i s p o s i t i v e l y c o r r e l a t e d w i t h p r o t e i n ) , succulence and p o s s i b l e reduced leachi n g . Again, species s p e c i f i c adaptations must be considered. However, these e f f e c t s are not u n i v e r s a l . In the present study, crude p r o t e i n f o r four species was unaffected where the only d i f f e r e n c e i n s i t e s was canopy cl o s u r e . Peek (1971h) found th a t p r o t e i n i n twig samples from beaked h a z e l , trembling aspen and pussy w i l l o w were not c o r r e l a t e d w i t h canopy c l o s u r e . D i f f e r e n c e s i n n u t r i e n t l e v e l s among p l a n t s of the same species i s a l s o p a r t l y c o n t r o l l e d g e n e t i c a l l y . Although genetic v a r i a b i l i t y i n shrubs i s w e l l recognized and e x p l o i t e d f o r h o r t i c u l t u r a l purposes, the genetic e f f e c t on n u t r i e n t s i s l e s s w e l l s t u d i e d {of. McKell e t a l . 1972). However, Welch et a l . (1977) r e c e n t l y described research i n progress t h a t demonstrated s i g n i f i c a n t d i f f e r e n c e s i n crude p r o t e i n l e v e l s of sagebrush [Artemisia tridentata). For twigs c o l l e c t e d from p l a n t s grown under uniform c o n d i t i o n s , crude p r o t e i n l e v e l s v a r i e d from 8.7.percent to 17.1 percent between i n d i v i d u a l p l a n t s , and from 10.9 percent to 14.2 percent between three subspecies of sagebrush. Presumably, s i m i l a r v a r i a b i l i t y e x i s t s w i t h i n p l a n t s and subspecies of shrubs eaten by moose. These i n d i v i d u a l p l a n t d i f f e r e n c e s may help e x p l a i n why one p l a n t i s used h e a v i l y while 330 adjacent ones of the same species are.not (see a l s o Section ,4.4.2).. A l s o , t h i s n a t u r a l v a r i a t i o n i n n u t r i e n t l e v e l s provides an opportunity to develop v a r i e t i e s high i n crude p r o t e i n and other valuable components. N u t r i e n t v a r i a t i o n s a l s o occur w i t h i n p l a n t s . Two types of d i f f e r e n c e s can be d i s t i n g u i s h e d : between t i s s u e s or organs,, and w i t h i n t i s s u e s or organs. Both these types have n u t r i e n t l e v e l s t h a t o f t e n exceed.those due to s i t e or species. For moose, probably the most r e l e v a n t \"between\" d i f f e r e n c e s i s between twigs and leaves. Leaves have higher p r o t e i n and lower l i g n i n (higher d i g e s t i b i l i t y ) than twigs except j u s t p r i o r to l e a f f a l l (Table 8.4, Aldous 1945, B a i l e y 1967, B l a i r and Epps 1967, Cook and H a r r i s 1950, Dietz 1972, F l o w e r - E l l i s 1971, L i n d l o f 1974, Tew 1970, and many o t h e r s ) . As noted before, t h i s t i s s u e d i f f e r e n t i a l i s l i k e l y one major reason why moose s t r i p leaves i n summer. The p r e d i c t a b l e r e l a t i o n s h i p between l e a f and stem CP (crude protein) i s a p o t e n t i a l l y u s e f u l technique. The r e g r e s s i o n of l e a f CP on stem CP was s i g n i f i c a n t f o r the P r i n c e George species (see p. 296). A n a l y s i s of data i n Dietz (1972) and F l o w e r - E l l i s (1971) a l s o y i e l d e d s t a t i s t i c a l l y s i g n i f i c a n t r e l a t i o n s h i p s , v i z . , y = 9.22 + 0.53x; where n = 12, r 2 = 0.71, S = 1.89 y. x (P < 0.01), and y = -9.19 + 2.49x; where n = 5, r 2 = 0.74, S = 1.69 (P = 0.05), r e s p e c t i v e l y . 331 N u t r i e n t gradients a l s o occur w i t h i n a t i s s u e . These trends may help e x p l a i n why w i n t e r i n g moose i n Pri n c e George and elsewhere w i l l o ften remove only p o r t i o n s of the cur r e n t annual twigs. The exis t e n c e of CP gradients i n n a t i v e browse was known at l e a s t by 1945. Aldous (1945) found t h a t t e r m i n a l p o r t i o n s of cu r r e n t annual growth of b i t t e r b r u s h (Purshia tridentata) contained more CP than proximal, p o r t i o n s . A l s o , s h o r t e r annual shoots contained more CP than long ones. Cowan e t a l . (1970) examined t h i s phenomenon i n greater d e t a i l than d i d Aldous. The t e r m i n a l 2.5 cm of black cherry and red maple contain 2.5 and 1.7 times the 2 3 - 30 cm p o r t i o n , r e s p e c t i v e l y . In a d d i t i o n , t e r m i n a l buds contained approximately .1.5 and 1.3 times the CP as the term i n a l 2.5 cm of twig. Ahlen (19 75) showed s i m i l a r though l e s s marked gradients i n a v a r i e t y of Swedish browse species. Within t i s s u e d i f f e r e n c e s have a l s o been documented f o r other n u t r i e n t components such as c e l l u l o s e (Cowan e t a l . 1970), minerals (Ahlen 1975) and crude f i b e r (Aldous 1945). In general, the l e s s e a s i l y d i g e s t i b l e f r a c t i o n s decrease d i s t a l l y . N u t r i e n t l e v e l s are a l s o a f f e c t e d by the annual age of t i s s u e s , moreso f o r the mobile n u t r i e n t s than f o r s t r u c t u r a l elements such as l i g n i n . a n d c e l l u l o s e . Influence of age was not examined s p e c i f i c a l l y but other s t u d i e s support the statement that w i t h age, crude p r o t e i n d e c l i n e s and l i g n i n i n c r e a s e s . The most r e l e v a n t r e s u l t s were 332 provided by Cowan e t a l . (1950). Winter samples of one, two and three year o l d twigs from h a z e l showed crude p r o t e i n d e c l i n i n g from 8.0 percent to 4.1 percent, a change of 49 percent. S i m i l a r l y , w i n t e r samples of one and two year o l d twigs from upland w i l l o w showed crude p r o t e i n d e c l i n i n g from 7.2 percent to 5.7 percent, a change of 21 percent (the comparable two year change i n h a z e l was 36 percent). In Ala s k a , E l l i s o n (19 76) documented age-dependent decreases i n ni t r o g e n l e v e l s i n black and white spruce. Three year o l d needles of black spruce were 11 percent l e s s than c u r r e n t year's needles, w h i l e f o r white spruce the corresponding d i f f e r e n c e was 24 percent. Undoubtedly, l e a c h i n g caused some of t h i s d e c l i n e (Laycock and P r i c e 1970), although reduced metabolic a c t i v i t y of o l d e r t i s s u e s a l s o was operating. P o s i t i o n on the p l a n t a l s o modified n u t r i e n t l e v e l s . B a i l e y (1967), E l l i s o n (1976) and Tew (1970) demonstrated that crude p r o t e i n l e v e l s of cu r r e n t growth i s p o s i t i v e l y c o r r e l a t e d w i t h height. A l s o , Hoffman (1961) i n d i c a t e d that aspect i n p l a n t s can be important. These p o s i t i o n s e f f e c t s probably r e f l e c t m i c r o c l i m a t i c and l i g h t d i f f e r e n c e s and consequently, t h e i r e f f e c t s w i l l probably vary according to stand f e a t u r e s . Many h a b i t a t manipulation methods a f f e c t n u t r i e n t l e v e l s i n browse. Many of these are as s o c i a t e d w i t h f o r e s t r y p r a c t i c e s and are discussed i n s e c t i o n 10.2. 9. EFFECTS OF FORESTS ON WINTER CLIMATE 9.1 I n t r o d u c t i o n Moose used h a b i t a t s d i f f e r e n t i a l l y throughout the winte r . These temporal and s p a t i a l p a t t e r n s were d e s c r i b e d and documented i n the f o r e g o i n g s e c t i o n s on h a b i t a t u t i l i z a t i o n ( Section 3), food h a b i t s (Section 4), wi n t e r browsing ( S e c t i o n 5), and bed s i t e s e l e c t i o n (Section 6). B r i e f l y , moose began c o n c e n t r a t i n g on w i n t e r ranges i n e a r l y December. From t h i s b u i l d u p phase u n t i l mid-winter, moose use was g r e a t e s t i n deciduous f o r e s t stands, c l e a r c u t s and p a r t i a l c u t o v e r s . During the l a t t e r h a l f of winte r , mixed wood and, e s p e c i a l l y , mature c o n i f e r o u s stands were used most f r e q u e n t l y . As snow melted i n A p r i l and May, moose d i s p e r s e d and began u s i n g both open and f o r e s t e d h a b i t a t s . C l e a r l y , moose made these s h i f t s i n h a b i t a t i n response to a stimulus o r combination of f a c t o r s . Major f a c t o r s were t e n t a t i v e l y i d e n t i f i e d i n the h a b i t a t u t i l i z a t i o n d i s c u s s i o n (Section 3.4.2), v i z . , amount of food, n u t r i t i v e value of food, and wi n t e r c l i m a t e , p a r t i c u l a r l y snow. The two pr e v i o u s s e c t i o n s (7 and 8) examined the f i r s t two p r i n c i p a l f a c t o r s . T h i s s e c t i o n examines w i n t e r c l i m a t e . V i r t u a l l y a l l e c o l o g i c a l s t u d i e s o f moose emphasize 333 334 the importance of winter c l i m a t e . The e a r l y , comprehensive p u b l i c a t i o n s by Formosov (1946) and Nasimovitch (1955) served to focus a t t e n t i o n on the many e f f e c t s of w i n t e r c o n d i t i o n s on moose and other b o r e a l mammals. S h o r t l y a f t e r , r e p o r t s on moose i n winter appeared f o r North America i n i n c r e a s i n g numbers (Edwards and R i t c e y 1956, Des Meules 1964 and 1965, T e l f e r 1967 and 1970, P r e s c o t t 1968, K e l s a l l and T e l f e r 1971, K e l s a l l and P r e s c o t t 1971, Peek 1971, Coady 19 74). The aspects of winter climate that were researched evolved over t h i s p e r i o d . E a r l i e s t s t u d i e s g e n e r a l l y d e a l t only w i t h snow depth. More recent research has demonstrated the need to consider other snow c h a r a c t e r i s t i c s such as d e n s i t y , hardness and compaction, and other c l i m a t i c elements such as temperature, wind, r e l a t i v e humidity and the combined e f f e c t of the l a s t three, c h i l l (Verme 19 68, Verme and Ozoga 1971). Moen (1973) and h i s students have drawn a t t e n t i o n to r a d i a t i o n , and emphasized the need f o r comprehensive examination of a l l c l i m a t i c parameters t h a t modify energy exchange between animals and t h e i r surround-ings . Winter climate a f f e c t s h a b i t a t s e l e c t i o n and use i n three p r i n c i p a l ways. In i n c r e a s i n g l e v e l of d e t a i l , they are: 1) a r r i v a l , occupancy and departure from winter ranges, 2) d i f f e r e n t i a l use between h a b i t a t s , and 335 3) d i f f e r e n t i a l use w i t h i n h a b i t a t s . The f i r s t l e v e l was examined by r e l a t i n g the general p a t t e r n of accumulation and melt of snow wi t h number of moose seen on the three i n t e n s i v e study areas. The second l e v e l was approached by assessing d i f f e r e n c e s i n snow depth and denr s i t y between major h a b i t a t - t y p e s on these winter ranges. F i n a l l y , the t h i r d l e v e l was described by documenting v a r i a b i l i t y i n snow parameters w i t h i n h a b i t a t - t y p e s , and by examining ecotonal climate. 9.2 Methods 9.2.1 The Es t i m a t i o n of Mi g r a t i o n and Occupancy of Winter Ranges Records of snow accumulation were obtained from standard snow courses at open s i t e s on the three i n t e n s i v e study areas. These methods are described i n the next sub-s e c t i o n , 9.2.2. Snowfall records were a l s o taken from the f e d e r a l Atmospheric Environment Service s t a t i o n s and those operated i n 1971-72 and 1972-73 by the Climatology S e c t i o n , Resource A n a l y s i s Branch of the p r o v i n c i a l government's M i n i s t r y of Environment. The c l i m a t i c data were c o r r e l a t e d w i t h i n d i c e s of moose numbers on the i n t e n s i v e study areas. The index used was number of moose seen per minute of f l y i n g . Methods used to c o l l e c t these data were described p r e v i o u s l y (Section 3.2.3) and so w i l l not be repeated here. 336 9.2.2 Snow C h a r a c t e r i s t i c s of Habitat-Types Snow surveys were conducted on the three i n t e n s i v e study areas, during the winters of 1971-72 and 1972-73. Sampling s i t e s were chosen to encompass a wide range of f o r e s t types. These types i n c l u d e d open burns, s e l e c t i v e l y -logged cutovers, deciduous stands, mixed woods, and open and dense c o n i f e r stands. The l o c a t i o n s and s i t e d e s c r i p t i o n s of a l l snow courses are i n Table 9.1. Where p o s s i b l e , l e v e l s i t e s at 750 \u00C2\u00B1 75 m were s e l e c t e d to minimize confounding e f f e c t s of slope, aspect, and e l e v a t i o n . Crown canopy cl o s u r e s were estimated from a e r i a l photographs i n most cases. A l l l o c a t i o n s were used by moose i n winter. The design and sampling of snow courses followed standard methods i n Water I n v e s t i g a t i o n s Branch (1968). Ten e q u i d i s t a n t s t a t i o n s were marked permanently along each snow course to enable accurate r e l o c a t i o n of sampling p o i n t s and so reduce sampling e r r o r . S i t e s were c l e a r e d or l e v e l l e d before s n o w f a l l except i n 19 71-72, when surveys d i d not begin u n t i l January. Snow courses were checked monthly. In 1972-73, sampling began i n October or November, when snow covered the ground. Sampling ended i n A p r i l or May, when a l l snow had melted. Depth and density of snow were measured w i t h a western snow sampler . (Figure 9.1a). Usually, one measurement was made at each s t a t i o n f o r a t o t a l of ten/ site/month. 337 Table 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 H a b i t a t of Snow Courses S i t e (snow course) Sampled i n winter of Study Area No. E l e v a t i o n (m) Ha b i t a t 1971-72 1972--73 Eagle E l 715 spruce-subalpine f i r f o r e s t X X E2 730 spruce-subalpine f i r f o r e s t X E3 700 open burn X X E4 685 p a r t i a l cutover X E5 700 paper b i r c h f o r e s t X Grove Gl 765 south-facing ecotone (burn) X G2 765 south-facing ecotone (pine f o r e s t ) X G3 765 west-facing ecotone (burn) X G4 765 west-facing ecotone (pine f o r e s t ) X G5 1,070 upland burn X G6 765 lowland burn - west s i t e X X G7 760 lowland burn - east s i t e X G8 850 lowland burn - south s i t e X G9 760 spruce-pine f o r e s t X G10 845 immature pine f o r e s t - west s i t e X G10 855 immature pine f o r e s t - east s i t e X G i l 775 spruce f o r e s t X Salmon S I * 680 aspen-pine f o r e s t X X S2 715 pine-spruce f o r e s t X X S3 775 p a r t i a l cutover X S4 535 cottonwood f o r e s t X X S5 670 cottonwood f o r e s t (logged) X S7* 695 aspen f o r e s t X ^Combined i n r e s u l t s . 337a Figure 9.1 Photographs showing the use of the western snow sampler and the penetrometer. 339 To assess snow hardness, surface p e n e t r a b i l i t y was measured, using an instrument employed s u c c e s s f u l l y i n the eastern United States (J. Peek, pers. comm.). The instrument c o n s i s t e d of a m e t a l l i c s h a f t attached to a s p r i n g . The s p r i n g was enclosed i n a c y l i n d e r c a l i b r a t e d i n 10 g i n t e r v a l s on the outer surface. A d i s c was f i t t e d on a small s h a f t that was attached about 1 cm from and p a r a l l e l t o , the c a l i b r a t e d c y l i n d e r . A small rod, fastened p e r p e n d i c u l a r l y to the main s h a f t , p r o j e c t e d out under the d i s c . Thus, when the instrument was pushed i n t o the snow, the s p r i n g compressed i n t o the c y l i n d e r while the small rod moved the d i s c upwards along the c a l i b r a t e d s c a l e . When the s h a f t broke through the snow c r u s t , the s p r i n g expanded and returned the small rod to zero. The d i s c remained i n p l a c e , i n d i c a t i n g the maximum force exerted. This instrument was s i m i l a r t o the N a t i o n a l Research C o u n c i l snow hardness gauge, and more convenient than the r a t c h e t m o d i f i c a t i o n r e f e r r e d to by Coady (1974). A l s o , i t was comparatively inexpensive. Four measurements were made at each s t a t i o n , s i t u a t e d e q u i d i s t a n t around the hole made by the snow sampler. Thus the sample s i z e f o r each snow course was 40 per month. The t e s t e r was pushed v e r t i c a l l y i n t o the snow, up to a maximum depth of 15 cm u n t i l the snow c r u s t broke (Figure 9.1b). Maximum force r e q u i r e d to break through the c r u s t was recorded i n increments of 10 g from 50 g to 1,000 g. In case of m u l t i p l e c r u s t i n g , the maximum force 340 r e q u i r e d over the 15 cm was taken as the surface p e n e t r a b i l i t y . Due to the great v a r i a b i l i t y of the o r i g i n a l measurements, and the measuring l i m i t a t i o n s of the instrument, data f o r surface p e n e t r a b i l i t y were transformed to a r a t i n g s c a l e ( o r d i n a l values as shown i n Appendix Table H - l ) . P e n e t r a b i l i t i e s of l e s s than 50 g and greater than 1,000 g were not recorded by the instrument. The wide v a r i a t i o n , even i n adjacent measurements, i s demonstrated by one t e s t i n a f o r e s t stand on the Salmon study area. Ten measurements made 10 cm apart had values ranging from 0 to > 1,000 g/cm. While t h i s transformation reduced the s e n s i t i v i t y of the measurements to changes i n surface p e n e t r a b i l i t y , I b e l i e v e that meaningful d i f f e r e n c e s were detectable. 9.2.3 Climate of the Forest Edge Climate across ah open-forest ecotone: was.. monitored at 765 m on two s i t e s of the Grove winter range. The main s i t e was a t r a n s e c t ( G l , G2) set perpe n d i c u l a r to a north-south f a c i n g ecotone, between an 11 y r o l d burn and a 100 y r o l d f o r e s t of lodgepole pine. The f i v e s t a t i o n s were s i t u a t e d along the t r a n s e c t , one at the ecotone, and one each at 30 m and 76 m from the ecotone i n both the burn and the f o r e s t . At each s t a t i o n a Stevenson screen was mounted 1.4 m above ground, to house maximum and minimum thermome-K 341 t e r s and a hygrothermograph: one s t a t i o n had only a thermograph (Figure 9.1c). Four t o t a l i z i n g anemometers were used; one each at 30 m and 76 m from the ecotone i n both d i r e c t i o n s . Snow courses were s i t e d at each s t a t i o n , w i t h f i v e measurements per s t a t i o n (2-0 f o r surface p e n e t r a b i l i t y ) . The second s i t e was l o c a t e d across the same ecotone, but on an east-west f a c i n g s e c t i o n . Here, only snow duration and depth were monitored. S i n g l e , 1-cm diameter stakes were placed at the ecotone and perpendicular to i t , along the t r a n s e c t at 15, 76 and 122 m from the ecotone i n the open, and i n the f o r e s t (G3, G4) . For comparative purposes, climate was a l s o monitored at the south end of the Grove burn at 1,050 m on an exposed south-facing slope (G5). Instrumentation was s i m i l a r to the f i r s t s i t e , w i t h a t o t a l i z i n g anemometer,, and a Stevenson that housed maximum and minimum thermometers and a thermograph. Snow d u r a t i o n , depth, d e n s i t y , and surface p e n e t r a b i l i t y was measured along a standard ten s t a t i o n snow course. 9.3 Results 9.3.1 M i g r a t i o n and Snow Accumulation On the study areas, snow accumulation and melt occurred i n three stages: r a p i d b u i l d u p , slow buildup or pl a t e a u , and r a p i d d e c l i n e (Figure 9.2). Snow accumulated 341a Figure 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 the Pr i n c e George a i r p o r t . Data f o r Pr i n c e George was from the Canada Atmospheric Environment S e r v i c e . NOV. DEC. J A N . FEB. MAR. APR. MAY 1972-73 343 r a p i d l y from near zero i n November u n t i l January, when approximately 85 percent of peak depths were a t t a i n e d . In the f o l l o w i n g two months snow cover e i t h e r d e c l i n e d s l i g h t l y (Grove) or increased s l i g h t l y (Eagle and Salmon). A f t e r March, snow melted r a p i d l y , w i t h v i r t u a l l y no snow by e a r l y May. These patterns p a r a l l e l e d longterm ones such as recorded at the P r i n c e George a i r p o r t . The snow c y c l e bore l i t t l e s i m i l a r i t y to the i n d i c e s of moose numbers on the study area, except at Grove (of;.. Figures 3.4 and 9.2) , At Salmon and Eagle, moose were most numerous i n November and December, before d e c l i n i n g s t e a d i l y or d e c l i n i n g to a r e l a t i v e l y s t a b l e l e v e l . At Grove, trends i n moose numbers and the snow c y c l e were g e n e r a l l y a l i k e , although they were not s i g n i f i c a n t l y c o r r e l a t e d (P > 0.05). Thus i f s n o w f a l l was important i n i n i t i a t i n g migration onto w i n t e r ranges, i t was s n o w f a l l at higher e l e v a t i o n s . Several p o i n t s support t h i s idea. In November 1971, migrating moose were being harvested at 700-750 m, where snow depths ranged from 11 - 15 cm. Snow depths at 990-1,06 0 m ranged from 19-2 8 cm. Thus migration began at snow depths much l e s s than the c r i t i c a l 100 cm depth when movements become r e s t r i c t e d by snow. A l s o , when moose i n d i c e s were i n c r e a s i n g estimated snow depths at higher e l e v a t i o n s was l e s s than c r i t i c a l . Snow depths were estimated using the r e g r e s s i o n equation: y = 0.04x - 12.95, 344 where y = snow depth i n cm, x = e l e v a t i o n i n m, S =4.04, y * ^ r 2 = 0.71 and F = 9.90 (P = 0.05). Golding (1969) found e l e v a t i o n and snow-water equi v a l e n t s s t r o n g l y c o r r e l a t e d . Data f o r t h i s equation were gathered on an e l e v a t i o n a l t r a n s e c t on the Salmon study area i n November 19 71. Assuming that t h i s r e l a t i o n s h i p h eld f o r the three study areas, the f o l l o w i n g snow depths were estimated: E l e v a t i o n (m) Snow pack (cm) 500 7 1,000 27 1,500 47 Thus, during November and December, when the moose index suggested t h a t migration onto the winter ranges was l a r g e l y completed, snow pack at higher e l e v a t i o n s was most l i k e l y l e s s than 50 cm. Departure from the w i n t e r ranges apparently occurred g r a d u a l l y over the l a t t e r h a l f of the winter and e a r l y s p r i n g (Figure 3.4). I t i s important to remember t h a t the index of moose seen i s based p r i m a r i l y on moose seen i n the unforested h a b i t a t . Thus a d e c l i n i n g index probably r e f l e c t e d two phenomena, v i z . , a s h i f t from open t o fo r e s t e d h a b i t a t , and departures from the w i n t e r ranges. The moose index d e c l i n e d g r a d u a l l y to a l e v e l i n A p r i l o r May t h a t probably remained r e l a t i v e l y steady u n t i l the f o l l o w i n g w i n t e r . This decrease occurred l a r g e l y e i t h e r before maximum snow depths or before s i g n i f i c a n t snow melt. Thus snow melt and decreases i n the index of moose abundance were poorly c o r r e l a t e d . However, the d e c l i n e i n number of moose seen d i d correspond to changes i n snow i n the open h a b i t a t s . At Eagle, the moose index d e c l i n e d most c o n s i s t e n t l y as the snow pack b u i l t up beyond 80 cm. During t h i s p e r i o d snow d e n s i t i e s i n the f o r e s t exceeded those i n the open. Thus moose apparently responded p r i m a r i l y to snow depth. At Grove, the moose index d e c l i n e d , beginning i n January when snow depths reached 51 cm. At t h i s time, snow density i n the f o r e s t was considerably.below than i n the open. Thus moose at Grove may have been responding to both depth and density d i f f e r e n t i a l s . At Salmon, the moose index d e c l i n e d from 1.2 8 to 0.2 7 moose/min between December and January: snow depth increased from 28 t o 76 cm, r e s p e c t i v e -l y . Forest snow d e n s i t i e s were approximately s i m i l a r to those i n open h a b i t a t . The moose w i n t e r i n g at Salmon, t h e r e f o r e , seemed to respond most c l o s e l y w i t h r a p i d b u i l d -up of snow, much l i k e moose at the Eagle area. 9.3.2 Snow C h a r a c t e r i s t i c s of Habitat-Types Before d e s c r i b i n g the r e s u l t s i n d e t a i l , i t i s important to emphasize that snow c o n d i t i o n s vary annually. These changes importantly modify h a b i t a t u t i l i z a t i o n and can e f f e c t a random, n a t u r a l range r o t a t i o n system ( G i l b e r t e t a l . 19 70). Between wi n t e r v a r i a t i o n was demonstrated 346 c l e a r l y by comparisons of snow depths from March of 19 72, 1973 and 1974, as recorded i n unforested or deciduous f o r e s t h a b i t a t s (Table 9.2). For consecutive March's, the open burn at Eagle had mean depths of 9 7, 104 and 142 cm, r e s p e c t i v e l y . Comparable data f o r the open burn s i t e at Grove were 86, 51 and 114 cm, r e s p e c t i v e l y . At the deciduous f o r e s t s i t e at Salmon, mean snow depths were 104, 89 and 140 cm. Thus the r a t i o of the maximum and minimum snow depths recorded i n March ranged from 1.5 to 2.2. Table 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 Study Area Year Mean (n = depth 10/site) Ratio of maximum CV to minimum depths Eagle 1972 97 \u00C2\u00B1 8* 21% 1973 104 \u00C2\u00B1 5 1.5:1.0 1974 142 \u00C2\u00B1 20 Grove 1972 86 \u00C2\u00B1 8 38% 1973 51 \u00C2\u00B1 5 2.2:1.0 1974 114 \u00C2\u00B1 5 Salmon 1972 104 \u00C2\u00B1 8 24% 1973 89 \u00C2\u00B1 5 1.6:1.0 1974 140 \u00C2\u00B1 5 Mean \u00C2\u00B1 sd. 347 V a r i a t i o n , expressed as c o e f f i c i e n t of v a r i a t i o n (CV), was comparable to that recorded f o r snow f a l l at the P r i n c e George a i r p o r t s t a t i o n . This suggests t h a t the study-years experienced t y p i c a l v a r i a t i o n s i n snow f a l l , although amount of snow d i f f e r e d (Figure 9.2). The snow c h a r a c t e r i s t i c s of depth, evenness of s u r f a c i n g , density and duration v a r i e d widely between h a b i t a t s on the three study areas (Tables 9.3 to 9.7). In comparing snow depth between h a b i t a t s , the t y p i c a l p a t t e r n was f o r greatest depths at the open or deciduous f o r e s t s i t e s , intermediate depths at the p a r t i a l cutover s i t e s , and lowest depths at the coniferous f o r e s t s i t e s . At the Eagle area, t h i s p a t t e r n was c o n s i s t e n t f o r a l l months th a t were sampled except f o r A p r i l i n 1971-72 and f o r May i n 1972-73 (Table 9.3). Moreover, the r e l a t i v e d i f f e r e n c e between h a b i t a t s was a l s o c o n s i s t e n t . Snow depths at the f o r e s t and p a r t i a l cutover s i t e s were approximately 5 0 percent and 75 percent of depths at the open s i t e , r e s p e c t i v e l y . At the Grove area, the two coniferous f o r e s t h a b i t a t s always had l e s s snow than both the upland and lowland burn h a b i t a t s (Table 9.4). A l s o , the immature f o r e s t had l e s s snow than the mature one. The t y p i c a l p a t t e r n was obscured on the Salmon winter range. Thus the p a r t i a l cutover had more rather than l e s s snow than the deciduous f o r e s t ; the coniferous f o r e s t had comparable depths to the deciduous f o r e s t r a t h e r than Table 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 Snow depth (mean \u00C2\u00B1 sd, i n cm) Snow density (mean \u00C2\u00B1 sd, i n g/cm^)* Habitats H a b i t a t s Winter and open p a r t i a l c o n i f e r open p a r t i a l c o n i f e r month burn cutover f o r e s t burn cutover f o r e s t 1971-72: Feb 117 \u00C2\u00B1 \u00E2\u0080\u00A2 8 nd 94 \u00C2\u00B1 1 3 + .27 \u00C2\u00B1 .03 nd .28 \u00C2\u00B1 . 02 + Mar 97 \u00C2\u00B1 + nd 89 \u00C2\u00B1 10** .32 \u00C2\u00B1 .07\"* nd .43 \u00C2\u00B1 .06** Apr 23 \u00C2\u00B1 1 8 * nd 48 \u00C2\u00B1 15** .47 \u00C2\u00B1 .02 4 nd .45 \u00C2\u00B1 .04** 1972-73: Dec 30 \u00C2\u00B1 3 20 \u00C2\u00B1 10 15 \u00C2\u00B1 2 nd nd nd Jan 81 \u00C2\u00B1 10 63 \u00C2\u00B1 18 43 \u00C2\u00B1 10 .33 \u00C2\u00B1 .04 .36 \u00C2\u00B1 .04 .44 \u00C2\u00B1 .17 Feb 91 \u00C2\u00B1 5 66 \u00C2\u00B1 13 43 \u00C2\u00B1 5 .36 \u00C2\u00B1 .03 .36 \u00C2\u00B1 .05 .39 \u00C2\u00B1 .05 Mar 104 \u00C2\u00B1 5 81 \u00C2\u00B1 18* 51 \u00C2\u00B1 10 .42 \u00C2\u00B1 .04 .42 \u00C2\u00B1 .04* .41 \u00C2\u00B1 .03 Apr 76 \u00C2\u00B1 10 61 \u00C2\u00B1 18 33 \u00C2\u00B1 8 .55 \u00C2\u00B1 .06 .49 \u00C2\u00B1 .10 .56 \u00C2\u00B1 .06 May 0 15 \u00C2\u00B1 13 0 0 .40 \u00C2\u00B1 .04 0 1973-74: Mar 142 \u00C2\u00B1 8 124 \u00C2\u00B1 18 107 \u00C2\u00B1 10 .40 \u00C2\u00B1 .04 .39 \u00C2\u00B1 .04 .40 \u00C2\u00B1 .03 Study s i t e s E2 E3 E4 E2 E3 E4 *n = 10 f o r each mean except n = 13 at **, n = 6 at +, n = 11 at ++, n = 8 at +4-4-, and n = 9 at *. 00 Table 9.4 Monthly Snow Depths f o r Four H a b i t a t s on the Grove Winter Range Snow depths.by h a b i t a t (mean \u00C2\u00B1 sd, i n cm) Winter and spruce-pine immature pine month lowland burn upland burn f o r e s t f o r e s t 1971-72: Feb 91 + 8* nd 61 + 5** 56 \u00C2\u00B1 3 Mar 86 + 8* nd 61 + 5 36 \u00C2\u00B1 5 Apr 20 + 13* nd 18 + 8* 0 1972-73: Dec 13 + 5 10 + o*** 8 + 3 nd Jan 51 + 2 36 + 2*** 28 + 5 nd Feb 43 + 2 23 \u00C2\u00B1 5 18 + 8 nd Mar 51 + 5 38 \u00C2\u00B1 5 15 + 10 nd Apr 36 + 8 33 \u00C2\u00B1 5 5 + 5 nd May 0 0 0 nd 1973-74: Mar 114 5* nd 94 5 nd n = 10 f o r a l l means except n = 20 at *, n = 16 at **, and n = 5 at ***. 350 Table 9.5 Monthly Snow D e n s i t i e s f o r Four H a b i t a t s on the Grove Winter Range Snow d e n s i t i e s by h a b i t a t (mean \u00C2\u00B1 sd, i n g/cm ) .Winter and spruce-pine immature pine month lowland burn upland burn f o r e s t f o r e s t 1971-72: Feb .28 1 .02* nd .27 \u00C2\u00B1 .02** .30 03 Mar .34 \u00C2\u00B1 .03* nd .39 \u00C2\u00B1 .06* .58 \u00C2\u00B1 . 15 Apr .47 \u00C2\u00B1 .07** nd .44 \u00C2\u00B1 .37* .34 \u00C2\u00B1 . 16** 1972-73: Dec nd nd nd nd Jan .32 \u00C2\u00B1 .02 .04 + 0 2 + .20 \u00C2\u00B1 .03 nd Feb nd .30 + . 08 nd nd Mar .36 \u00C2\u00B1 .02 .26 \u00C2\u00B1 . 03 .30 \u00C2\u00B1 .08 nd Apr .53 \u00C2\u00B1 .09 .37 + . 05 .40 1 .10 + nd May 0 0 0 nd 1973-74: Mar .34 \u00C2\u00B1 .03 .34 05 .37 \u00C2\u00B1 .02 nd n = 10 f o r a l l means except n = 20 at *, n = 16 at **, n = 3 at ***, and n = 5 at +. Table 9.6 Monthly Snow Depths f o r Five H abitats at the Salmon Winter Range Winter and month Snow depths by h a b i t a t (mean \u00C2\u00B1 sd, i n cm) 1 aspen-pine f o r e s t logged cottonwood* pine-spruce f o r e s t cottonwood-spruce p a r t i a l cutover 1971-72: Feb 119 \u00C2\u00B1 5* nd 107 + 3 nd nd Mar 104 1 8* nd 99 + 5 nd nd Apr 56 \u00C2\u00B1 10* nd 66 + 8 18 + 10 nd 1972-73: Dec 28 \u00C2\u00B1 3 nd 23 + 5 13 \u00C2\u00B1 5 .281+ 8 Jan 76 \u00C2\u00B1 8 66 1 13 58 + 18 41 1 8 94 \u00C2\u00B1 8 Feb 74 \u00C2\u00B1 8 74 \u00C2\u00B1 5 66 + 10 51 \u00C2\u00B1 8 104 \u00C2\u00B1 5 Mar 89 \u00C2\u00B1 5 84 + 8 81 + 8 38 1 8 114 \u00C2\u00B1 10 Apr 61 1 5 58 + 13 66 + 8 15 \u00C2\u00B1 8 91 + 10 May 0 0 20 + 8 0 20 \u00C2\u00B1 13 1973-74: Mar 140 1 5 nd 127 + 5 81 + 10 160 \u00C2\u00B1 5 Study s i t e SI S5 S2 S4 S3 \"*\"n = 10 f o r a l l means except f o r n = 20 at +, and n = 25 at *. * P a r t i a l l y logged. Table 9.7 Monthly Snow Den s i t i e s f o r Five H a b i t a t s at the Salmon Winter Range Snow d e n s i t i e s by h a b i t a t (mean \u00C2\u00B1 sd, i n g/cm ) aspen-pine logged pine-spruce ' cottonwood- p a r t i a l Winter and month f o r e s t cottonwood* f o r e s t spruce cutover 1971-72: Feb .27 1 .01* nd .27 \u00C2\u00B1 . 03 nd nd Mar .37 \u00C2\u00B1 .06* nd .41 08 .47 1 .12 nd Apr .46 1 .03* nd .43 + . 03 .56 1 .36 nd 1972-73: Dec nd nd nd nd nd Jan .39 1 .05 .39 + .03 .38 \u00C2\u00B1 . 04 .38 1 .04 .36 1 .05 + Feb .39 \u00C2\u00B1 .05 .43 \u00C2\u00B1 .06 .41 \u00C2\u00B1 . 08 .35 \u00C2\u00B1 .04 .34 \u00C2\u00B1 .02 Mar .47 1 .02 .44 \u00C2\u00B1 .04 .44 \u00C2\u00B1 . 08 .41 1 .04 .40 \u00C2\u00B1 .07 Apr .48 \u00C2\u00B1 .03 .46 \u00C2\u00B1 .03 .47 \u00C2\u00B1 . 06 .49 1 .07 .48 \u00C2\u00B1 .06 May .40 \u00C2\u00B1 07 .42 \u00C2\u00B1 .04 1973-74: Mar .37 1 .01 nd .40 \u00C2\u00B1 . 05 .38 1 .03 .37 \u00C2\u00B1 .03 n = 10 f o r a l l means except f o r n = 20 at *, n = 9 at **, and n = 25 at +. * P a r t i a l l y logged. OO ho 353 one-half as much. The most l i k e l y e x p l a n a t i o n s f o r these i n c o n s i s t e n c i e s were that the s i t e s were not a l l at the same e l e v a t i o n , and t h a t they were s i t u a t e d i n d i f f e r e n t s n o w f a l l regimes. For example, the p a r t i a l cutover s i t e was a t 775 m while the deciduous f o r e s t was a t 680 m (Table 9.1). The s i t e s were separated at l e a s t by 20 km i n an area where s n o w f a l l and o t h e r c l i m a t i c elements are changing. However, at two s i t e s where e l e v a t i o n and d i s t a n c e were of i n s i g n i f i -c ant e f f e c t , the aspen-pine f o r e s t and logged cottonwood, the expected p a t t e r n was r e c o r d e d (Table. 9.6). In comparing the r e s u l t s , d i f f e r e n c e s i n snow depths appeared to p a r a l l e l d i f f e r e n c e s i n c l o s u r e of the f o r e s t crown-canopy. The evenness or u n i f o r m i t y of the snow s u r f a c e v a r i e d between h a b i t a t s , e s p e c i a l l y i n the e a r l y p a r t of the w i n t e r . Assuming t h a t unevenness i s r e f l e c t e d by the c o e f f i c i e n t of v a r i a t i o n (CV), the data from the s i t e s a t the Eagle area showed CV of e i g h t , 17 and 50 p e r c e n t s , r e s p e c t i v e l y , f o r the open burn, f o r e s t and p a r t i a l cutover. As the snow s e t t l e d i n l a t e w i n t e r , the d i f f e r e n c e between h a b i t a t d e c l i n e d w i t h corresponding CV f o r A p r i l of 13, 2 3 and 2 9 p e r c e n t s . At the Grove area, the open h a b i t a t s had a more even snow s u r f a c e (lower CV) than f o r e s t e d ones (Table 9.4). The data from the Salmon w i n t e r range p a r a l l e l those from the Eagle range (Table 9.6). The t h i r d snow f e a t u r e examined, snow d e n s i t y , d i d not show obvious p a t t e r n s between h a b i t a t s (Tables 9.3, 9.5 354 and 9.7). At the Eagle area, snow density i n the f o r e s t was u s u a l l y greater than or equal t o d e n s i t i e s i n the open burn and p a r t i a l cutovers. At the Grove area, the tendency was i f o r the opposite, that i s , the density at the open s i t e s exceeded that of f o r e s t e d s i t e s . At the Salmon area, no patterns were d i s c e r n i b l e . Thus snow density i s e i t h e r l a r g e l y unaffected by f a c t o r s such as canopy and e l e v a t i o n , or i s i n f l u e n c e d by f a c t o r s that I d i d not monitor. The amount of r a i n f a l l and i t s i n t e r c e p t i o n by the f o r e s t r y canopy i s one such f a c t o r . The e f f e c t of more moderate temperatures i n the f o r e s t undoubtedly a f f e c t s snow density but i t s impact i s modified by other f a c t o r s . A t o t a l look at heat budgets f o r each h a b i t a t would probably e x p l a i n snow density v a r i a t i o n s . The l a s t snow feature compared was the duration of snow on the ground. Estimates of duration were only approximate since sampling s i t e s were only checked monthly. Nonetheless, the data revealed that snow duration d i d vary w i t h h a b i t a t . For a l l areas, my observations were that open h a b i t a t s are snow covered before f o r e s t e d areas but t h a t the time of disappearance v a r i e d between areas and h a b i t a t s . At the Eagle area, snow stayed longer i n the p a r t i a l cutover than i n e i t h e r the open burn or coniferous f o r e s t . Thus snow i n those h a b i t a t s w i t h the greatest or l e a s t amount of snow melted before that i n a h a b i t a t w i t h an intermediate amount. At the Grove range, the coniferous f o r e s t , the 355 h a b i t a t w i t h the l e a s t snow, was snow-free before the open burn. At the Salmon study area, snow disappeared the l a t e s t from those h a b i t a t s that had.the deepest snow i n l a t e winter. The d u r a t i o n of snow w a s . l e s s . a f f e c t e d by h a b i t a t , and showed a l e s s c o n s i s t e n t p a t t e r n than snow depth. The d i f f e r e n c e i n duration may be explained as f o l l o w s . Snow appears e a r l i e r i n open h a b i t a t s because the f o r e s t canopy i n t e r c e p t s the u s u a l l y l i g h t s nowfalls of e a r l y w i n t e r . Snow disappearance appeared to be r e l a t e d to both f o r e s t canopy and the amount of snow present i n the l a t e w i n t e r . Although the open s i t e s had the most snow, they had the most exposure to s u n l i g h t , r a i n f a l l , probably the two p r i n c i p a l m e l t i n g agents. The f o r e s t s s h i e l d e d t h e i r snow from both of these agents. The p a r t i a l cutover represented an intermediate case. Although t h i s h a b i t a t had' l e s s snow than the open, the presence of a p a r t i a l canopy i n t e r c e p t e d some r a i n and sunshine and consequently retarded snow melt. 9.3.3 Climate of the Forest Edge Winter c l i m a t e was documented most i n t e n s i v e l y at the south-facing ecotone between a lodgepole pine f o r e s t and the 1961 Grove Burn ( s i t e G5). A second s i t e , the Bowron c l e a r c u t provided backup data. A t h i r d one, the Buckhorn s i t e (G5), provided comparative information f o r what I considered to represent one of the harshest c l i m a t i c 356 extremes on a winter range - an upland, exposed and unforested h a b i t a t . Before p r e s e n t i n g r e s u l t s of the ecotonal c l i m a t e , data c o l l e c t e d at the 76 m open s t a t i o n from Grove are compared w i t h data from the P r i n c e George a i r p o r t to i l l u s t r a t e d i f f e r e n c e s and s i m i l a r i t i e s between these two sampling p o i n t s (Table 9.8). Except f o r December, Grove experienced milder temperatures. The d i f f e r e n c e averaged 12 percent. Grove always had lower r e l a t i v e humidity, averaging 81 percent of the P r i n c e George a i r p o r t data. Grove experienced snow depths more than three times those at the a i r p o r t . The d i s p a r i t y was e s p e c i a l l y pronounced i n the l a t t e r h a l f of the w i n t e r . Conversely, the a i r p o r t experienced almost f i v e times as much wind run as Grove. To summarize, the Grove s i t e was warmer, and experienced more snow while the P r i n c e George s i t e was w i n d i e r and more humid. Temperature v a r i e d across the ecotone, but not i n a s t r a i g h t - f o r w a r d manner (Figure 9.3, Appendix Table H-2). Both highest and lowest mean temperatures were recorded i n the open at the 76 m and 15 m s t a t i o n s , r e s p e c t i v e l y . Temperatures at the ecotone and i n the f o r e s t stand were intermediate, averaging s l i g h t l y lower than those i n the open. Graphical p r e s e n t a t i o n of the data demonstrated that moose could enjoy temperatures at the ecotone comparable to those f u r t h e r i n t o the burn. The p a t t e r n of the temperature gradient was e s s e n t i a l l y the same f o r a l l w i n t e r months. 357 Table 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 P r i n c e George A i r p o r t Month (1972-73) C l i m a t i c parameter Nov. Dec. Jan. Feb. Mar. Apr. May Mean temp. (C) PG -2 -5 -9 -6 1 4 10 G -1 -10 -7 -3 2 5 8 Re l . hum. (%) PG 85 79 69 71 69 58 57 G m* 63 59 56 52 39 55 Snow depth (cm) PG 0 7 13 10 13 0 0 G t * 10 48 43 51 30 0 Windage (km) PG 231 247 332 251 228 189 224 G m 33 64 39 46 66 m :m = missing, t = trace amounts. T h a t f o r e s t t e m p e r a t u r e s a v e r a g e d l e s s t h a n t h o s e i n t h e open was s u b s t a n t i a t e d b y d a t e f r o m t h e Bowron c l e a r c u t ( T a b l e 9 . 9 ) . The mean f o r e s t t e m p e r a t u r e was a p p r o x i m a t e l y 15 p e r -c e n t l o w e r t h a n t h e open mean. The c o m p a r a b l e p e r c e n t a g e d i f f e r e n c e a t t h e G r o v e s t u d y a r e a was 6 p e r c e n t . R e l a t i v e h u m i d i t y was g r e a t e r i n a n d a d j a c e n t t o t h e f o r e s t s t a n d t h a n i n t h e open ( F i g u r e 9.3, A p p e n d i x T a b l e H - 2 ) . F o r m o s t m o n t h s , l o w e s t h u m i d i t y was i n t h e open a n d h i g h e s t h u m i d i t y a t t h e e c o t o n e . S i m i l a r t o t e m p e r a t u r e , p e a k v a l u e s w e r e r e c o r d e d a t t h e e c o t o n e . A g a i n , d a t a f r o m t h e Bowron s i t e p a r a l l e l e d t h o s e f r o m t h e G r o v e : r e l a t i v e 357a Figure 9.3 Some temperature and r e l a t i v e humidity gradients across the forest-open ecotone at the Gove study area during the 1972-73 winter. Table 9.9 Mean Monthly Temperature, R e l a t i v e Humidity, Snow Pack, Wind f o r Clearcut and Adjacent Forest S i t e s * at the Bowron Study Area, 19 72-73 Data** and Temperature (C) R e l a t i v e humidity (%) Snowpack (cm) Wind (km/h) Month c l e a r c u t f o r e s t c l e a r c u t f o r e s t c l e a r c u t f o r e s t c l e a r c u t f o r e s t Aug 13.8 12. 8 Sept 6.9 6.6 Oct 2.6 2.0 Nov -4.7 -2.4 Dec*** -5.8 -5.8 28 Jan*** -3.0 -2. 7 17 Feb -5.9 -6.5 3 Mar 0.2 -0.1 8 15 Apr 3.8 3.3 4 17 May 9.8 9.2 59 63 Jun 11.6 11.2 64 71 J u l \u00E2\u0080\u00A2 15.8 14.9 59 55 4.6 0.2 Aug 13.0 12.3 61 72 4.2 0.4 Means 4.9 4.2 61 65 6 16 4.4 0.3 *S t a t i o n s were located m from the f o r e s t edge i n t o the c l e a r c u t and f o r e s t stand. Stand height was **Data s u p p l i e d courtesy of J . Po w e l l , Canadian Forestry S e r v i c e , Edmonton, A l b e r t a and G. Cheeseman, Climatology S t a t i o n , Resource A n a l y s i s Branch, B.C. M i n i s t r y of Environment, P r i n c e George. ***Incomplete records f o r December (15-31 o n l y ) , and f o r January (1, 13-31 o n l y ) . 360 humidity i n the f o r e s t u s u a l l y exceeded t h a t of the open. The percentage d i f f e r e n c e s , based on humidity values i n the open, were 2 6 percent and 7 percent f o r the Grove and Bowron s i t e s , r e s p e c t i v e l y . The f o r e s t was an e f f e c t i v e b a r r i e r t o wind (Figure 9.4, Appendix Table H-3). At the Grove study s i t e , wind run i n the f o r e s t averaged 52 percent of that i n the open f o r the p e r i o d from December 1972 to A p r i l 1973. Least wind run was recorded w i t h i n 15 m of the ecotone, both i n the open (33 percent) and i n the f o r e s t (29 percent). Thus the f o r e s t stand created a zone of minimal wind run i n a band that s t r a d d l e d the ecotone. Seventy-six m i n t o the f o r e s t , wind run had increased 8 3 percent over the run at 15 m i n t o the stand. S i m i l a r l y , by 76 m i n t o the open, wind run had increased 200 percent over the run at 15 m i n t o the open. L i m i t e d summer data from the Bowron s i t e a l s o showed the p r o t e c t i v e e f f e c t of f o r e s t s on wind. Wind run i n the f o r e s t averaged 7 percent that i n the open. The d i f f e r e n c e s between Grove and Bowron s i t e s demonstrated that the s i z e and probably the shape of an opening i n f l u e n c e s the i n t e r a c t i o n of wind and f o r e s t s . Comparison of wind run at the Buckhorn s i t e (95) on an upland (1,0 70 m) exposed area w i t h the ecotone s i t e a l s o p o i nted to the geographic v a r i a b i l i t y of wind. For the three periods f o r which comparable data e x i s t e d , wind run Buckhorn was 256 percent, 266 percent, and 205 percent 360a Figure 9.4 Wind run and snow depths across the f o r e s t -open ecotone at the Grove study area during the 1972-73 winter. See Figure 9.3 caption f o r l o c a t i o n s of instruments. 362 greater than data f o r the 76 m open s i t e at Grove. Snow features were also modified by the f o r e s t . In the e a r l y w i n t e r , snow depths i n the f o r e s t and i n the open were e i t h e r s i m i l a r (south-facing s i t e ) or l e s s i n the f o r e s t (west-facing s i t e ) (Figure 9.4, Table 9.10 and Appendix Table H-4). For the r e s t of the winter months, snow depth i n f o r e s t s were p r o p o r t i o n a t e l y l e s s and l e s s than depths i n the open. By e a r l y A p r i l , the p r o p o r t i o n was approximately 15 - 18 percent f o r both s i t e s , and by e a r l y May a l l snow had melted i n both the open and i n the f o r e s t . Snow pack was most v a r i a b l e i n the f o r e s t . Snow accumulation patterns d i f f e r e d between the f o r e s t and the adjacent unforested areas. Maximum f o r e s t snow pack occurred i n January and then d e c l i n e d s t e a d i l y throughout the r e s t of the win t e r . By comparison, maximum snow cover i n the open was reached i n January and remained r e l a t i v e l y steady u n t i l March. Snow then melted r a p i d l y . I t appeared that i n the l a s t h a l f of w i n t e r , f a l l i n g snow was e f f i c i e n t l y i n t e r c e p t e d by the f o r e s t canopy, while the snow on the ground i n the f o r e s t melted or compacted or both. Maximum f o r e s t snow depths were about 6 0 percent of maximum open snow depths f o r both the ecotone s i t e s . At the f o r e s t edge, snow depth was g e n e r a l l y most s i m i l a r to open snow depths. Snow depth at the ecotone (both s i t e s ) averaged 9 0 percent of snow cover 15 m i n the open but 2 33 percent of snow cover 15 m i n the f o r e s t . The 363 Table 9.10 Mean Monthly Depth, Density, and Penetrance of Snow Across the South-Facing, Forest-Burn Ecotone at the Grove Study Area, 1972-73 Winter Month Distance i n t o f o r e s t Forest edge Distance i n t o open 76 m 15 m 15^ m. 76 m SNOW DEPTH (cm)* Nov t t . t t t Dec 13 \u00C2\u00B1 3 13 \u00C2\u00B1 5 15 \u00C2\u00B1 3 13 \u00C2\u00B1 8 10 \u00C2\u00B1 5 Jan 28 \u00C2\u00B1 5 30 \u00C2\u00B1 5 46 \u00C2\u00B1 5 51 \u00C2\u00B1 3 48 \u00C2\u00B1 3 Feb 15 \u00C2\u00B1 3 23 \u00C2\u00B1 8 38 \u00C2\u00B1 5 43 \u00C2\u00B1. 3 43 \u00C2\u00B1 3 Mar 13 \u00C2\u00B1 8 20 \u00C2\u00B1 10 43 \u00C2\u00B1 8 48 \u00C2\u00B1 8 51 \u00C2\u00B1 3 Apr 5 \u00C2\u00B1 5 5 \u00C2\u00B1 5 23 \u00C2\u00B1 5 ' 38 \u00C2\u00B1 3 30 \u00C2\u00B1 8 SNOW DENSITY (g/cm 3) Nov t t t t t Dec nd nd nd nd nd Jan 0.34 \u00C2\u00B1 0.02 0.39 \u00C2\u00B1 0. 02 0.29 \u00C2\u00B1 0.01 0 .31 \u00C2\u00B1 0.02 0 .32 \u00C2\u00B1 0.01 Feb m m m m m Mar 0.25 \u00C2\u00B1 0.09 0.35 \u00C2\u00B1 0. 03 0.36 \u00C2\u00B1 0.03 0 .36 \u00C2\u00B1 0.03 0 .36 \u00C2\u00B1 0.01 Apr 0.43 \u00C2\u00B1 0.12 0.41 \u00C2\u00B1 0. 02 0.46 \u00C2\u00B1 0.06 0 .57 \u00C2\u00B1 0.08 0 .48 \u00C2\u00B1 0.08 SNOW PENETRANCE (1-11 r a t i n g ) * * Nov 0 0 0 0 0 Dec nd nd nd nd nd Jan nd nd nd nd nd Feb 11(3-11) 11(3-11) 5(3-11) 4(2-11) 5(2-11) Mar 11(4-11) 7(3-11) 5(2-11) 6(3-11) 5(3-11) Apr 3(0-11) 0(0-11) 4(1-8) 2(1-10) 2(1-11) *Mean \u00C2\u00B1 SD (n = 5). **Median (n = 20). 364 r e s u l t s i n d i c a t e d a rat h e r sharp t r a n s i t i o n i n snow cover across the ecotone. The l i m i t e d data on snow den s i t y and p e n e t r a b i l i t y a l s o r e f l e c t e d f o r e s t - r e l a t e d d i f f e r e n c e s . In e a r l y w i n t e r , snow density i n the f o r e s t was greater than i n the open. By l a t e w i n t e r , the reverse was t r u e . The values f o r penetrance were always greatest i n the f o r e s t u n t i l A p r i l when d i f f e r e n c e s between the s i t e s were not detectable (Table 9.10 and Appendix Table H - l ) . The snow c h a r a c t e r i s t i c s across the forest-burn ecotone can be summarized as f o l l o w s : 1. A coniferous f o r e s t reduces snow cover by over one h a l f . This s h i e l d i n g e f f e c t extends to the ecotone. The snow cover of the adjacent openings can be reduced by approximately 10 percent w i t h i n 15 m of the ecotone. Snow cover i n the f o r e s t i s more v a r i a b l e than i n the open. 2. Snow dens i t y i s s i m i l a r l y i n f l u e n c e d by the f o r e s t being l e s s dense i n e a r l y w i n t e r and more dense i n l a t e winter than i n the open. The zone of greatest t r a n s i t i o n i s r e l a t i v e l y narrow, i . e . w i t h i n 46 m of the f o r e s t edge. 3. Snow p e n e t r a b i l i t y i s l e s s i n the open than in.the f o r e s t i n l a t e w i n t e r , but f o r e s t e f f e c t s tend to disappear when snow melt i s w e l l e s t a b l i s h e d . An abrupt t r a n s i t i o n zone a l s o appears to occur f o r p e n e t r a b i l i t y . 9.4 Discussion 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 The i n d i c e s of moose numbers on the i n t e n s i v e study areas showed l i t t l e r e l a t i o n s h i p to snow pack. Moose were on t h e i r w i n t e r ranges even before snow depths at higher e l e v a t i o n s became r e s t r i c t i v e . Therefore snow apparently t r i g g e r s a migr a t i o n response before i t becomes a p h y s i c a l b a r r i e r . The r o l e of snow i n causing migrations has been examined by other workers. Mi g r a t i o n s to winter ranges i n so u t h - c e n t r a l B r i t i s h Columbia c o i n c i d e d w i t h gradual snow build-ups on summer ranges; movements were e s s e n t i a l l y completed once higher e l e v a t i o n s had snow depths of 75 cm (Edwards and Ritcey 1956). Cold temperatures i n t e r a c t e d w i t h snow to hasten these downward movements. S i m i l a r l y , S h i r a s moose g r a d u a l l y moved to wi n t e r h a b i t a t s i n response to i n c r e a s i n g snow on summer ranges in.Montana (Knowlton 1960, Stevens 1970) and Wyoming (Houston 1968). In the l a t t e r study, departures from summer ranges d i d not begin u n t i l snow depths reached 80 cm. Gradual downward movements i n response to 60 cm snow depths apparently occur i n Alaska (Coady 1974). Abrupt downward migration were also recorded w i t h depths of 90 cm and 60 em f o r i n t e r i o r and western i n t e r i o r A l a s k a , r e s p e c t i v e l y . P o s s i b l y i n these l a t t e r i n s t a n c e s , c o l d temperatures or c r u s t i n g ( K r a f f t 19 64) were i n v o l v e d (see Edwards and R i t c e y 1956, above). Russian st u d i e s by Nasimovitch (1955) suggest t h a t m i g r a t i o n can begin at depths of 25 to 45 cm, while Coady (1974) st a t e d that i n Alaska some migrations begin even before snow f a l l s . In most of the above s t u d i e s , however, migrations were c o r r e l a t e d w i t h snow depths around 70 cm. According to Nasimovitch (1955, quoted by K e l s a l l and T e l f e r 1974), moose probably cannot survive i n areas where 7 0 cm or more of s o f t snow i s common and l o n g - l a s t i n g . Departure from the winter ranges appeared unrelated to changes i n the snow pack. This agrees w i t h the l i t e r a -t u r e . While upward s p r i n g migrations are g e n e r a l l y more r a p i d than downward f a l l migrations ( K r a f f t 1964, Edwards and R i t c e y 1956, R i t c e y 1967), the nature of snow cover at departure appears v a r i a b l e . In Scandinavia and Ala s k a , returns to summer range c o i n c i d e d w i t h the appearances of patches of ground ( K r a f f t 1964, Coady 1974); i n B r i t i s h Columbia, w i t h melting snow of from 30 t o 45 cm and up to 125 cm; and i n Montana, w i t h the appearance of snow-free southern exposures and c r u s t s capable of supporting moose. Changes i n other snow parameters such as reduced hardness and lowered d e n s i t i e s , and s p r i n g green-up have also been considered as f a c t o r s l e a d i n g to s p r i n g migration. In general, changes i n snow pack i s an important f a c t o r , but others such as forage a v a i l a b i l i t y a l s o play a r o l e . 367 9.4.2 R e l a t i o n s Between Snow, Habitats and Winter Use Throughout much of i t s range, moose use shrub and deciduous f o r e s t stands in, e a r l y w i n t e r , and dense coniferous stands i n l a t e w i n t e r . In Minnesota f o r example, moose u t i l i z e d dense stands of balsam f i r [Abies balsamea) and spruce (Picea spp.) when a d i f f e r e n t i a l i n snow hardness, d e n s i t y , or depth occurred. This happened i n January when snow depths were l e s s than 51 cm (Peek 1971). Temperature apparently had no e f f e c t on the timing of t h i s s h i f t . S p r u c e - f i r stands were p r e f e r r e d to pine f o r e s t s . In Quebec, moose began u t i l i z i n g small and medium openings i n c o n i f e r stands when snow depths i n cutovers reached 76 to 86 cm (Des Meules 1964). In New Brunswick, heavy use of c o n i f e r stands began w i t h snow depths of 91 cm ( T e l f e r 1968). Also i n New Brunswick, K e l s a l l and P r e s c o t t (1971) noted t h a t at mean snow depths of 9 7 cm and gre a t e r , v i r t u a l l y a l l moose tra c k s were seen under coniferous canopies. In Nova S c o t i a , moose tended to yard as snow depths neared 76 cm (Prescott 19 66). In general, these s h i f t s i n h a b i t a t s occurred when the snow depths reached approximately 90 cm, except i n Minnesota where s i g n i f i c a n t d i f f e r e n t i a l s i n hardness and density c o i n c i d e d w i t h h a b i t a t s h i f t s . In n o r t h - c e n t r a l B r i t i s h Columbia, the s h i f t from open to f o r e s t e d h a b i t a t was c o r r e l a t e d p r i m a r i l y w i t h a r a p i d build-up of snow that approached c r i t i c a l depths. 368 Snow den s i t y appeared to be of l i m i t e d importance, except at the Grove area where snow d e n s i t i e s i n f o r e s t s were notably l e s s than those i n open h a b i t a t s . I t may be that i n high s n o w f a l l areas, moose respond p r i m a r i l y to depth while i n low s n o w f a l l areas, moose respond to other c h a r a c t e r i s t i c s such as density. The r e l a t i o n s h i p s between snow c h a r a c t e r i s t i c s and h a b i t a t s e l e c t i o n are complex f o r b o r e a l ungulates. They are perhaps best i n t e g r a t e d by the g e n e r a l i z a t i o n t h a t through h a b i t a t s e l e c t i o n these ungulates attempt to reduce energy l o s s e s . Thus, moose may move i n t o timber when energy demands req u i r e d to remain i n open h a b i t a t s become too great. Probably the most common f a c t o r causing excessive energy demands i s snow depth: other f a c t o r s such as snow hardness, wind and r a d i a t i o n a l heat l o s s e s operate i n some circumstances, depending upon the magnitude of t h e i r energy d r a i n and the animal's c o n d i t i o n . (Of course, moose use timber f o r other reasons such as escape from predators and i n d i v i d u a l spacing.) Viewed i n t h i s f u n c t i o n a l context, i t i s obvious th a t the elements of winter climate cause various b e h a v i o r a l adjustments so t h a t moose can remain i n a thermoneutral environment. These adjustments w i l l vary vary temporally, g e o g r a p h i c a l l y , and w i t h the age, sex and n u t r i t i o n a l status of the animal. I t i s a l s o apparent t h a t these adjustments w i l l be modified by other f a c t o r s such as the amount and 369 n u t r i t i v e value of a v a i l a b l e food, and the s o c i a l s t a t u s of the animal. For example, a subordinate animal may be denied access to some c r i t i c a l p a r t of w i n t e r h a b i t a t by the presence of a dominant one. The n u t r i t i o n a l h i s t o r y experienced the previous summer probably a l s o has an e f f e c t since i t can modify d i g e s t i v e e f f i c i e n c y i n the subsequent winter (Hebert 1973). Perhaps the most u s e f u l ways to approach e f f e c t s of c l i m a t e on moose i s by studying i t s annual energy r e q u i r e -ments. Once these are understood, parameters of d i f f e r e n t w i n t e r c l i m a t i c environments can be i n t e r f a c e d w i t h the energy demands using s i m u l a t i o n modelling techniques (e.g., R u s s e l l 1976, Hudson e t a l . 1976). Then the c r i t i c a l c l i m a t i c elements and how they might change over the winter could be i d e n t i f i e d . From t h i s , the w i l d l i f e manager could d i r e c t h a b i t a t management p r e s c r i p t i o n s t h a t would amelior-ate e f f e c t s of these c r i t i c a l f a c t o r s . This approach has been employed f o r domestic l i v e s t o c k under bo r e a l c o n d i t i o n s , e.g., Webster (1971). A l i t t l e work of t h i s k i n d has been done f o r white-t a i l deer (Moen 1968, 1973), f o r cariboo and reindeer (Bunnell e t a l . 1973, R u s s e l l 1976), and f o r bighorn sheep (R. Hudson, pers. comm.). Energy requirements of w i n t e r i n g moose, e s p e c i a l l y f o r thermo-regulation have not been determined, however. Gasaway and Coady (1974) developed an i n i t i a l model by drawing e x t e n s i v e l y upon the l i t e r a t u r e f o r 370 other species. Based on Scholander e t a l . (1950) and stud i e s of reindeer (Rangifertarandus) and w h i t e t a i l deer (Odocoileus vivginianus), they b e l i e v e d t h a t metabolic thermoregulation i s l i k e l y a minor energy requirement of moose. However, estimates of energy needed f o r locomotion under v a r y i n g snow c o n d i t i o n s , are l a c k i n g f o r moose. U n t i l s u f f i c i e n t e n e r g e t i c research has been completed, the option i s t o proceed e m p i r i c a l l y and t r y t o define c r i t i c a l f a c t o r s through f i e l d s t u d i e s . I d e a l l y , systems models, lab experimentation and f i e l d s t u d i e s should occur together. Snow depth appeared to be the most important c l i m a -t i c f a c t o r a f f e c t i n g w i n t e r h a b i t a t s e l e c t i o n by sub-boreal moose. This was based on data i n the present study and on the reported l i t e r a t u r e where comparable snow regimes e x i s t . Thus the main concern of moose h a b i t a t management i n the P r i n c e George area should be to reduce the impact of snow depth, rather than the impact of c h i l l , snow hardness, or other f a c t o r s . Before management becomes e f f e c t i v e , the cause and e f f e c t r e l a t i o n s h i p between a l t e r a t i o n s i n f o r e s t mantle and snow depths must be understood. The two aspects presumed most important were the r e l a t i o n s h i p between the f o r e s t canopy and snow depth, and the e f f e c t s of openings on snow ( i n c l u d i n g climate of the ecotone). Before d i s c u s s i n g snow depth and f o r e s t canopy, s e v e r a l remarks should be made regarding methodology and 371 terminology. Golding (1968) noted that e s t i m a t i n g i n t e r c e p t i o n by measuring d i f f e r e n c e s between snow i n the open and i n an adjacent f o r e s t i s hazardous, due p r i m a r i l y to the v a r i a b l e e f f e c t s of wind on canopy-caught snow, and the e f f e c t s of wind turbulence i n , r e d i s t r i b u t i n g snow i n openings. Golding (pers. comm.) documented as much as fo u r -f o l d d i f f e r e n c e s i n snow depths along a c e n t r a l l y - p l a c e d t r a n s e c t i n A l b e r t a n c l e a r c u t s . V a r i a t i o n i n the adjacent f o r e s t was much l e s s marked. Thus the l o c a t i o n of sampling s i t e s i n openings requires considerable care. In my study, data f o r open s i t e s were based on s t a t i o n s w e l l removed from the i n f l u e n c e of the adjacent f o r e s t stand, but the aspect of the ecotone d i d a f f e c t snow depths. Concerning terminology, Golding (1968) d i s t i n g u i s h e d between i n t e r c e p t i o n and i n t e r c e p t i o n l o s s . The former i s snow i n t e r c e p t e d by f o r e s t crowns, while the l a t t e r i s only that snow evaporated from crowns. C l e a r l y , l o s s e s are always l e s s than or r a r e l y equal to i n t e r c e p t i o n . In t h i s t h e s i s , concern was p r i m a r i l y f o r i n t e r c e p t i o n r a t h e r than evaporative l o s s e s . Coniferous f o r e s t s c l e a r l y had l e s s snow on the ground than unforested s i t e s . D i f f e r e n c e s were e s p e c i a l l y pronounced i n the l a t t e r h a l f of the w i n t e r , averaging 64 percent 1 21 percent (n = 6) at maximum depths (February 1972, March i n . 1973 and 1974 at Grove and Eagle areas). Therefore, approximately 36 percent of the s n o w f a l l was 372 i n t e r c e p t e d . This i n t e r c e p t i o n rate compared favourably w i t h other s t u d i e s . Connaughton (1935) recorded a 30 percent i n t e r c e p t i o n i n a ponderosa pine (Pinus ponderosa) stand s i t u a t e d at 1370 m i n Idaho. The i n t e r c e p t i o n r o l e of understory vegetation was a l s o revealed where only 25 per-cent was i n t e r c e p t e d i n a s i m i l a r stand without an under-s t o r y . On northern Vancouver I s l a n d , Jones (1975) observed that snow was about twice as deep i n logged h a b i t a t s as i n adjacent mature f o r e s t s , an i n t e r c e p t i o n of 50 percent. Moen (19 73:72) s t a t e d that a moderately dense coniferous f o r e s t , i n an area r e c e i v i n g 76 - 12 7 cm annual p r e c i p i t a -t i o n , may i n t e r c e p t 15 percent t o 30 percent of the t o t a l w i n t e r p r e c i p i t a t i o n . However, he provided no reference f o r t h i s statement, nor d i d he d i s t i n g u i s h the proportions of snow and r a i n i n the t o t a l p r e c i p i t a t i o n f i g u r e . In Minnesota, Peek (1971b)found t h a t snow depths i n dense coniferous stands averaged 19 percent l e s s than stands w i t h sparse or absent canopies. In New Brunswick, T e l f e r (1970) i n d i c a t e d t h a t dense-canopied stands had 19 percent t o 42 percent l e s s snow than open s i t e s , and K e l s a l l and P r e s c o t t (1971) reported that open snow depths averaged 45 percent deeper than sub-canopy depths. The foregoing estimates of i n t e r c e p t i o n averaged about 33 percent. Meiman (1968) provided many a d d i t i o n a l references documenting f o r e s t canopy i n t e r c e p t i o n of snow. 373 I n t e r c e p t i o n rates vary between f o r e s t types (Tables 9.3, 9.4 and 9.6). In northeastern B r i t i s h Columbia, t h i s was demonstrated by S i l v e r . (1976:75) as f o l l o w s : Average maximum snow pack (cm) Crown Forest type 1972-73 1973-74 canopy cover % Open type 53 75 0 Young aspen 54 (102%)* 71 (95%) 31 Older aspen 53 (100%) 80 (107%) 41 Black spruce 60 (113%) 81 (108%) 44 Mature pine 44 (83%) 64 (85%) 64 Maturing spruce 32 (60%) 53 (71%) 74 *proportion of snow depth i n the open type. C l e a r l y , deciduous f o r e s t s i n t e r c e p t e d l i t t l e snow while coniferous f o r e s t s were q u i t e v a r i a b l e . These data suggest t h a t a coniferous f o r e s t canopy does not e f f e c t i v e l y i n t e r -cept snow u n t i l c losure exceeds 44 percent, as determined by dot and g r i d counts of photographs (Brown and Worley(1962) method) . I t a l s o appears t h a t f o r e s t types and years i n t e r a c t (see above data from S i l v e r (1976)). These probably r e f l e c t d i f f e r e n c e s i n snow q u a l i t y , and i n a s s o c i a t e d phenomena such as wind v e l o c i t y and d i r e c t i o n , temperature, and r a i n f a l l . However, the major f a c t o r appears to be associa^. ted w i t h the f o r e s t type i t s e l f . P r e d i c t i n g i n t e r c e p t i o n therefore r e quires a s t r a t i f i c a t i o n of types, since these vary i n t h e i r . i n t e r c e p t i o n a b i l i t i e s . A necessary pre-374 r e q u i s i t e i s to define c l i m a t i c zones t h a t are broadly-homogeneous (see Chapman (1955) f o r t h i s p r o v i n c e ) . Secondly, a r e l i a b l e r e l a t i o n s h i p between commonly or e a s i l y determined stand parameters and i n t e r c e p t i o n i s needed to account f o r changes over time. For example, the i n t e r c e p t i o n c h a r a c t e r i s t i c of a 100 year o l d lodgepole pine stand i s l i k e l y d i f f e r e n t from when, i t was 50 years o l d . S e l e c t i n g a r e l i a b l e stand parameter has r e c e i v e d some study. On northern Vancouver I s l a n d , Jones (1975) used crown c l o s u r e as determined from photographs w i t h a 17 mm fisheye lens to p r e d i c t snow depths i n f o r e s t s from open depths (P = 0.04). The U.S. Army (1956, quoted by Moen 19 7 3) a l s o used canopy coverage to derive a standard i n t e r c e p t i o n rate f o r the north-western United States. This standard must provide only a general i n d i c a t i o n , given the c o m p l e x i t i e s of i n t e r c e p t i o n and the climate i n t h i s mountainous area. I examined t h i s p o s s i b i l i t y w i t h data c o l l e c t e d on bed s i t e s (Section 6) and found a weak though 2 s i g n i f i c a n t r e l a t i o n s h i p of y = 65.82-0.49x, where r = 0.15, S = 25.54, n = 33, F r a t i o . = 5.40; and x = canopy clo s u r e and y = snow depth. One of the major problems f a c i n g the use of crown canopy closure i s i n choosing the best technique to measure, i t , t h a t i s , one t h a t i s meaning-f u l and r e s i s t a n t to observer-dependent v a r i a t i o n s . The photographic method i s promising but i s not commonly 375 measured i n f o r e s t surveys. Stand b a s a l area i s another candidate. I t i s a standard f o r e s t measurement that can be o b j e c t i v e l y obtained. Using the B i t t e r l i c h point-sampling method, Golding and Harlan (19 72) found that snow water equ i v a l e n t s were s i g n i f i c a n t l y c o r r e l a t e d w i t h b a s a l area i n a r e l a t i v e -l y uniform stand of Engelmann spruce and subalpine f i r . I examined t h i s p o s s i b i l i t y using data c o l l e c t e d from the McKenzie area but found no s i g n i f i c a n t r e l a t i o n s h i p . My data base was q u i t e l i m i t e d so that t h i s r e s u l t should be confirmed w i t h f u r t h e r sampling and p o s s i b l y a r e g r e s s i o n model th a t u t i l i z e s s e v e r a l other v a r i a b l e s . I n t e r c e p t i o n i s a f f e c t e d by f a c t o r s i n a d d i t i o n to f o r e s t canopy. Golding (1968) l i s t s these as stand height, depth of canopy, and t e x t u r e or roughness of the surface canopy. Developing r e g r e s s i o n models to i n c l u d e some of these would l i k e l y account f o r more of the v a r i a t i o n i n i n t e r c e p t i o n than s o l e l y f o r e s t canopy. However, Peek (1971b)found no c o n s i s t e n t d i f f e r e n c e s i n snow depth that were a t t i b u t a b l e to stand height i n Minnesota. I n c l u s i o n of m e t e o r o l o g i c a l and p h y s i o g r a p h i c a l f a c t o r s would a l s o improve the models by i n c r e a s i n g t h e i r geographic a p p l i c a b i l i t y . Golding (1974) accounted f o r 48 percent of the v a r i a t i o n i n snow water e q u i v a l e n t values by i n c l u d i n g f a c t o r s of stand, physiographic and weather. S i m i l a r m u l t i p l e component models have, explained from 80 percent to 376 92 percent of snow depth v a r i a t i o n s (Anderson 1967, and Packer 1962, r e s p e c t i v e l y ) . In a d d i t i o n to the e f f e c t of f o r e s t canopy on the average i n t e r c e p t i o n of snow, trees a l s o modify the evenness of the snow surface. The Eskimo term \"quamaniq\" i s used to describe the t y p i c a l depressions i n the snow surface found under t r e e s . Des Meules (1964.) examined t h i s phenomenon i n Quebec and found that the depth and diameter of quamaniqs were a f f e c t e d by crown width, height above the ground of the lower edge of the crown, and whether a tre e was coniferous or deciduous. In areas of deep snow, quamaniqs are obviously e x p l o i t e d by moose. I observed a p a r t i c u l a r l y s t r i k i n g example of t h i s at Limestone Creek where snow depths i n the open exceeded 100 cm yet the bases of trees were snow-free. Tracks i n the snow revealed that moose moved from one quamaniq to another i n s t r a i g h t l i n e s , r a t h e r than i n the meandering p a t t e r n t y p i c a l of moose i n areas w i t h l e s s snow. In these heavy snow areas, unplanned logging would have considerable adverse consequences on w i n t e r i n g moose. These v a r i a t i o n s i n the snow surface appear t o play an important r o l e i n h a b i t a t s e l e c t i o n w i t h i n h a b i t a t s , t h a t i s , s e l e c t i o n of mi c r o h a b i t a t s . Many observations i n the Princ e George f o r e s t s i n d i c a t e that use by moose of an apparently uniform f o r e s t type or h a b i t a t i s d i s t i n c t l y non-random. This s e l e c t i o n of m i c r o s i t e s deserves f u r t h e r study 377 to p r o p e r l y evaluate the bases of s e l e c t i o n , and the r e l a t i v e r o l e s of snow depth and hardness, food supply and other f a c t o r s . Snow depths i n one.habitat are also i n f l u e n c e d by adjacent h a b i t a t s (see Section 9.3.3). This has important i m p l i c a t i o n s i n some logged areas because snow depth, d i s t r i b u t i o n , and duration can, be modified through the s i z e , shape and d i s t r i b u t i o n of cutovers. Regarding the d e p o s i t i o n and melt of snow i n f o r e s t openings, much h y d r o l o g i c a l research i s a p p l i c a b l e . However, a major problem i s wind since wind d i r e c t i o n i s not c o n s i s t e n t during e i t h e r s n o w f a l l or l a t e r r e d i s t r i b u t i o n . A r e l a t i o n s h i p between a north wind and snow depth i n open and adjacent f o r e s t s i s d i f f e r e n t from one w i t h an east wind. At best, l o c a t i o n , o r i e n t a t i o n s i z e , and shape of cutover should be r e l a t e d t o the p r e v a i l i n g winds t h a t are mostly responsible f o r b r i n g i n g snow and r e d i s t r i b u t i n g i t . This poses immediate problems f o r some areas such as P r i n c e George where wind d i r e c t i o n s are v a r i a b l e . A f u r t h e r complication i s the i n f l u e n c e of topography on wind. For example, a p r e v a i l i n g west wind at a climate s t a t i o n (the source of information) may be d e f l e c t e d to a n o r t h e r l y one at the study area. Thus an understanding of orographic e f f e c t s i s necessary to i n t e r p r e t and apply weather s t a t i o n records to the area of i n t e r e s t . The above d i f f i c u l t i e s notwithstanding, there are 378 some c o n s i s t e n t r e l a t i o n s h i p s . In an A l b e r t a study, Swanson and Stevenson (1971), greatest snow accumulation occurred i n c i r c u l a r openings twice the diameter of the mean height of the adjacent f o r e s t (H). (The range i n diameters was from 0.25 H to 6 H.) A b l a t i o n rates were slowest i n openings < 1 H i n s i z e and increased w i t h i n c r e a s i n g opening s i z e above 1 H. Snow accumulation rates were greatest i n the center of openings and l e a s t at the edge. A s i m i l a r edge e f f e c t was noted f o r the Grove ecotones. A l s o , accumulation and a b l a t i o n rates v a r i e d according to the d i r e c t i o n they faced, presumably r e f l e c t i n g the e f f e c t of wind and sun. Powell (1971) reviewed the e f f e c t s of c l e a r c u t t i n g on meteorological f a c t o r s . Climate of the ecotone i t s e l f i s a l s o important since i t o f f e r s a compromise to moose. In the simplest s i t u a t i o n , open areas have the most food but a l s o the harshest climate - deeper snow, co l d e r and more extreme temperatures, greatest wind run, and highest p o t e n t i a l f o r r a d i a t i o n a l heat l o s s e s . Conversely, f o r e s t s have l e a s t food but a moderated c l i m a t e . Ecotones provide an e c o l o g i -c a l compromise t h a t allows moose to optimize b e n e f i t s of food - and cover-producing h a b i t a t s . A d d i t i o n a l l y , ecotones have c l i m a t i c features d i s t i n g u i s h a b l e from both f o r e s t and open types. For example at Grove, temperatures were almost as warm as i n the open, but wind run was markedly lower and snow cover reduced s l i g h t l y . 10. DISCUSSION 10.1 Habitat R e l a t i o n s h i p s i n Moose Management Throughout the b o r e a l regions, the h i s t o r y of moose and man has been i n t e r t w i n e d . Many h i s t o r i c a l accounts describe the value of moose to man. In the U.S.S.R., at l e a s t as e a r l y as the 17th century, moose provided food f o r c o l o n i z i n g Caucasians. P u l l i a i n e n (19 74) mentioned the great importance of f l e s h and hides of moose f o r Europeans, and Markgren (19 74) noted that i n famines, moose were e x t i r p a t e d l o c a l l y . In North America, moose f i g u r e d i n the h i s t o r y of both indigeneous peoples and s e t t l e r s from the Old World. Peek (1971) noted that remains and pictographs of moose date back to at l e a s t 500 B.C. i n eastern America. K r e f t i n g (19 74) summarized b r i e f l y the r o l e of moose to man's welfa r e i n c e n t r a l North America. Dodds (1974) quoting Benson (unpubl. ms.) remarked t h a t moose provided meat and hides f o r the Indians, the white s e t t l e r s and the merchant v e s s e l s . P a r t i c u l a r l y i n southern Europe and eastern North America, i n d i s c r i m i n a t e hunting and h a b i t a t changes have reduced the h i s t o r i c a l d i s t r i b u t i o n and abundance of moose. I t i s a l s o true that man's a c t i v i t i e s have increased 379 380 moose populations. The r a p i d and remarkable increases of Fennoscandian moose have been a t t r i b u t e d to major changes i n land use p r a c t i c e s , e s p e c i a l l y logging and abandonment of a g r i c u l t u r a l lands (Ahlen 1975, Lykke and Cowan 1968, Markgren 1974). Range extensions to Labrador and Newfound-land r e s u l t e d from t r a n s p i a n t s ; and the success of moose i n the l a t t e r area i s due l a r g e l y to logging. Peek (19 71) concluded f o r Minnesota that pulpwood logging i n a d v e r t e n t l y created very good moose h a b i t a t . In B r i t i s h Columbia, the remarkable extension by moose i n t o the n o r t h - c e n t r a l area and subsequently southward and westward r e s u l t e d p r i m a r i l y from man's a c t i v i t i e s (Hatter 1950). Thus, s h i f t s i n a g r i c u l t u r e and logging p r a c t i c e s , and man-induced f i r e s occur as repeated themes i n build-ups and range extensions of moose throughout Fennoscandia and North America (Bedard et a l . 1974). Probably the most amazing aspect of these enhance-ments f o r moose was t h a t they were v i r t u a l l y a l l inadvertent. Almost without exception, major increases i n d i s t r i b u t i o n and abundance of moose were chance by-products of other land-use a c t i v i t i e s . H i s t o r i c a l l y , land was used f o r s i n g l e purposes such as lumbering, mining, farming or ranching. Most l i k e l y , the cause and e f f e c t r e l a t i o n s h i p , between vegetation disturbance and moose abundance was e i t h e r neglected or overlooked. . Ha b i t a t manipulation f o r the e x p l i c i t purpose of moose production i s a recent concept. 381 P r e s e n t l y , although the a b i l i t y and value of managing a c t i v e l y i s recognized by w i l d l i f e managers, the future welfare of moose i s s t i l l l a r g e l y determined by other land use a c t i v i t i e s . The present s i t u a t i o n f o r north-c e n t r a l B.C. was described i n the I n t r o d u c t i o n : r a p i d l y i n c r e a s i n g human populations occupy valuable h a b i t a t , encourage the development of u t i l i t y and t r a n s p o r t a t i o n c o r r i d o r s across migration routes, and place great demands upon r e c r e a t i o n a l resources; h y d r o e l e c t r i c developments destroy e s s e n t i a l w i n t e r h a b i t a t s , and s t a b i l i z e the down-stream r i v e r systems so reducing t h e i r value to moose; a g r i c u l t u r e continues to expand on f i n e - t e x t u r e d s o i l s that formerly produced long-term s u p p l i e s of w i n t e r browse; and f o r e s t r y seeks to c u r t a i l w i l d f i r e , probably the s i n g l e most important n a t u r a l f a c t o r i n the ecology of moose. H i s t o r i c a l l y , i t i s obvious t h a t man's land use a c t i v i t i e s were l a r g e l y f o r t u i t o u s l y b e n e f i c i a l f o r moose. The nature and e s p e c i a l l y the sc a l e of contemporary a c t i v i t i e s are s t i l l l a r g e l y f o r t u i t o u s but, except f o r f o r e s t r y , g e n e r a l l y d e t r i m e n t a l . While i t i s u n l i k e l y that moose 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 w i l l be exterminated by current land uses, i t appears h i g h l y l i k e l y t h a t moose could be reduced i n abundance and i n d i s t r i b u t i o n . Whether or not t h i s i s d e s i r a b l e , i s p r i m a r i l y a s o c i a l and economic matter. In many ways, the present-day d e c i s i o n s bear importantly upon the future of 382 moose and t h e i r value to people of the province. Yet c l e a r statements of management o b j e c t i v e s f o r moose appear l a c k i n g . In s i t u a t i o n s where d e c i s i o n s must be made i n the absence of c l e a r l y s t a t e d o b j e c t i v e s , the future of moose i s on an u n s a t i s f a c t o r y f o o t i n g . Despite the lack of o b j e c t i v e s , the future p u b l i c demand f o r moose can be assumed w i t h some confidence. A l s o , the development of i n t e g r a t e d resource management and the gradual acceptance of moose management i n the process, suggests t h a t moose w i l l continue to be an important f a c e t of the area's ecosystem. The extent and impact of resource development places e v e r - i n c r e a s i n g demands on a . f i n i t e land base. I t f o l l o w s t h a t more i n t e n s i v e management i s req u i r e d to face these future demands, issues and c o n f l i c t s . The requirement i s e s p e c i a l l y important f o r the n o r t h - c e n t r a l region since i t produces 40 - 50 percent of the p r o v i n c i a l moose harvest. As the t e r r a i n g e n e r a l l y poses few b a r r i e r s t o access and many resource development a c t i v i t i e s , problems between w i l d -l i f e (moose) and other resource sectors w i l l a r i s e f r e q u e n t l y . The need f o r land use. zoning and f u n c t i o n a l l y i n t e g r a t e d management i s e s s e n t i a l . . Given the area's importance as a moose producer and the i n e v i t a b l e problems r e s u l t i n g from increased development by other resource s e c t o r s , i t i s an important p r e r e q u i s i t e that moose management o b j e c t i v e s be s t a t e d c l e a r l y . I t i s 383 as important t h a t management be on a sound b i o l o g i c a l b a s i s . The scale of knowledge must be commensurate w i t h the scale of use. The b i o l o g i c a l b a s i s begins w i t h a conceptual model of f a c t o r s r e g u l a t i n g moose numbers; and ends w i t h an a b i l i t y to i n t e r p r e t and apply the model i n r e g i o n a l and sub-regional contexts. Apparently, only Houston (19 68) has attempted to develop a model showing probable mechanisms t h a t regulate moose numbers (Figure 10.1). His proposal was based on research i n t o the Shiras moose i n the Jackson Hole, Wyoming area, but i t i s s u f f i c i e n t l y general t o be a p p l i c a b l e elsewhere. In the r e s t of t h i s s e c t i o n , I discuss r e s u l t s of my study w i t h respect to Houston's model and suggest ways to improve h i s proposal. As my t h e s i s i s h a b i t a t o r i e n t e d , my remarks deal mostly w i t h the environmental r a t h e r than population mechanisms. Gross (undated) renamed these mechanisms h a b i t a t and population responses, r e s p e c t i v e l y . At the outset, the f o l l o w i n g quote from Gross (undated:5-7) provides an important.perspective on the model and how i t should be viewed: The components are arranged i n a multi-branched h i e r a r c h y . . . . F u n c t i o n a l information i s i n t e g r a t e d i n each component and i s passed on to the next higher compartment to be i n t e g r a t e d i n t o i n c r e a s i n g l y complex in f o r m a t i o n . In t h i s manner, the i n t e r n a l complexity of i n d i v i d u a l components increases nearer the apex of the pyramid. Con-v e r s e l y , the comparative complexity of l i n k s between components decreases nearer the apex of the pyramid. For example, the i n t e r n a l complexity . (near the apex) of Natality Rate i s greater than the I n t e r n a l complexity of a component (near the base) 383a Figure 10.1 Major f a c t o r s and how they i n t e r r e l a t e to a f f e c t moose population l e v e l s (modified from Houston 1968). AVAILABLE ENERGY AND NUTRIENTS SNOW FEATURES SPECIES COMPOSITION AND ABUNDANCE c CLIMATE PLANT SUCCESSION ^ORGANISMfP) TYPE 3 /ECOLOGICAL\" PLANT PROPERTIES, to oo 385 such as Quantity of Forage. However, the e f f e c t of Natality Rate on Population Size i s e a s i e r to determine than i s the. e f f e c t of Quantity of Forage on e i t h e r an adjacent component such as Energy Available or a d i s t a n t component such as Population Size. In general, the d i f f i c u l t y of determining cause and e f f e c t increases as more l i n k s (progres-s i n g toward the base) a r e . i n c l u d e d i n the f u n c t i o n a l process. Two I m p l i c a t i o n s f o r studying competition are evident i n the p e r s p e c t i v e of h i e r a r c h i c a l components. F i r s t , there seems to be l i t t l e chance of d e s c r i b i n g a s i g n i f i c a n t f u n c t i o n a l i n t e r a c t i o n (such as l e a s t squares regression w i t h confidence and c o r r e l a t i o n statements) which operate between a low-order-complexity component such as Quantity of Forage and the highest-order-complexity component such as Population Size by attempting to measure simultaneous changes i n the two components. The many components th a t are i n t e g r a t e d to produce a p a r t i c u l a r population s i z e probably: (1) change independently of each other, (2) change t h e i r r e l a t i v e e f f e c t on the population over time, and (3) change t h e i r r e l a t i v e e f f e c t on the population as population density changes. Thus, the e f f e c t on the population of any s p e c i f i c low-order-complexity component would probably be confounded by the e f f e c t s of other components. Such confounding would reduce the chances of showing a reasonable c o r r e l a t i o n between a s i n g l e low-order-complexity component and a high-order-complexity component. Second, i f the foregoing deductions are c o r r e c t , the best approach f o r d e s c r i b i n g the e f f e c t of low-order-complexity components on Population Size i s not to begin the a n a l y s i s by attempting to describe the e f f e c t of a component such as Quantity of Forage on Population Size. A more promising approach i s t o begin analyses by showing the e f f e c t of an adjacent high-order-complexity component ( f i r s t -order- i n . t e r a c t i on) such as Natality Rate, Mortality Rate, or Dispersal on Population Size. A f t e r f i r s t - o r d e r i n t e r a c t i o n s are described, the l o g i c a l p o i n t f o r a d d i t i o n a l work i s the next higher order (second-order i n t e r a c t i o n s ) of i n t e r a c t i o n s . In t h i s manner, i n c r e a s i n g l y complex i n t e r a c t i o n s are explained as more low-order-complexity components are i n t e g r a t e d i n t o the mechanism. The approach i s fundamentally c o n s i s t e n t w i t h that of H o l l i n g (1966) : 386 \"The j o i n t need f o r p r e c i s i o n and r e a l i s m l e d to the development of the experimental components a n a l y s i s so t h a t the explanation could be b u i l t i n many small steps, each step being taken only when the explanation posed i n previous steps had been experimentally v e r i f i e d . In t h i s way, the form of the explanation i s d i c t a t e d not as much by the need f o r mathematical neatness, but by the process i t s e l f . \" Although the e f f e c t on Population Size of a component such as Quantity of Forage w i l l - n o t be explained by d e s c r i b i n g the e f f e c t of Natality Rate on Population Size, the e f f e c t of Quantity of Forage on Population Size w i l l be i n c l u d e d to the extent t h a t i t a f f e c t s Natality Rate. For example., i f Quantity of Forage exerted the major c o n t r o l l i n g i n f l u e n c e on Natality Rate., then e x p l a i n i n g the. e f f e c t of Natality Rate on Population Size would e x p l a i n the e f f e c t of Quantity of Forage on Population Size. But i f Quantity of Forage had l i t t l e e f f e c t on Population Size, then e x p l a i n i n g the e f f e c t of Natality Rate on Population Size would provide l i t t l e i n f o r m a t i o n on how Quantity of Forage a f f e c t s Population Size. The above quotation should be q u a l i f i e d by rec o g n i z i n g that the degree of i n t e r n a l complexity of various components depend l a r g e l y upon an obse r v e r 1 s p o i n t of view. P l a n t e c o l o g i s t s could argue j u s t i f i a b l y t h a t p l a n t succession i s as complex as n a t a l i t y r a t e , and they could develop a s i m i l a r compartment model w i t h a su c c e s s i o n a l parameter at the apex. The study's o b j e c t i v e s and the researcher's p e r s p e c t i v e determine the perception of r e a l i t y . A v a i l a b l e energy i s i l l u s t r a t e d as the i n t e r f a c e between the population and h a b i t a t responses (Figure 10.1). This s i n g l e parameter i s somewhat s i m p l i s t i c and should be 387 expanded to i n c l u d e at l e a s t p r o t e i n . For other n u t r i e n t s , Robbins (19 73) suggested t r e a t i n g d e f i c i e n c i e s or t o x i c i t i e s on a g e o g r a p h i c a l l y l o c a l i z e d b a s i s . For example, f u n c t i o n -a l copper d e f i c i e n c i e s may occur i n some areas of north-c e n t r a l B.C. where i t s a v a i l a b i l i t y may be impaired by molybdenum.. Moose wit h overgrown hooves, a symptom of copper d e f i c i e n c y (Flynn e t al.. 1977) , have been reported to r e g i o n a l o f f i c i a l s of the F i s h and W i l d l i f e Branch. A l s o , the linkages between a v a i l a b l e energy and other population responses may be more e f f e c t i v e l y and r e a l i s t i c a l l y portrayed using the animal n u t r i t i o n approach of Robbins (1973) and Moen (1973). Thus the amount of a v a i l a b l e energy (Robbin's (1973) range supply) u t i l i z e d i s determined by the requirements of the animal. The h a b i t a t responses or environmental f a c t o r s can be subdivided i n t o those that are mainly long-term and those that are mainly short-term or annual. The c e n t r a l long-term f a c t o r i s p l a n t succession, since i t determines the q u a n t i t y of forage, the long-term forage a v a i l a b i l i t y ( h o r i z o n t a l and v e r t i c a l ) , and a l s o the q u a n t i t y and a v a i l a b i l i t y of cover. Succession a l s o appears to i n f l u e n c e forage q u a l i t y . As noted i n s e c t i o n 7, succession i s determined by f a c t o r s i n a d d i t i o n to c l i m a t e . In t h i s p a r t of the model, the major f a c t o r s i n f l u e n c i n g succession need (to be s t a t e d since they a f f e c t s u c c e s s i o n a l rates and patterns and since some are manageable. For the P r i n c e George area, substrate was shown 388 to be an important f a c t o r f o r management. C l i m a t i c regions are a l s o important, though f u r t h e r study would be u s e f u l on t h i s p o i n t . Types of disturbance i s another s i g n i f i c a n t f a c t o r since the h a b i t a t produced,by d i f f e r e n t disturbances vary i n t h e i r a t t r a c t i v e n e s s to moose. The disturbance f a c t o r was not st u d i e d i n d e t a i l i n t h i s t h e s i s , but should be examined i n a subsequent study. Another long-term f a c t o r i s d i v e r s i t y of the p l a n t community patterns (pattern d i v e r s i t y ) since i t a f f e c t s ways i n which moose use h a b i t a t s (forage a v a i l a b i l i t y ) and the a v a i l a b i l i t y of some p l a n t s f o r c o l o n i z a t i o n of cutover and burned lands. Attempts t o q u a n t i f y t h i s d i v e r s i t y i n terms meaningful to ungulates have been unsuccessful. I t s importance has been recognized at l e a s t since Leopold (19 33) s t a t e d h i s law of i n t e r s p e r s i o n . McGinnes (1969) discussed p a t t e r n w i t h respect to w h i t e t a i l deer and r e f e r r e d to the much quoted work of Reynolds (1966a and b). He a l s o noted the need to consider home range s i z e i n c o n s i d e r i n g s i z e and d i s t r i b u t i o n of cuts. Kelker (19 64) developed an index of edge but as Patton (1975) pointed out, perimeter-based i n d i c e s are not expressed i n a form that can be r e l a t e d to area. Patton (1975) then proposed a d i v e r s i t y index that r e q u i r e s t e s t i n g . In the present study, the apparent importance of h a b i t a t d i v e r s i t y to moose was demonstrated by the heavy use of p a r t i a l l y logged cutovers (Section 3). Other s t u d i e s discussed i n Section .3 suggest t h a t the 38.9 p r e f e r e n c e o f m o o s e f o r h a b i t a t s w i t h a c o m p l e x o f p l a n t c o m m u n i t i e s i s a g e n e r a l o n e . H o w e v e r , t h e f u n c t i o n a l r e l a t i o n s h i p b e t w e e n h a b i t a t d i v e r s i t y a n d s o m e i n d e x o f m o o s e u s e h a s n o t r e c e i v e d t h e a t t e n t i o n i t d e s e r v e s . Home r a n g e , c a r r y i n g c a p a c i t y a n d h a b i t a t s e l e c t i o n a r e t h r e e b a s i c i n g r e d i e n t s t o t h e d e v e l o p m e n t o f t h i s f u n c t i o n a l r e l a t i o n s h i p . Home r a n g e i s t h e a r e a i n w h i c h a n a n i m a l p e r f o r m s i t s n o r m a l a c t i v i t i e s . (A c o m p r e h e n s i v e b i b l i o g r a p h y o n h o m e r a n g e s a n d d i s p e r s a l w a s p r e p a r e d b y H a r e s t a d a n d B u n n e l l (1977)). H o w e v e r , t h i s i s n o t a n o p e r a t i o n a l d e f i n i t i o n . T h e r e v i e w s b y H a r e s t a d (1975) a n d S c h o e n e r (1968) s h o w e d s t r o n g c o r r e l a t i o n s b e t w e e n b o d y w e i g h t t o t h e 0.75 p o w e r a n d h o m e r a n g e s i z e . F o r m o o s e , t h e p r e d i c t e d s i z e i s 1609 h a ( H a r e s t a d a n d B u n n e l l 1977). H o w e v e r , t h e f e w d e t a i l e d s t u d i e s o n m o o s e s h o w t h a t h o m e r a n g e s v a r y i n s i z e a c c o r d i n g t o s e a s o n , a g e , s e x a n d g e o g r a p h i c a r e a . U n d o u b t e d l y , h o m e r a n g e a r e a s a l s o v a r y a n n u a l l y a n d w i t h h a b i t a t d i v e r s i t y a n d c o m p o s i t i o n , s o c i a l s t a t u s a n d m i g r a t o r y p a t t e r n ( L e R e s c h e 19 74). P e r h a p s t h e k e y p o i n t h e r e i s w h a t d e t e r m i n e s h o m e r a n g e s i z e . T h e a b o v e r e v i e w s s u g g e s t t h a t i t i s u l t i m a t e l y d e t e r m i n e d b y e n e r g e t i c s . C a r r y i n g c a p a c i t y i s a s e c o n d b a s i c c o n c e p t . I t h a s b e e n d e f i n e d v a r i o u s l y a n d i s i n s o m e w a y s a n u n u s e a b l e t e r m ( E d w a r d s a n d F o w l e 1955, G r o s s u n d a t e d ) . T h e t e r m i s d e r i v e d f r o m t h e l o g i s t i c m o d e l o f p o p u l a t i o n g r o w t h . T h e 390 rate of population increase i s a density-dependent f u n c t i o n that decreases as the population s i z e approaches \"K\", an upper numerical l i m i t of numbers or c a r r y i n g c a p a c i t y (Odum 19 71). Thus capacity i s defined i n terms of numbers of animals or density. However,,this d e f i n i t i o n of l i m i t needs to be q u a l i f i e d w i t h respect t o s p e c i f i e d c o n d i t i o n s , a p a r t i c u l a r area and a set time p e r i o d (e.g., Wilson 1971). Otherwise, confusion develops. Gross (undated:16-17) s t a t e d t h a t \"concepts of c a r r y i n g c a p a c i t y . . .may c o n t r i b u t e valuable i n s i g h t f o r understanding the general workings of ecosystems, but. . . ( t h e i r ) b a s i c inadequacy. . . i s t h e i r f a i l u r e to d i r e c t l y r e l a t e to any population v i t a l s t a t i s t i c that can be p r e c i s e l y measured.\" Hab i t a t s e l e c t i o n i s a t h i r d b a s i c concept i n answering the problem of h a b i t a t d i v e r s i t y and use. Habitat s e l e c t i o n i s the process of choosing or p r e f e r r i n g a h a b i t a t (\"a place to l i v e \" (Andrewartha and B i r c h 1954)) from a number of p o t e n t i a l l y a v a i l a b l e h a b i t a t s . I t has two aspects. F i r s t , an element of choice i s necessary to d i s t i n g u i s h s e l e c t i o n from u t i l i z a t i o n . Choice probably has a b e h a v i o r a l component th a t provides the mechanism f o r choice, and an e v o l u t i o n a r y component that confers s u r v i v a l value i n h a b i t a t s e l e c t i o n (Krebs 1972). Second, the f a c t o r s or s t i m u l i that provide the b a s i s of choice must be i d e n t i f i a b l e . The i d e n t i f i c a t i o n of r e l e v a n t f a c t o r s i s an e s p e c i a l l y important matter f o r w i l d l i f e managers since i t 391 can determine whether or not to attempt h a b i t a t manipulation and what to do. One subtle d i s t i n c t i o n should be made here: f a c t o r s may act as cues or t r i g g e r s to the animal that are i n themselves, not.otherwise f u n c t i o n a l l y s i g n i f i c a n t . Thus, i n a d d i t i o n to d e f i n i n g the proximate cues, t h e i r u l t i m a t e b e n e f i t should a l s o be known. Generally, t h i s d i s t i n c t i o n i s not made i n most w i l d l i f e s t u d i e s . Habitat s e l e c t i o n probably operates at d i f f e r e n t l e v e l s and the operative f a c t o r s determining choice may vary f o r each l e v e l . At l e a s t three l e v e l s were d i s t i n g u i s h a b l e f o r moose i n sub-boreal f o r e s t s , v i z . , 1) s e l e c t i o n of seasonal ranges, 2) s e l e c t i o n of h a b i t a t s w i t h i n a seasonal range, and 3) s e l e c t i o n of m i c r o s i t e s w i t h i n a h a b i t a t . For example, s e l e c t i o n of w i n t e r h a b i t a t i s probably made i n response to snow c o n d i t i o n s . The long-term b e n e f i t i s presumably th a t s u r v i v a l of migrants i s b e t t e r than that of non-migrants. Once on a win t e r range, moose s e l e c t amongst h a b i t a t s . Since the needs of moose are v a r i e d , the c r i t e r i a f o r choosing h a b i t a t s i s l i k e w i s e v a r i e d . However, of prime importance are the needs f o r food, space, s h e l t e r , and escape from predators. Within each h a b i t a t , a f u r t h e r s e l e c t i o n appears to occur. Thus i n a h a b i t a t chosen f o r feeding, moose may s e l e c t the wetter s i t e s where forage p r o t e i n l e v e l s are higher than forage on mesic and x e r i c s i t e s . I t a l s o appears that the \" c r i t i c a l n e s s \" of s e l e c t i o n 392 may vary, p a r t i c u l a r l y as the animal's c o n d i t i o n d e c l i n e s during w i n t e r . This i n c r e a s i n g d i s c r i m i n a t i o n probably acts mostly at the l e v e l of m i c r o s i t e s e l e c t i o n . Thus i n e a r l y w i n t e r , moose may s e l e c t a f o r e s t e d h a b i t a t f o r bedding. I t would not show s e l e c t i o n f o r m i c r o s i t e s . s i n c e snow co n d i t i o n s are not severe,, and since the animal i s l i k e l y i n good p h y s i c a l c o n d i t i o n . In l a t e w i n t e r , the same moose i n the same h a b i t a t may s e l e c t f o r m i c r o s i t e s to minimize energy demands since snow c o n d i t i o n s are adverse, and since the a f f e c t s of even low energy demands are p r o p o r t i o n a t e l y much greater. Human a c t i v i t i e s and disturbance was the l a s t long-term f a c t o r i d e n t i f i e d i n Houston's model (Figure 10.1). A c t u a l l y , human a c t i v i t i e s a f f e c t , most of the short- and long-term f a c t o r s . Since f o r e s t r y i s the dominant human a c t i v i t y a f f e c t i n g moose h a b i t a t , and since these a c t i v i t i e s are v a r i e d and have many e f f e c t s , d i s c u s s i o n i s r e f e r r e d to the next s e c t i o n (10.2). S i x short-term or annual f a c t o r s i n t e r a c t t o modify a v a i l a b l e energy, v i z . , g r a z i n g by other ungulates, annual snow c o n d i t i o n s , forage p l a n t c o n d i t i o n , q u a n t i t y of forage, q u a l i t y of forage used and annual forage a v a i l a b i l i t y (Figure 10.1, Houston 1968). A v a i l a b l e energy and n i t r o g e n depend upon the f i r s t order f a c t o r s of q u a l i t y and q u a n t i t y of a v a i l a b l e forage. Forage n u t r i e n t q u a l i t y v a r i e s according to s e v e r a l 393 f a c t o r s . At l e a s t f o r some n u t r i e n t s , composition i s p a r t l y determined by l e v e l s a t t a i n e d i n the previous growing season (see Section 8). Shoot: l e a f r a t i o s are al s o important since leaves and buds contain p r o p o r t i o n a t e l y much more ni t r o g e n than twigs (see al s o Section 8, Cowan e t a l . 19 70). In t u r n , these r a t i o s are a f f e c t e d by the l e v e l and time of use. Growing season parameters of temperature and p r e c i p i t a t i o n a l s o modify n u t r i e n t l e v e l s . Most of the f a c t o r s i n f l u e n c i n g forage n u t r i t i v e value are impervious to a l l but e x c e p t i o n a l management p r a c t i c e s . In a few cases, a l t e r i n g l e v e l s of forage use by domestic gr a z i n g may be a method of improving forage q u a l i t y but f o r much of the n o r t h - c e n t r a l region, ranges used by c a t t l e and moose are not managed j o i n t l y . S i m i l a r l y , forage q u a n t i t y v a r i e s according to s e v e r a l f a c t o r s . Grazing by other herbivores modifies q u a n t i t y both d i r e c t l y by removing vegetation and i n d i r e c t l y by i n f l u e n c i n g c o n d i t i o n of the range p l a n t s . For moose, snow i s probably the main f a c t o r determining forage a v a i l a b i l i t y i n winter. L i k e g r a z i n g , i t a c t s d i r e c t l y and i n d i r e c t l y . Since moose r a r e l y paw through snow, snow pack e f f e c t i v e l y define the lower v e r t i c a l l i m i t of a v a i l a b l e browse. As noted i n s e c t i o n 9, however, snow pack i s q u i t e v a r i a b l e even w i t h i n a small area. The nature of f o r e s t canopies, the j u x t a p o s i t i o n of open and f o r e s t e d patches, and the s i z e of openings a l l a f f e c t snow d e p o s i t i o n and 394 melt. The p o t e n t i a l s f o r manipulating f o r e s t stands to a l t e r snow behaviour are many (see Section 9). Snow c o n d i t i o n s play an important i n d i r e c t r o l e by i n f l u e n c i n g the d i s t r i b u t i o n and d e n s i t i e s of w i n t e r i n g moose, and the occupancy p e r i o d on winter ranges. I t i s al s o l i k e l y t h a t snow and probably other c l i m a t i c features modify forage a v a i l a b i l i t y by i n f l u e n c i n g use of open areas adjacent to f o r e s t s . However, i t i s important to f i r s t assess the r e l a t i v e impact of snow on moose before developing h a b i t a t management plans. For example, i n areas where mean s n o w f a l l does not exceed about 80 cm, snow i s l i k e l y not a major h a b i t a t management c o n s i d e r a t i o n . In high s n o w f a l l areas, such as Quebec where mean snow depths exceed 200 cm, snow i s c l e a r l y a c r i t i c a l f a c t o r and w i l l a f f e c t management p r e s c r i p t i o n s on such issues as s i z e and shape of c l e a r c u t s . 10.2 The E f f e c t s of Timber Management on Moose Habitat F o r e s t r y a c t i v i t i e s have diverse impacts on moose h a b i t a t . These a c t i v i t i e s range from s i t e - s p e c i f i c e f f e c t s such as slashburning, to general ones such as the d i s t r i b u -t i o n of cut blocks and the sequence and season of c u t t i n g ; a l l are re g u l a t e d by annual allowable cuts (A.A.C.). Thus f o r e s t r y p r a c t i c e s can a f f e c t i n d i v i d u a l or very l o c a l i z e d groups of moose as w e l l as herds or populations. E v a l u a t i n g such a comprehensive and wide-ranging land use i s u s u a l l y 395 done i n the context of a general model, the approach adopted f o r t h i s s e c t i o n ( f o l l o w i n g Bunnell and Eastman 19 76). S i m i l a r to many f o r e s t - d w e l l i n g w i l d l i f e s pecies, moose have f i v e b a s i c resource requirements, v i z . , energy, n u t r i e n t s , water, t r a n s i e n t p r o t e c t i o n from c l i m a t i c elements ( s h e l t e r ) , and escape cover and space. To t h i s l i s t could be added such s p e c i a l i z e d needs as r u t t i n g grounds, c a l v i n g areas, and mineral l i c k s , but the f i r s t f i v e encompass most of the primary requirements, p a r t i c u l a r -l y i n the c r i t i c a l w i n t e r season. These resources vary i n t h e i r h o r i z o n t a l and v e r t i c a l d i s t r i b u t i o n , and whether they are used as ingested or s t r u c t u r a l elements (Table 10.1). Table 10.1 General Features of B a s i c Resources Required by Moose Resource Resource A t t r i b u t e Amount D i s t r i b u t i o n ingested s t r u c t u r a l h o r i z o n t a l v e r t i c a l Energy N u t r i e n t s Water Temporary s h e l t e r (from wind and snow) Escape cover and space x x x x x X X X X X X X X X 396 F o r e s t r y p r a c t i c e s r e s t r u c t u r e and r e d i s t r i b u t e these resources i n diverse ways. By co n s i d e r i n g these many impacts i n the context of a general model, they are r e l a t e d to the b a s i c resources i n a way tha t allows r e s u l t s of the present study to be c l e a r l y r e l a t e d w i t h those of many others. This approach aids i n t e g r a t i o n of knowledge plus f a c i l i t i e s i d e n t i f i c a t i o n of research needs. Development of the general model was based on a sy n t h e s i s of w i l d l i f e and f o r e s t r y l i t e r a t u r e (Bunnell and Eastman 19 76). Impacts of f o r e s t management p r a c t i c e s are considered according to the f o l l o w i n g s i x t o p i c s : 1. F e l l i n g : s i l v i c u l t u r a l systems, logging systems, residues. 2. S i t e p r e p a r a t i o n : s l a s h burning, s c a r i f i c a t i o n . 3. Stand establishment: species choice, monocultures, h e r b i c i d e s , stand conversion, p l a n t i n g , seeding. 4. Stand tending: spacing,, t h i n n i n g , f e r t i l i z a t i o n . 5. Stand p r o t e c t i o n : f i r e suppression and e x c l u s i o n , p e s t i c i d e s . 6. General management c o n s i d e r a t i o n s : r o t a t i o n length, annual allowable cut, landscape p a t t e r n , road b u i l d i n g . A l l these p r a c t i c e s are employed to varying degrees i n the n o r t h - c e n t r a l region. Those not examined i n t h i s study are evaluated w i t h moose research conducted elsewhere. 397 10.2.1 F e l l i n g The f e l l i n g systems commonly c a l l e d c l e a r c u t t i n g arid s e l e c t i v e c u t t i n g represent p o i n t s on a continuum t h a t ranges from nothing cut to everything cut. For moose, the s i g n i f i c a n t p o i n t i s t h a t p a r t i a l l o gging r e d i s t r i b u t e s w i l d l i f e resources over a wider range of h o r i z o n t a l and v e r t i c a l dimensions than occurs e i t h e r i n succession a f t e r c l e a r c u t t i n g or i n mature f o r e s t s . While the absolute amounts of forage or cover are l e s s than that f o r e a r l y s u c c e s s i o n a l or mature f o r e s t stands, r e s p e c t i v e l y , w i n t e r i n g moose i n n o r t h - c e n t r a l B r i t i s h Columbia p r e f e r r e d s e l e c t i v e l y logged f o r e s t s to c l e a r c u t s (Section 3). A s i m i l a r preference was evident i n summer. In other areas, productive moose ranges have been c h a r a c t e r i z e d by t h e i r h a b i t a t d i v e r s i t y . Winter concentration areas i n Nova S c o t i a were t y p i c a l l y p a r t i a l cutovers ( T e l f e r 1967, P r e s c o t t 1968). In Newfoundland, 2 mid-winter d e n s i t i e s of at l e a s t 6 moose/km were estimated f o r cutover balsam f i r - white b i r c h {Abies balsamea - Betula papyrifera) stands (Bergerud and Manuel 1968). Although commercially c l e a r c u t , these logged stands resembled p a r t i a l cutovers since a l l white b i r c h and a l l c o n i f e r s l e s s than 9 cm diameter were l e f t uncut. In Alas k a , LeResche e t a l . (19 74) noted t h a t the 19 4 7 Kenai burn, one of the most 2 productive large areas of moose h a b i t a t (12 moose/km ), was c h a r a c t e r i z e d by great v e g e t a t i v e d i v e r s i t y and edge. A 398 2 t y p i c a l sample p l o t of 2.5 km contained an estimated 112 km of edge. An apparent c o n t r a s t i s Sweden, where, the r a p i d increase i n moose numbers was a t t r i b u t e d to the s h i f t from s e l e c t i v e c u t t i n g to c l e a r c u t t i n g (Markgren 1974). However, these cutovers were approximately 2 ha a decade ago and at present range from f i v e to 25 ha. Thus the r e s u l t a n t vegetation p a t t e r n tends towards a p a r t i a l cutover mosaic r a t h e r than the very l a r g e c l e a r c u t s of western North America. These apparent preferences f o r d i v e r s i t y can be i n t e r p r e t e d as a p o s i t i v e response to the f i n e - g r a i n e d mosaic of cover- and food-producing patches created by s e l e c t i v e logging as compared t o the r e l a t i v e l y homogeneous stands created by c l e a r c u t t i n g . P a r t i a l l o gging represents the p r a c t i c a l extreme of what Hatter (1950:134) considered to be i d e a l w i n t e r h a b i t a t f o r moose: A good d i s p e r s i o n of climax stands and new burns, . . ., probably c o n s t i t u t e s the very best winter h a b i t a t f o r moose. The preference f o r stands of. intermediate food and cover valued a l s o re-emphasizes the important d i s t i n c t i o n between absolute amounts and the animal's perception of a v a i l a b i l i t y . Other species w i t h c a t h o l i c d i e t s a l s o appear to p r e f e r p a r t i a l l y d i s t u r b e d stands. Freddy (1974), i n a general e c o l o g i c a l study of mountain c a r i b o u , suggested that extensive c l e a r c u t t i n g of Englemann spruce-subalpine f i r f o r e s t s would be d e t r i m e n t a l to t h i s species through removal of o l d e r lichen-producing stands. He recommended s e l e c t i v e and small patch logging as a l t e r n a t i v e s to extensive c l e a r c u t t i n g . Swanson (19 70) found l e v e l s of use by Roosevelt e l k (Cervus canadensis roosevelti) to be higher i n o l d e r , salvage-logged stands than i n unlogged timber (297 p e l l e t groups/ha) compared to 126/ha, but lower than i n c l e a r c u t s (330-699 groups/ha.). Any logging system creates s i t e disturbance. The nature and extent of disturbance caused by d i f f e r e n t logging methods i s v a r i a b l e , depending l a r g e l y upon the type used (Table 10.2), season of l o g g i n g , t e r r a i n , and c a l i b e r of the logging c o n t r a c t o r . Thus impacts are a l s o v a r i a b l e , but are mediated p r i m a r i l y through energy, n u t r i e n t s and water. Tr a c t o r logging i s the most p r e v a l e n t type of logging i n the study area. Deep disturbances commonly as s o c i a t e d w i t h t r a c t o r logging (Table 10.2), can lead to severe erosion and adversely a f f e c t moose through l o s s of h a b i t a t and lowered s i t e p r o d u c t i v i t y . Shallower disturbances t h a t s t i l l expose mineral s o i l probably have a b e n e f i c i a l e f f e c t through incre a s e d h o r i z o n t a l d i v e r s i t y of ingested resources and the i n i t i a t i o n of e a r l y s e r a i stages.that contain important food species. In the n o r t h - c e n t r a l r e g i o n , most logging occurs i n the w i n t e r . Logging i n t h i s season i s probably l e s s u s e f u l than summer logging i n producing moose h a b i t a t unless i t i s followed by slash-burning or s c a r i f i c a t i o n , due to the 400 Table 10.2 S o i l Disturbance and S l a s h 1 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 et a l . 1975) Type of l o g g i n g * * Type of Disturbance* M i n e r a l Amount Forest s o i l of f l o o r Shallow Deep Compacted T o t a l exposed* s l a s h * Horse (1) Jammer (1) Tract o r (1, 2,3,5) High-lead (2,3,4,5) S k y l i n e (4) H e l i c o p t e r (5) 10 13 13 25 11 12 15 50 32 12 15 37 20 31 36 17 30 15 14 23 45 *As percentage of area sampled. **Parentheses enclosure sources of data: (1) Garri s o n and Rummell (1951), (2) Wooldridge (1960, (3) Dyrness (1965), (4) Ruth (1967), (5) Bockheim et a l . (1975). Number of areas examined: f o r horse = 2; jammer = 3; t r a c t o r = 22; h i g h - l e a d = 19; s k y l i n e = 4; h e l i c o p t e r = 1. p r o t e c t i v e cover of snow, and frozen ground. My observations f o r winter-logged cutovers suggested a greater abundance of e a r l y s e r a i shrub species such as w i l l o w and paper b i r c h on the more d i s t u r b e d s k i d t r a i l s and haul roads than on the balance of the cutover. The preference by moose f o r s k i d t r a i l s and roads (Section 3) may be p a r t l y e x plained by t h i s d i f f e r e n t i a l disturbance. Even i n c l e a n l y 4Q1 logged cutovers where p o s t - l o g g i n g residues provided l i t t l e impediment to movement, s k i d roads and t r a i l s were p r e f e r r e d . Swanson (1970) found s i g n i f i c a n t l y higher e l k use on moderately or h e a v i l y d i s t u r b e d s i t e s than on l i g h t l y d i s t u r b e d s i t e s (P < .01). T e r r a i n a l s o modifies the type and extent of disturbance. On the gently r o l l i n g topography of the study, t e r r a i n . e f f e c t s were minimal. However, s t u d i e s i n mountainous t e r r a i n i n southern B r i t i s h Columbia i n d i c a t e considerable s i t e damage (Smith and Wass 1976). I b e l i e v e that the extent of s i t e disturbance was a l s o s t r o n g l y i n f l u e n c e d by the a b i l i t y and concern of the logging company (also mentioned by R. C l i f f o r d , per. comm.). As w e l l as s i t e disturbance, logging bequeaths l i v i n g and n o n - l i v i n g residues to the f o r e s t system. L i v i n g remains such as branches can provide an important, though temporary, food supply f o r w i n t e r i n g moose. C a r e f u l logging i n moose w i n t e r i n g areas t h e r e f o r e could provide short-term b e n e f i t . Dimock (1974) b e l i e v e d t h a t residues are g e n e r a l l y b e n e f i c i a l to w i l d l i f e . N o n - l i v i n g residues vary according to timber type, age of stand and logging method (Dimock (1974). Their impact on moose was not stud i e d but work on other ungulates suggests t h a t anything but l i g h t s l a s h would be de t r i m e n t a l to moose, both by reducing access to forage and reducing a v a i l a b l e space f o r p l a n t s . The impact of s l a s h i n decreasing amounts of understory vegetation has 402 been documented f o r c o a s t a l and i n t e r i o r f o r e s t s (see review by Garrison and Smith 19 74). In c o a s t a l Douglas f i r cutovers, s l a s h was the main f a c t o r reducing v e g e t a t i v e cover from approximately 70 percent to 10 percent (Dyrness 1965). In eastern Oregon and Washington, vegetative cover was reduced by 33 percent i n a s e l e c t i v e l y logged stand by a combination of s l a s h accumulation and s o i l disturbance (Garrison and Rummell 1951). In ponderosa pine stands of northern C a l i f o r n i a , Hormay (1940) recorded a 19 percent re d u c t i o n i n forage r e s u l t i n g from a s i m i l a r combination of s l a s h and s o i l disturbance. The d i s b e n e f i t s of residues i n n o r t h - c e n t r a l B.C. would be greater f o r spruce-subalpine f i r types than f o r lodgepole pine stands. The l a t t e r are u s u a l l y whole-tree logged w i t h t r e e s f e l l e d by shearing close to the base. The l a t t e r types u s u a l l y c o n tain a higher p r o p o r t i o n of unsound wood t h a t i s l e f t on the ground, and tr e e s are t y p i c a l l y limbed before s k i d d i n g to the landing. 10.2.2 S i t e P r e p a r a t i o n The commonest s i t e p r e p a r a t i o n techniques used i n the P r i n c e George region are slashburning, p i l e and burn and s c a r i f i c a t i o n . Slashburning i s g e n e r a l l y p r a c t i s e d i n spruce-subalpine f i r stands, though i t s prevalence i s d e c l i n i n g . L i m i t e d data comparing moose use on burn and unburned cutovers suggested that moose p r e f e r burned areas, 403 at l e a s t f o r the s i t u a t i o n s sampled (Section 3). A s i m i l a r preference f o r slashburned cutovers has been documented f o r other ungulates, v i z . , b l a c k t a i l deer (Odoooileus hemionus columbianus) (Gates 19 68) , and Roosevelt e l k {Cervus canadensis voosevelti) (Harper 1971, Swanson 1970) . P r e f e r e n t i a l use i s of l i m i t e d d u r a t i o n , u s u a l l y disappearing w i t h i n f i v e years a f t e r burning. Explanations f o r heavier use of slashburned areas are u s u a l l y given a posteriori (Eddleman and McLean 19 69) . Increased n u t r i t i v e value of p o s t - f i r e forages i s l i k e l y of l i m i t e d and short term importance (Einarsen 19 64, Taber 1973, Gates 1968). The most probable b e n e f i c i a l e f f e c t s are i n promoting growth of fire-dependent species such as f i r e -weed, and i n s l i g h t l y d e l a y i n g s u c c e s s i o n a l development. Removal of p o s t - l o g g i n g debris a l s o increases area a v a i l a b l e f o r p l a n t establishment and a c c e s s i b i l i t y of forage to moose. As w i t h n u t r i e n t l e v e l s , these assets of s l a s h -burning are of r e l a t i v e l y minor s i g n i f i c a n c e when compared to other h a b i t a t f a c t o r s such as cover and s i t e . M o d i f i c a t i o n s to broadcast burning, such as windrow and burn, p i l e and burn, are probably more b e n e f i c i a l to moose as they tned to increase h a b i t a t d i v e r s i t y . I found no q u a n t i f i c a t i o n of t h i s i n the l i t e r a t u r e f o r moose, but Swanson (19 70) noted that e l k use was s i g n i f i c a n t l y higher i n patchy, windrowed s l a s h than i n uniformly d i s t r i b u t e d s l a s h . Probably these observations were due to a more 404 favorable d i s t r i b u t i o n of food and cover resources i n the former s l a s h type. S c a r i f i c a t i o n i s becoming more common as a s i t e p r e p a r a t i o n technique i n n o r t h - c e n t r a l B r i t i s h Columbia. F u n c t i o n a l l y , t h i s p reparation method acts to a l t e r and most often r e t a r d s u c c e s s i o n a l rates and the p r o v i s i o n of food. I t i s t y p i c a l l y employed.in lodgepole pine stands, and has been advocated as a means of improving regeneration of white spruce (Eis 1967). Exposure of mineral s o i l s t i m u l a t e d growth of important browse species such as w i l l o w s and paper b i r c h and a l s o increased h a b i t a t d i v e r s i t y . S c a r i f i c a t i o n i s analogous to s k i d t r a i l s and minor logging roads. As noted p r e v i o u s l y , moose used these d i s t u r b e d access ways i n preference to adjacent u n t r a v e l l e d p a rts of cutovers. Thus moose would probably b e n e f i t from s c a r i f i c a t i o n . Studies by Young e t a l . (1967) showed that forage produced improved w i t h moderate mechanical disturbance. Conversely, S t e l f o x e t a l . (1976) found that forage production on s c a r i f i e d cut-overs lagged behind that on undisturbed cutovers f o r up to 17 years a f t e r logging. These v a r i a b l e conclusions demonstrate that the nature and degree of impact depends upon s e v e r i t y of the disturbance and the p r o d u c t i v i t y of the s i t e s . 405 10.2.3 Stand Establishment This a c t i v i t y has two f a c e t s : the e f f e c t of establishment on moose, and the e f f e c t of moose on stand establishment. With respect to the l a t t e r , moose r a r e l y were a problem i n e s t a b l i s h i n g the d e s i r e d commercial species of white spruce and lodgepole pine. As noted i n Sec t i o n 4, these species were r a r e l y taken and thus the impact of moose was n e g l i g i b l e . Douglas f i r i s planted o c c a s i o n a l l y but again, moose browsing on t h i s species was unimportant. The other commercially valuable c o n i f e r , subalpine f i r , was u t i l i z e d e x t e n s i v e l y by moose i n winter. In l o c a l i z e d areas, damage was severe enough t o cause death of subalpine f i r regeneration. However, t h i s t r e e species i s n e i t h e r p l a n t e d nor encouraged by extant s i l v i c u l t u r a l p r a c t i c e s . I t has ranked as a \"second c l a s s c i t i z e n \" i n the f o r e s t e r ' s t r e e community, and. t h e r e f o r e moose-induced damage was e s s e n t i a l l y i n c o n s e q u e n t i a l . However, t h i s s i t u a t i o n i s changing. Adverse impact of moose browsing appears more important i n eastern North America and Scandinavia than i n western America. Bergerud and Manuel (1968) reported that moose damage on balsam f i r i n Newfoundland was severe enough to prevent establishment of t h i s species. S p e c i a l hunting seasons were developed i n order to lower moose d e n s i t i e s and therefore reduce damage to acceptable l e v e l s . Damage to c o n i f e r s has a l s o been reported elsewhere i n eastern Canada 406 (Peterson 1955, P i m l o t t 1965) and the United States ( K r e f t i n g 1951, Murie 1934). Several Scandinavian workers examined the d e l e t e r i o u s e f f e c t s of moose browsing on Scotch pine {Pinus sylverstris) and spruce {Picea abies) (Kangas 1949, Lykke 1964, Westman 1958, a l l quoted by Markgren 1974). Much of the e a r l y concern on damage has been a l l a y e d by such studi e s as Lykke (19 64). Only a very low percentage of stems were k i l l e d and those that were browsed e x h i b i t e d compensatory growth once they reached beyond the browsing height of moose (Markgren 1974). The other f a c e t of stand establishment i s i t s a f f e c t on moose h a b i t a t . Three i n t e r - r e l a t e d p o i n t s are of main concern, v i z . , the rate of establishment, the tree species e s t a b l i s h e d , and stand conversion. I n c r e a s i n g the rate of establishment shortens the duration of the food-producing s u c c e s s i o n a l stage and therefore disadvantages moose. I t i s accomplished by a r t i f i c i a l r egeneration, immediately a f t e r the s i t e i s s u i t a b l y prepared, and, where necessary, c o n t r o l of competing vegetation by the most economical means ( h e r b i c i d e s ) . P r e s e n t l y , these manifestations of i n t e n s i v e f o r e s t r y are g e n e r a l l y absent i n the study area and t h e i r impact i s correspondingly minute. Regeneration i s s t i l l h e a v i l y dependent upon n a t u r a l seed sources. However, as f o r e s t r y becomes more i n t e n s e , these p r a c t i c e s w i l l become commoner and cause i n c r e a s i n g l y adverse impacts on moose h a b i t a t , p a r t i c u l a r l y i n the high s i t e , lowland f o r e s t s that 407 provide important year-round h a b i t a t f o r moose. The second m a n i f e s t a t i o n of i n t e n s i v e management i s the trend toward s i n g l e species stands or monocultures. In n o r t h - c e n t r a l B.C., the species of choice are lodgepole pine f o r t i l l and other coarse textured s o i l s , and white spruce f o r the heavy l a c u s t r i n e m a t e r i a l s . (Douglas f i r i s a l s o p l a n t e d but i n very l i m i t e d amounts). These species are e s s e n t i a l l y u n u t i l i z e d by moose, while the commercially u n a t t r a c t i v e species such as paper b i r c h , trembling aspen and subalpine f i r are taken. Thus developing commercial stands tend t o be homogeneous, even-aged populations of non-browse species. Concern f o r the long-range e f f e c t s of f o r e s t monocultures on moose have a l s o been expressed f o r Minnesota by Peek e t a l . (1976). An i n t e r e s t i n g , p a r t i a l l y c o n t r a d i c t o r y expectation was voiced by Ahlen (1975) who f e l t t hat the t r a n s i t i o n from spruce.to Scotch pine p l a n t a t i o n s would be b e n e f i c i a l to moose. The t h i r d f a c t o r of stand establishment i s stand conversion. P r e s e n t l y , t h i s a c t i v i t y i s p r i m a r i l y aimed at r e p l a c i n g o l d p a r t i a l l y logged stands ( \" s i l v i c u l t u r a l slums\") w i t h even-aged monocultures. Two adverse a f f e c t s are apparent: valuable p r e f e r r e d h a b i t a t s f o r moose are l o s t and areas s l a t e d f o r changeover can be q u i t e large (> 200 ha). Stand conversion through use of h e r b i c i d e s remains at the discussion/experimental stage. However, as wood production i n t e n s i f i e s , t h i s o p t i o n w i l l gain i n a t t r a c t i v e -408 ness. R e g i o n - s p e c i f i c i n v e s t i g a t i o n s on w i l d l i f e r e a c t i o n s should begin now i n s t e a d of a f t e r the f a c t . The e f f e c t s of stand establishment procedures w i l l l i k e l y have greatest adverse e f f e c t on l a c u s t r i n e h a b i t a t s . On these areas, seres are often c h a r a c t e r i z e d by a prolonged deciduous phase, p r i m a r i l y aspen, where white spruce, often the f i r s t major c o n i f e r , does not become dominant u n t i l approximately age 50 years. Thus, food production i s correspondingly prolonged. A r t i f i c i a l p l a n t i n g w i l l v i r t u a l l y e l i m i n a t e t h i s deciduous phase i n h a b i t a t s t h a t t y p i c a l l y are s i t u a t e d i n lands w i t h high c a p a b i l i t i e s f o r moose. I t should a l s o be noted that these h a b i t a t s are a l s o o f t e n subjected t o land c l e a r i n g f o r a g r i c u l t u r e . 10.2.4 Stand Tending The l i m i t e d amount of stand.tending p r a c t i c e d i n B r i t i s h Columbia i s r e s t r i c t e d almost e x c l u s i v e l y t o second growth f o r e s t s of the south c o a s t a l region. Thus stand tending i s not p r a c t i c e d i n moose h a b i t a t i n B.C. S i m i l a r l y , most North American moose, range i n commercial f o r e s t s has not yet experienced t h i s f o r e s t r y p r a c t i c e . However, some inferences can be derived from the l i t e r a t u r e on the two dominant tending p r a c t i c e s : t h i n n i n g or spacing and f e r t i l i z a t i o n . The f o l l o w i n g paragraphs of t h i s s e c t i o n were taken from Bunnell and Eastman (1976). Since thinnings reduce ba s a l area and canopy cover 409 of the overstory, the understory vegetation should respond w i t h increased biomass, p r o d u c t i v i t y and perhaps n u t r i e n t l e v e l s i n some species. However, the r e l a t i o n s h i p between biomass of the understory vegetation and crown closure f o l l o w s a negative e x p o n e n t i a l form (Jameson 1967), and a r e l a t i v e l y l a r g e reduction i n crown c l o s u r e i s necessary to obtain a s i g n i f i c a n t response i n understory vegetation (see F f o l l i o t t and Clay (1972) f o r an annotated b i b l i o g r a p h y on t h i s t o p i c ) . Given the v a r y i n g r e l a t i o n s h i p between crown closure and b a s a l area we expect the reduction necessary to s t i m u l a t e s i g n i f i c a n t understory production to d i f f e r markedly between f o r e s t types. L i t t l e i nformation i s a v a i l a b l e from f o r e s t types i n B r i t i s h Columbia. Dodd (1969) documented the general c u r v i l i n e a r r e l a t i o n s h i p between understory vegetation and tree overstory i n p a r k - l i k e , r e l i c t stands of i n t e r i o r Douglas f i r . Quenet (1973) and Kemper (1971) documented understory changes r e l a t e d to t r e e crown cl o s u r e i n s e r a i and r e l i c t Douglas f i r and ponderosa pine stands i n south-eastern B r i t i s h Columbia. Other s t u d i e s outside the province have documented the overstory:understory r e l a t i o n -ship and r e l a t e d i t t o w i l d l i f e use. In Arizona ponderosa pine, C l a r y and Lawson (19 71) documented an inverse r e l a t i o n s h i p between e l k p e l l e t groups and b a s a l area. In p o l e - s i z e d stands of Douglas f i r i n northern Idaho, Marsh (1954, i n Hungerford 1969) found t h a t b a s a l area had to be 410 reduced by 25-30 percent before a s i g n i f i c a n t response occurred i n the understory forage f o r white t a i l e d deer and r u f f e d grouse. Hungerford (1969) s t a t e d that these reductions were about 10 percent heavier than normal t h i n n i n g i n t e n s i t i e s f o r that area, but d i d not provide pre-t h i n n i n g b a s a l areas. In western white pine {Pinus monticola) stands of the same area, j u d i c i o u s t h i n n i n g can improve w i l d l i f e forage values f o r 30 to 40 years (Hungerford 1955). Thus, the p e r i o d of high browse biomass a f t e r . c l e a r c u t t i n g could be extended from the normal 10 to 15 years p o s t - l o g g i n g , up to 40 or 50 years i f t h i n n i n g took place 20 to 25 years a f t e r logging (Hungerford 1969). . Dealey (1975) reported s i m i l a r observations from a 4 7-year old.lodgepole pine f o r e s t i n c e n t r a l Oregon. Thinning resulted, i n a l o s s i n p r o d u c t i v i t y f o r the f i r s t two years due to shock of overstory removal. By the t h i r d year understory vegetative cover had exceeded p r e - t h i n n i n g l e v e l s at a l l spacings, and a f t e r nine years the stands were producing, from 300 to 1,000 percent more understory vegetative than before t h i n n i n g , w i t h the 4 m and 5.8 m pine spacings being most productive. Dealey (1975) d i d not report w i l d l i f e response to t h i n n i n g s , and the work of Pearson (1968) and C l a r y and. Lawson (1971) are e q u i v o c a l . For the mixed coniferous-deciduous, b o r e a l f o r e s t s of north-eastern North America, T e l f e r . (1974) found that r e s i d u a l 2 basal area had to be reduced to 17.2 m /ha before browse 411 production was s u b s t a n t i a l l y increased. Thinning has been observed to have det r i m e n t a l e f f e c t s on ungulates. D e l l and Ward (1969) noted t h a t s l a s h from pre-commercial th i n n i n g s of suppressed ponderosa pine i n eastern Oregon and Washington can reach depths of > 1 m, thus i m p a i r i n g access to large ungulates and reducing under-s t o r y growth. Working wi t h the same tree species i n c e n t r a l Oregon, Dealey (1975) noted that untreated s l a s h from 5.8 m spacings r e s u l t e d i n a c e s s a t i o n of deer use, regardless of forage production. For f o r e s t f e r t i l i z a t i o n the l i t e r a t u r e r e l a t i n g to w i l d l i f e s u f f e r s from an analogous \" c o n f l i c t of i n t e r e s t \" that permeates the l i t e r a t u r e on stand establishment. The reason i s the same: w i l d l i f e often, feed on commercial tr e e species. The f e r t i l i z e r treatment.which enhances t r e e growth ( i d e a l from the f o r e s t e r manager's perspective) often s t i m u l a t e s browsing (encouraging from the w i l d l i f e manager's p e r s p e c t i v e , but d i s q u i e t i n g to the f o r e s t e r ) . Behrend (1973) concluded t h a t these opposing p e r s p e c t i v e s are the main reason f o r the lack of a comprehensive approach r e l a t i n g f o r e s t f e r t i l i z a t i o n to w i l d l i f e management. In h i s short review, Behrend (1973:108) noted: There i s an abundance of info r m a t i o n about the e f f e c t s of f e r t i l i z a t i o n on f o r e s t p l a n t s , and there i s some information about the e f f e c t s of f e r t i l i z a t i o n on the n u t r i t i o n a l q u a l i t y of w i l d -l i f e food. Some information i s a l s o a v a i l a b l e about the r e l a t i v e a t t r a c t i v e n e s s of f e r t i l i z e d and u n f e r t i l i z e d p l a n t s t o herbivores. Thus i t might be concluded that management systems using 412 t h i s information are c u r r e n t l y employed i n f o r e s t resource management. However, t h i s i s not tr u e : l i t t l e more than crude s p e c u l a t i o n has a c t u a l l y been accomplished about the r e l a t i o n s h i p of f o r e s t f e r t i l i z a t i o n to w i l d l i f e management. While s t u d i e s i n mixed f o r e s t types of eastern North America have suggested that f e r t i l i z i n g could increase c a r r y i n g c a p a c i t y f o r deer i n forested.areas (Wood and Lindzey 1967; Ward and Bowersox 1970), con d i t i o n s i n the simpler c o n i f e r f o r e s t s of the west suggest the c o n f l i c t between w i l d l i f e and f o r e s t r y o b j e c t i v e s w i l l continue. 10.2.5- Stand P r o t e c t i o n In B r i t i s h Columbia, as over much of North America, a major o b j e c t i v e of f o r e s t management has been the c o n t r o l and p a r t i a l e l i m i n a t i o n of w i l d f i r e . U n t i l very r e c e n t l y t h i s o b j e c t i v e was pursued i n d i s c r i m i n a t e l y throughout the province. Arguments f o r f i r e c o n t r o l have been economic i n nature and only r e c e n t l y have the e c o l o g i c a l r a t i o n a l e and longer term economics been considered. As Heinselman (19 71) noted, the primeval c o n i f e r f o r e s t s of North America, w i t h t h e i r a s s o c i a t e d deciduous components, were l a r g e l y f i r e -dependent ecosystems. F i r e was an important f a c t o r i n e s t a b l i s h i n g succession, species.composition and age s t r u c t u r e of these f o r e s t s . The almost u n i v e r s a l a p p l i c a -t i o n of f i r e e x c l u s i o n has i n i t i a t e d succession which i s \"unnatural\" and o f t e n undesirable, from the perspect i v e of both timber and w i l d l i f e management. Some f o r e s t e r s are 413 becoming concerned that c e r t a i n stands are growing i n c r e a s i n g l y \" f i r e - p r o n e \" through accumulation of l i t t e r and n a t u r a l d e b r i s , and th a t chances f o r holocausts are i n c r e a s -i n g . W i l d l i f e managers are concerned that e a r l y s e r a i stages are being reduced i n extent. I f an i n t e g r a t e d approach to both w i l d l i f e and f o r e s t r y resources i s to be implemented, some zoning of f i r e suppression e f f o r t i s e s s e n t i a l . Where w i l d l i f e adapted to e a r l y s u c c e s s i o n a l stages are to be encouraged and f o r e s t values w i l l not be severely damaged, f i r e suppression may not be necessary or advantageous; where important f o r e s t values or climax-adapted w i l d l i f e are present, f i r e e x c l u s i o n i s an appropriate p o l i c y . Such zoning i s now present i n Montana ( A l d r i c h and Mutch 19 72), Yellowstone N a t i o n a l Park (Houston 1973), Alaska ( P r a s i l 1971) and elsewhere. In B r i t i s h Columbia costs of f i g h t i n g f i r e s i n low c a p a b i l i t y lands f a r removed from access are being evaluated, and attempts to zone f o r e s t lands according to the type and extent of f i r e suppression are underway. These e f f o r t s o f f e r opportunity f o r more e f f e c t i v e f o r e s t and w i l d l i f e management. A second major o b j e c t i v e of North American f o r e s t r y has been c o n t r o l of f o r e s t i n s e c t s and diseases. In the Pri n c e George d i s t r i c t , considerable concern e x i s t s regarding the spruce budworm. 414 10.2.6 General Management Considerations The establishment of the allowable annual cut (AAC) and r o t a t i o n age n a t u r a l l y i n f l u e n c e f o r e s t management p r a c t i c e s discussed above. S i m i l a r l y , they are i n t e r r e l a t e d and generate patterns of cut and uncut f o r e s t s w i t h extensive road networks. Except f o r one recent example i n the B e l l a Coola region, annual allowable cuts (AAC's) c a l c u l a t e d by the B r i t i s h Columbia Forest Service have been based s o l e l y upon timber production o b j e c t i v e s . These u n i l a t e r a l terms of reference e f f e c t i v e l y reduce options f o r other resource managers, p a r t i c u l a r l y when the AAC i s f u l l y committed to the f o r e s t i n d u s t r y and o p p o r t u n i t i e s f o r r e - a l l o c a t i n g , reducing or r e - d i s t r i b u t i n g the cut are l i m i t e d . As f o r e s t management becomes more i n t e n s i v e , r o t a t i o n lengths w i l l become sh o r t e r ; s i t e c o n t r o l and prompt regeneration w i l l become more important o b j e c t i v e s . High-y i e l d f o r e s t r y w i l l \"streamline\" e c o l o g i c a l succession and shorten the e a r l i e r s u c c e s s i o n a l stages. For moose, e l i m i n a t i o n of o l d e r stands w i l l probably prove det r i m e n t a l i n areas of high snow, but b e n e f i c i a l i n areas of low sn o w f a l l providing a s u i t a b l e p a t t e r n of cutover and f o r e s t e d land e x i s t s . Lawrence (19 69) suggested t h a t shortened r o t a t i o n length and i t s c o r o l l a r y , increased rate of cut, would probably b e n e f i t b l a c k - t a i l e d deer. However, the 415 shortened r o t a t i o n length almost c e r t a i n l y w i l l be accompanied by a need f o r e a r l y s i t e c o n t r o l and h e r b i c i d a l reduction of browse species commonly ass o c i a t e d w i t h e a r l y s u c c e s s i o n a l stages. Also Lawrence (1969) was d e s c r i b i n g c o n d i t i o n s i n an area of r e l a t i v e l y low s n o w f a l l . In B r i t i s h Columbia, b l a c k - t a i l e d deer commonly i n h a b i t areas of high s n o w f a l l where older f o r e s t s appear important i n p r o v i d i n g both food and s h e l t e r (Jones 1974, 1975). The d i s t r i b u t i o n of cuts i s as important as, and r e l a t e d t o , the rate of c u t t i n g . Landscape p a t t e r n i n v o l v e s not only the d i s t r i b u t i o n of c u t s , but the s i z e and shape of cut areas. The question \"How b i g should a cut block be?\" i s unanswerable unless the w i l d l i f e management o b j e c t i v e s are c l e a r l y s t a t e d , f o r example,.how many of what species. The d i s t r i b u t i o n of logging profoundly i n f l u e n c e s h o r i z o n t a l d i s t r i b u t i o n of w i l d l i f e resources, so t h a t home ranges of the species i n question must be considered when s p e c i f y i n g s i z e , shape and d i s t r i b u t i o n of cutovers. T e l f e r (1974), i n r e c o g n i z i n g t h i s p r i n c i p l e , suggested that f o r moose, logged patches could be up to 100 ha. Grazing i s another f a c t o r a f f e c t i n g n u t r i e n t l e v e l s . I t s main e f f e c t appears to be i n preventing e l a b o r a t i o n of o l d e r l e s s n u t r i t i o u s t i s s u e , i n modifying l e a f : shoot r a t i o s arid i n prolonging growth. Thus gra z i n g e f f e c t s are probably b e t t e r i n t e r p r e t e d and p r e d i c t e d by understanding w i t h i n p l a n t v a r i a t i o n s i n n u t r i e n t s . However, Oldenmeyer (1974) 416 found e s s e n t i a l l y s i m i l a r . d i g e s t i b i l i t i e s f o r w i nter samples of h e a v i l y used and normally used paper b i r c h . 10.3 Overview and Recommendations The management of moose i s at a crossroads i n north-c e n t r a l B r i t i s h Columbia. In the past, the net e f f e c t of man's a c t i v i t i e s on moose has been b e n e f i c i a l , although remarkably f o r t u i t o u s . The h i s t o r i c a l p a tterns of development as summarized i n the I n t r o d u c t i o n , revealed that human-caused f i r e , l o g g i n g , and a g r i c u l t u r a l l a n d - c l e a r i n g created s u i t a b l e h a b i t a t f o r moose; and the c i r c u m s t a n t i a l evidence i s t h a t these anthropogenic changes c o n t r i b u t e d s u b s t a n t i a l l y to the r a p i d southern and westward expansion of moose throughout the province.. To be sure, not a l l a c t i v i t i e s were b e n e f i c i a l , but the o v e r a l l assessment must be that they enhanced h a b i t a t f o r moose. That these same a c t i v i t i e s can have d i s b e n e f i t s f o r moose i s a recent r e a l i z a t i o n . Human-caused f i r e s are much reduced i n number, c l o s e l y c o n t r o l l e d i n extent, and c a r e f u l l y planned i n l o c a t i o n . This c o n t r a s t s sharply w i t h the h i s t o r i c a l p a t t e r n . Moreover, the i n t e g r a l r o l e of n a t u r a l f i r e i n these sub-boreal f o r e s t s i s d i m i n i s h i n g i n c r e a s i n g l y . The technology f o r f i r e d e t e c t i o n , c o n t a i n -ment and suppression improves apace. These trends have reduced the rate of production of s e r a i ranges. The impact of timber h a r v e s t i n g on moose has 417 c h a n g e d . D u e t o a c o m b i n a t i o n , o f p a r t i a l c u t t i n g m e t h o d s , s l o w r a t e o f h a r v e s t ( l a b o r - i n t e n s i v e ) , a n d t h e e x p l o i t a t i o n o f w e l l s t o c k e d s t a n d s p r i m a r i l y a t l o w e l e v a t i o n s , e a r l i e r t i m b e r h a r v e s t i n g c r e a t e d l i m i t e d , t h o u g h e x c e l l e n t w i n t e r h a b i t a t f o r m o o s e . D u e t o a c o m b i n a t i o n o f c l e a r c u t t i n g , a n i n c r e a s e d r a t e o f h a r v e s t ( h i g h l y m e c h a n i z e d ) , a n d t h e e x p l o i t a t i o n o f s t a n d s o v e r a w i d e e l e v a t i o n a l r a n g e , p r e s e n t - d a y t i m b e r h a r v e s t i n g i s c r e a t i n g e x t e n s i v e a r e a s o f h a b i t a t i n b o t h s u m m e r a n d w i n t e r r a n g e s . T h e i m p a c t o f a g r i c u l t u r a l l a n d c l e a r i n g h a s a l s o m o d i f i e d i n w a y s t h a t p a r a l l e l f o r e s t r y . E a r l y l a n d c l e a r -i n g w a s s m a l l i n a r e a a n d r e s t r i c t e d t o t h e b e t t e r , l o w e l e v a t i o n s o i l s . T h e h i g h r a t e o f a b a n d o n m e n t r e s u l t e d i n s m a l l p a t c h e s o f s e r a i v e g e t a t i o n o n p r o d u c t i v e l o w l a n d m o o s e w i n t e r h a b i t a t . P r e s e n t - d a y l a n d c l e a r i n g i s l a r g e i n e x t e n t t h o u g h s t i l l m a i n l y o n l o w l a n d s i t e s . T h e m a j o r d i f f e r e n c e i n a d d i t i o n t o i n c r e a s e d c l e a r i n g i s t h a t t h e a b a n d o n m e n t i s l e s s f r e q u e n t . T h u s h i g h c a p a b i l i t y m o o s e h a b i t a t i s m a i n t a i n e d i n f o r a g e c r o p s - v e g e t a t i o n t h a t i s e s s e n t i a l l y u s e l e s s f o r w i n t e r i n g m o o s e i f i t w a s a v a i l a b l e . T h e g r o w i n g a d v e r s e , o r a t l e a s t e q u i v o c a l , e f f e c t s o f t h e s e l a n d u s e p r a c t i c e s h a s b e e n a c c o m p a n i e d b y a g r o w i n g h u m a n p o p u l a t i o n , p r o l i f e r a t i n g a c c e s s , a n d a n i n c r e a s i n g d e m a n d f o r m o o s e . . T h e c u r r e n t e f f o r t s o f man o n m o o s e i n s u b - b o r e a l f o r e s t s a r e n o l o n g e r f o r t u i t o u s l y b e n e f i c i a l . 418 C l e a r l y , h i s t o r i c a l trends do not n e c e s s a r i l y p r e d i c t the f u t u r e . Nevertheless, these trends do i n f l u e n c e the choice of management options, and do provide some b a s i s f o r a ssessing patterns t h a t may occur. These trends i n d i -cate that i n n o r t h - c e n t r a l B r i t i s h Columbia the human population w i l l continue.to increase r a p i d l y . Both primary and secondary i n d u s t r i e s w i l l grow and resource demands and commitments w i l l expand. These developments must l e a d t o more c a r e f u l conservation of the region's n a t u r a l resources. As the demand f o r resources approach the ecosystem's a b i l i t y to meet them, options are fewer and accidents or mistakes l e s s e a s i l y absorbed. The f r o n t i e r environment w i l l be replaced by the managed environment. In t h i s context of a managed environment, the f o l l o w i n g question can be r a i s e d , \"What place do moose have i n i t ? \" The answer must be i n part a p h i l o s o p h i c a l one that supports the view that moose and other animals have a l e g i t i m a t e r o l e i n the scheme, of thi n g s . The~ answer must al s o i n p a r t r e l a t e to the demand f o r moose. Once i t i s accepted that moose do have a p l a c e , the next c o n s i d e r a t i o n i s how many moose are enough. This i s a d i f f i c u l t issue to decide since methods f o r assessing demand and f o r paying the costs of meeting t h i s demand are not s a t i s f a c t o r i l y solved. Notwithstanding these problems.with a non-market resource such as moose, some assessment of demand i s necessary i f management o b j e c t i v e s are to be formulated. 419 As t h i s study showed and many others show, moose are adaptable animals. They endure adverse w i n t e r s , s u b s i s t on a widespread.though comparatively i n d i g e s t i b l e resource, u t i l i z e v i r t u a l l y a l l stages of f o r e s t succession, and e x p l o i t h a b i t a t s ranging from tidewater wetlands t o a l p i n e meadows. As K e l s a l l and T e l f e r (1974) remarked, only extended periods of deep (> 70 cm) snow and high temperature appear to l i m i t moose d i s t r i b u t i o n i n North America. The a d a p t a b i l i t y of moose can be e x p l o i t e d through adequate planning and management of the sub-boreal landscape i f o b j e c t i v e s have been defined. Since timber h a r v e s t i n g i s the dominant land use, i n t e g r a t i o n of the two resources of f o r e s t r y and w i l d l i f e i s probably the s i n g l e most important task f a c i n g contemporary resource managers i n the region. . Some attempts at planning have begun (see below) but i t i s c r i t i c a l to place planning i n a pe r s p e c t i v e that considers b i o p h y s i c a l parameters (the c a p a b i l i t y f o r p r o d u c t i o n ) , s o c i a l and economics f a c t o r s (the demand and costs of pr o d u c t i o n ) , and the su c c e s s i o n a l a t t r i b u t e s of time and p a t t e r n . For n o r t h - c e n t r a l B r i t i s h Columbia, the f o l l o w i n g b i o p h y s i c a l a t t r i b u t e s bear c o n s i d e r a t i o n . F i r s t , the t e r r a i n i s uniform when compared to much of the province. R e l a t i v e l y few p h y s i c a l b a r r i e r s modify the patterns of resource development. The ease of access and the r o l l i n g t e r r a i n encourage continued development of i n t e n s i v e , 420 mechanized f o r e s t r y over large areas. Second, i t i s a region of gradual environmental gradients. Due to the r e l a t i v e l y slow rate of change across the landscape, these gradients may tend to be overlooked. Such oversig h t s could be b i o l o g i c a l l y unsound and economically i n e f f i c i e n t when i t comes to formulating management g u i d e l i n e s f o r moose and f o r e s t r y . For example, adopting, a c e r t a i n s i z e of c l e a r c u t across a snow gradient could l e a d t o cutovers too lar g e t o enhance moose production i n heavy snow areas, and not large enough i n l i g h t snow areas (but more c o s t l y ) . T h i r d , the region has a bor e a l c l i m a t e . The c l i m a t i c regime has important i m p l i c a t i o n s f o r the. n a t u r a l resources. As noted e a r l i e r , climate places severe r e s t r i c t i o n s on a g r i c u l t u r a l development. For f o r e s t r y , i t determines the p a t t e r n , s e a s o n a l i t y and l o c a t i o n of cut b l o c k s . For moose, i t i m p l i e s the need and importance of s u i t a b l e w i n t e r h a b i t a t as w e l l as i n f l u e n c i n g over-winter s u r v i v a l , forage production and migration r a t e s , timing and p a t t e r n s . Bearing i n mind the above p e r s p e c t i v e , the need f o r planning i n paramount i f a d e s i r e d resource mix i s t o develop i n the r i g h t places at the r i g h t times and i n the r i g h t amounts. A t t a i n i n g the \" r i g h t \" combinations i s a c h a l l e n g i n g , d i f f i c u l t task but i t i s c l e a r t h a t any planning f o r moose must be done i n conjunction w i t h f o r e s t r y . I t i s a l s o c l e a r t h a t planning w i l l have t o address landscape management, that i s , the mosaic and 421 p a t t e r n of vegetation r a t h e r than the more narrow c o n s i d e r a t i o n of p l a n t succession alone. The need f o r f o r e s t land planning has been recognized (Pearse 1976). Four l e v e l s of f o r e s t planning have been proposed (see Figure 19-1 i n Pearse 1976:262), but i t i s the o p e r a t i o n a l l e v e l that i s most p r e s s i n g . Two planning systems are p r e s e n t l y being a p p l i e d - F o l i o Planning and Coordinated Resource Planning. These systems meet some needs but f a i l to address others. For example, the main o b j e c t i v e s i n f o r e s t r y such as the annual allowable cut, are determined at the l e v e l of P u b l i c Sustained Y i e l d Units and Tree Farm Licences while management o b j e c t i v e s f o r moose and other vertebrates are determined at the l e v e l of sub-units or Timber Sale Harvesting Licences. While t h i s procedure has d e f i n i t e b e n e f i t s f o r moose (Eastman 1973), i t al s o denies p o t e n t i a l options f o r w i l d l i f e ( J . Dick, pers. comm.). Moreover, sub-unit plans often cover land areas that are too small to encompass the. f u l l annual range occupied by moose. Thus, i n t e g r a t e d management on p r i m a r i l y a moose summer range may be .wasted i f equal a t t e n t i o n i s not devoted to the management of c r i t i c a l w i n t e r ranges (Eastman 19 7 3). As these winter h a b i t a t s a l s o overlap p o t e n t i a l a g r i c u l t u r e land, the need a l s o a r i s e s to i n v o l v e other resource i n t e r e s t s . Coordinated Resource Planning attempts to draw a l l a f f e c t e d users i n t o preparing plans. However, i t a l s o 422 s u f f e r s from the problem of land u n i t s i z e . Coordinated Resource Planning has a u s e f u l way of i n t e g r a t e d d e c i s i o n -making but often r e l i e s upon inadequate inventory: F o l i o Planning has a u s e f u l way of i n t e g r a t e d inventory c o l l e c t i o n but l a c k s a corresponding way of decision-making (J. Dick, pers. comm.). What i s needed i s a s u c c e s s f u l combination of the best parts of each. Then t h i s \"hybrid\" should be placed i n context w i t h i n a management u n i t ( i . e . , the l e v e l of P u b l i c Sustained Y i e l d Units) and other l e v e l s of planning. U n t i l t h i s happens, i n t e g r a t e d management of moose and f o r e s t r y i s incomplete. Information about the planned resources i s an e s s e n t i a l p a r t of planning. This t h e s i s i s aimed at p r o v i d i n g information on h a b i t a t - r e l a t e d matters needed f o r planning. Some of the main p o i n t s t h a t r e l a t e to t h i s are as f o l l o w s (These assume that moose production i s a main goal) : 1. Moose appear to use h a b i t a t s d i f f e r e n t l y according to snow regimes. Therefore, management g u i d e l i n e s should r e f l e c t these d i f f e r e n c e s , e.g., i n s i z e of cutovers. 2. Moose appear t o p r e f e r s e l e c t i v e l y logged h a b i t a t s to other types of cutovers. The p o s s i b i l i t i e s of applying recent research on p a r t i a l c u t t i n g should be explored, e.g., Alexander (1973) Alexander and Edminster (1977) e s p e c i a l l y i n productive moose h a b i t a t s . 3. The major winter food species of moose were i d e n t i f i e d . 423 Habitat manipulations should favour these species. 4. Notwithstanding the above, moose a l s o eat a wide v a r i e t y of species. While the importance of t h i s v a r i e d d i e t i s not yet understood, i t should be considered i n planning such items as leave s t r i p s and cutover s i z e . 5. Since moose appear to s e l e c t c e r t a i n trees f o r bedding, these trees or p o r t i o n s of stands c o n t a i n i n g these, should be reserved from c u t t i n g u n t i l s u i t a b l e r e p l a c e -ments have grown up. 6. In browsing, moose do not remove a l l the p o t e n t i a l l y a v a i l a b l e forage, at l e a s t on the ranges studied. Thus, i t seems moose are below the c a r r y i n g c a p a c i t y of the forage resource. Moose tend to rebrowse shrubs but not i n d i v i d u a l twigs. Management of shrubs should encourage methods th a t s t i m u l a t e production of stems r a t h e r than twigs. 7. In de c i d i n g cutover s i z e and d i s t r i b u t i o n , the s i z e of home ranges and the concept of range mix should be kept i n mind. 8. The production of forage has an e a r l y , abrupt and high peak during f o r e s t succession. Thus, important winter ranges l i k e Grove w i l l be out of s i g n i f i c a n t production i n l e s s than 20 years. To ensure adequate w i n t e r forage f o r moose using these areas, though should be given to e i t h e r managing these burns f o r production of browse, or i n s t i t u t i n g experimental logging on adjacent, s i m i l a r areas to ensure adequate forage. 9. The occupancy of open.portions of winter ranges reaches peak l e v e l s i n December and January. Unless w i n t e r counts bear t h i s i n mind, year-to-year v a r i a t i o n s i n timing surveys may y i e l d f a l s e data regarding numbers of moose and trends i n numbers. 10. Many of the important winter h a b i t a t s of moose co i n c i d e w i t h areas where human a c t i v i t i e s have and w i l l con-tinue to modify the landscape. Some of these a c t i v i -t i e s are incompatible w i t h w i n t e r i n g moose, but others can accommodate moose i f s u f f i c i e n t thought i s given. Thus areas e x i s t close to and even w i t h i n the boundaries of c i t i e s such as P r i n c e George where moose can winter s u c c e s s f u l l y and with minimal e f f e c t on the settlements. For example, r i v e r breaks, flood-prone i s l a n d s where there are p a r t i c u l a r l y rough and unstable s o i l s , and even some of the l e s s developed parks can provide winter h a b i t a t . Such s i t e s have an a d d i t i o n a l advantage as a convenient outdoor l a b o r a t o r y f o r students. Future research needs r e l a t e d to moose and t h e i r h a b i t a t were a l s o i d e n t i f i e d . These needs are considered to be important gaps that need to be overcome i f research i s to a n t i c i p a t e f u t u r e demands f o r knowledge. Some of the p o t e n t i a l t o p i c s have been mentioned i n appropriate s e c t i o n s of the t e x t , but, f o r the sake of convenience, these and rs are l i s t e d b r i e f l y below: Continue study of s u c c e s s i o n a l patterns and rates t o i n c l u d e an improved sample f o r mesic environments, and t o i n c l u d e x e r i c and h y d r i c environments. This research should be done c o o p e r a t i v e l y by f o r e s t r y and w i l d l i f e s ectors on both enclosed and open permanent p l o t s . Study e f f e c t s of logging, methods and s i t e treatments on p l a n t succession, e.g., species composition, rates of coniferous establishment, t h i n n i n g regimes. Determine home range s i z e s f o r various sex and age c l a s s e s of moose, f o r a l l seasons, and f o r broadly d i f f e r e n t environments. Conduct d e t a i l e d s t u d i e s of moose a c t i v i t y along eco-tone s. Evaluate the r o l e and importance of summer range, e s p e c i a l l y aquatic h a b i t a t s . I d e n t i f y migration routes and y e a r l y home ranges. I d e n t i f y herds so t h a t management can be done on a b i o l o g i c a l l y more meaninful herd-basis. Determine through appropriate experiments those f a c t o r s t h a t i n f l u e n c e forage production and n u t r i e n t content. I n v e s t i g a t e the i n f l u e n c e of s o c i a l behaviour on h a b i t a t use and s e l e c t i o n . I n i t i a t e n u t r i t i o n a l and b i o e n e r g e t i c s t u d i e s . Explore computer s i m u l a t i o n as an approach to s e t t i n g 426 c a r r y i n g c a p a c i t y l i m i t s (e.g., Robbins 1973, Wallmo et a l . , 1977), and as a way of assessing f o r e s t r y impacts. 12. Conduct d e t a i l e d s t u d i e s of moose, using telemetry, i n a f o r e s t stand before, during and a f t e r logging. 13. Develop and t e s t r e l i a b l e methods of assessing demands f o r moose. 14. Develop and t e s t r e l i a b l e methods f o r assessing the s t a t u s and trend of moose populations. The foregoing suggestions f o r management and research w i l l improve the a b i l i t y of w i l d l i f e b i o l o g i s t s to improve management of moose. They are s p e c i f i c suggestions that should be considered w i t h one o v e r - r i d i n g f a c t i n mind about the sub-boreal ecosystem. 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Pap. presented at 30th Ann. Meeting Soc. Range Manage., P o r t l a n d , Oregon. Westman, H. 1958. Algens skadegorelse pa ungskogen (on moose damage to f o r e s t r y ) . K. Skogshogsk. Skr. 28. 14 8 pp. Whitford, H. N., and R. D. C r a i g . 1918. Forests of B r i t i s h Columbia. Commission of Conservation Canada. King's P r i n t e r , Ottawa. 409 pp. Whittaker, R. H. 1962. Net production r e l a t i o n s of shrubs i n the Great Smoky Mountains. Ecology 43:35 8-377. . 19 70. Communities and ecosystems. The Macmillan Company, C o l l i e r - M a c m i l l a n Canada L t d . , Toronto. 162 pp. Wilson, D. E. 1971. Ca r r y i n g c a p a c i t y of the key browse species f o r moose on the north slopes of the Uinta Mountains, Utah. Utah Dept. Nat. R e s o u r c , Div. W i l d l . R e s ourc, Pub. 71-9. S a l t Lake C i t y . 57 pp. Wood, A. J . , H. C. Nordan, and I. McT. Cowan. 19 62. P e r i o d i c i t y of growth i n ungulates as shown by deer of the genus Odocoileus. Can. J . Zool. 40(4):59 3-603. Wood, G. W., and J . S. Lindzey. 1967. T h e e f f e c t s of f o r e s t f e r t i l i z a t i o n on the crude p r o t e i n , calcium and phosphorous content of d e e r browse i n a mixed oak f o r e s t . N a t u r a l i s t e can. 94:335-346. Wooldridge, D. D. 1960. Watershed disturbance from t r a c t o r and s k y l i n e crane logging. J . For. 58:369-372. 459 Young, H. E. 19 71. Biomass sampling methods f o r puckerbrush stands i n f o r e s t biomass s t u d i e s . Univ. of Maine, College of L i f e Sciences and A g r i c , Orono. Young, J . A., D. W. Hedrick, and R. F. Keniston. 1967. Forest cover and logging. J . For. 65:807-813. Z a v i t k o v s k i , J . 1976. Ground vegetation biomass, production, and e f f i c i e n c y of energy u t i l i z a t i o n i n some northern Wisconsin f o r e s t ecosystems. Ecology 57(4):694-706. i 460 APPENDIX A SCIENTIFIC AND COMMON NAMES OF PLANT SPECIES RECORDED IN THE STUDY AREA 461 As much as p o s s i b l e s c i e n t i f i c and common names were taken from the f o l l o w i n g references: Brayshaw (1976) f o r w i l l o w and b i r c h f a m i l i e s (Salicaceae and Betulaceae), Clark and Frye (1928) f o r the l i v e r w o r t s (Hepatica), Crum et a l . (1973) and Lawton (1971) f o r the mosses (Musci), Garman (1973) f o r trees and shrubs not covered by the other, more s p e c i f i c works, Howard (1950) f o r the l i c h e n s , Szczawinski (1962) f o r the heather f a m i l y ( E r i c a c e a e ) , Szczawinski (1970) f o r the o r c h i d f a m i l y (Orchidaceae), T a y l o r (1971) f o r the ferns and f e r n - a l l i e s ( P t e r i d o p h y t a ) , Taylor (1973) f o r the rose f a m i l y (Rosaceae), Taylor (1974a) f o r the l i l y f a m i l y ( L i l i a c e a e ) , T a y l o r (1974b) f o r the pea f a m i l y (Leguminoseae), and Tay l o r (1974c) f o r the fi g w o r t f a m i l y (Scrophulariaceae). These references were supplemented by Hulten (1968) and Hitchcock et a l . (1955-1969). P l a n t species are l i s t e d i n a l p h a b e t i c a l order based on t h e i r s c i e n t i f i c names. Common names, as used i n the t e x t or given i n the foregoing r e f e r e n c e s , are also l i s t e d . Major s u b d i v i s i o n s are by forage c l a s s e s , f o l l o w i n g the format used i n the t e x t . EVERGREEN TREES AND SHRUBS: Abies lasiooarpa (Hook.) Nutt. Ledum palustra L. subsp. groenlandioum (Oeder) H u l t . Mahonia nervosa (Pursh) Nutt. Mahonia repens ( L i n d l e . ) G. Don Paohystima myrsinites Raf. Picea glauoa (Moench) Voss ssp. glauoa Picea glauoa (Moench) Voss ssp. engelmanni (Parry) Taylor Pioea mariana ( M i l l . ) B. S. P. Pinus oontorta Dougl. ex Loud. Pseudotsuga menziesii (Mirb.) Franco Thuja plioata Donn Tsuga heterophylla (Raf.) Sarg. DECIDUOUS TREES AND SHRUBS: Acer glabrum Torr. Alnus orispa ( A i t ) Pursh subsp. sinuata (Regel) H u l t . Alnus tenuifolia Nutt Amelanchier alnifolia Nutt. Betula glandulosa Michx. Betula papyrifera Marsh Cornus stolonifera Michx. Corylus oornuta M a r s h a l l Lonioera involucrata (Richards) Banks Lonicera utahensis (Sm.) Miq. Gplopanax horridus (Sm.) Miq. Populus balsamifera L. subsp. triohocarpa (Torr. and Gray) H u l t . Populus tremuloides Michx. Prunus virginiana L. Ribes glandulosum Grauer Ribes lacustra (Pers.) P o i r Ribes L., spp. Rosa aoioularis L i n d l . Rubus idaeus L. Rubus parviflorus Nutt. Rubus L., spp. Salix bebbiana Sarg. var. depilis Raup Salix lasiandra Benth. Salix pedioellaris Pursh Salix sitchensis Bongard Salix L., spp. Sambucus racemosa L. Shepherdia canadensis (L.) Nutt. Sorbus scopulina Greene Sorbus sitchensis Roemer 462 Subalpine f i r Labrador tea Oregon grape Creeping mahonia False-box White spruce Engelmann spruce Black spruce Lodgepole pine Douglas f i r Western red cedar Western hemlock Douglas maple S i t k a a l d e r Mountain a l d e r Saskatoon Bog b i r c h Paper b i r c h Red-osier dogwood Beaked h a z e l nut Black twinberry Red twinberry D e v i l s ' club Black cottonwood Trembling aspen Chokecherry Skunk currant B r i s t l y b l ack currant Gooseberries and currants Rose Raspberry Thimbleberry B l a c k b e r r y , raspberry Bebb's w i l l o w P a c i f i c w i l l o w Bog w i l l o w S i t k a w i l l o w Willow E l d e r b e r r y Soapberry Mountain ash S i t k a mountain ash Spiraea dougtasi Hook. Spiraea lucida Dougl. ex Hook. Symphorioarpos albus (L.) Blake Vaaainium caespitosum Michx. Vaccinium membranaceum Dougl. Vaccinium myrtilloides Michx. Vaccinium myrtillus L. Vaecinium L., spp. Viburnum edule (Michx.) Raf. FORBS AND DWARF SHRUBS: Achillea millefolium L. Aotaea rubra ( A i t . ) W i l l d . Anaphalis margaritaoea (L.) Benth and Hook f. Antennaria neglecta Greene Aquilegia formosa F i s c h . Aralia nudioaulis L. Aratostaphylos uva-ursi (L.) Spreng. Arnica ohamissonis Less. Arnica cordifolia Hook Aster conspicuus L i n d l . Aster foliaceus L i n d l . Aster junciformis Rydb. Aster modestus L i n d l e . Aster L., spp. Calypso bulbosa ( 1 . ) Rchb. f:. Camassia quamash (Pursh) Greene Castilleja miniata Dougl. Cerastium arvense L. Chimaphila menziesii (R. Br.) Spreng. Chimaphila umbellata (L.) Barton Cirsium undulatum (Nutt.) Spreng. CUntonia uniflora (Schult.) Kunth Comandra livida Richards Corallorhiza striata L i n d l . Cornus canadensis L. Corydalis sempervirens (L.) Pers. Disporum oreganum (Wats.) Q. Jone Epilobium angustifolium L. Epilobium latifolium L. Epilobium minutum L i n d l . ex Hook. Erigeron acris L. Erigeron L., spp. Eriogonum Michx., spp. Fragaria virginiana Duch. Galeopsis tetrahit L. Galium boreale L. Galium triflorum Michx. Gaulthe.ria hispidula (L.) Muhl. Hardhack F l a t - t o p s p i r e a Waxberry Dwarf blueberry Mountain b i l b e r r y V e l v e t - l e a f blueberry Canada blueberry V a c c i n i a Squashberry Yarrow Banebe'rry P e a r l y e v e r l a s t i n g Pussytoe Western columbine S a r s a p a r i l l a Bearberry A r n i c a Heat-leaf a r n i c a A s t e r A s t e r Aster A s t e r Aster Calypso o r c h i d Common camas Indian paintbrush Chickweed L i t t l e p r i n c e ' s pine P r i n c e ' s pine T h i s t l e Queen's cup Bastard t o a d - f l e x S t r i p e d c o r a l - r o o t Bunchberry Pale c o r y d a l i s Oregon f a i r y b e l l s Fireweed R i v e r beauty Fireweed Fleabane Fleabanes Umbrella p l a n t W i l d strawberry Hemp n e t t l e Northern bedstraw Sweet-scented bedstraw Creeping snowberry 464 Gentiana amarella L. Geranium bicknelli B r i t t . Geranium L., spp. Geum maorophy Hum W i l l d . Goody era oblongi folia Raf. Goody era repens (L.) R. Br. Eabeneria orbiculata (Pursh) Torr. Habenaria unalasoensis (Spreng.) Wats. Habenaria W i l l d . , spp. Eeuchera L., spp. Eieracium albiflorum Hook. Hieraoium oanadense Michx. Eieracium cynoglossoides Arv.-Touv. Eypochaeris radioata L. Lathyrus ochroleucus Hook. Lilium columbianum Hanson Linnaea borealis L. Listeria oaurina :Piper Lycopus uniflorus Michx.. Maianthemum dilatatum (How) Nels. and MacBr. Matricaria matricarioides (Less.) P o r t e r Melampyrum lineare Desr. Mimulus guttatus D.C. Mitella nuda L. Myriophyllum L. , spp. Nuphar variegatum Engelm. Parnassia palustris L. Pedicularis L., spp. Penstemon albertinus Greene Petasites frigidus (L.) French. Polygonum amphibium L. Potamogeton alpinus B a l b i s Potentilla palustris (L.) Scop. Prunella vulgaris L. Pyrola elliptica Nutt. Pyrola minor L. Pyrola seounda L. Pyrola virens Schweigg Pyrola L., spp. Ranunculus oooidentalis Nutt. Rubus aroticus L. Rubus chamaemorus L. Rubus pedatus J . E. Smith Scheuchzeria palustris L. Scutellaria galericulata L. Senecio indecorus Greene Smilacina raoemosa (L.) Desf. Solidago canadensis L. Sparganium euryoarpum Engelm. Gentian B i c k n e l l ' s geranium Geranium Avens Rattlesnake p l a n t a i n Rattlesnake p l a n t a i n Re i n - o r c h i d Re i n - o r c h i d R ein-orchid Heuchers Hawkweed Canada hawkweed Hawkweed Cat's ear Cream-flowered peavine Tiger l i l y Twinflower Northwest twayblade Bugleweed Two-leaved Solomon's s e a l Pineapple weed Cow-wheat Monkey-f1 owe r Mitrewort Water m i l f o i l Water l i l y Northern grass-of-Parnassus Lousewort Penstemon C o l t s f o o t Water smartweed Pondweed Marsh f i v e f i n g e r S e l f - h e a l W i l d l i l y - o f - t h e - v a l l e y Lesser wintergreen One-sided wintergreen Greenish flowered wintergreen Wintergreen Buttercup A r c t i c bramble Cloudberry T r a i l i n g rubus Scheuchzeria Skullcap Groundsel False Solomon's s e a l Goldenrod Bur-reed Sparganium minimum F r i e s Sparaganium multipedunaulatim ' (Morong) Rydg. Streptopus amplexifolius (L.) D.C. Taraxacum officinale Weber Tiarella unifoliata Hook. Trifolium repens L. Typha latifolia L. Urtica dioica L. Valeriana sitchensis Bong. Veratrum viride A i t Veronica americana Schw. Vicia americana Muhl. Viola glabella Nutt. Zannichellia palustris L. GRAMINOIDS: Agropyron caninum (L.) Beauv. Agropyron repens (L.) Beauv. Agrostis scabra W i l l d . Agrostis L., spp. Aira caryophyllea (L.) Nash Alopecurus aequalis Sobol. Bromus secalinus L. Calamagrostis canadensis (Michx.) Beauv. Calamagrostis inexpansa Gray Calamagrostis montanensis S c r i b n . Calamagrostis rubescens Buckl. . Carex aena Fern. Carex arcta Boott Carex limosa L. Carex microptera Mack. Carex rostrata Stoke Carex L., spp. Cinna latifolia (Trev.) Griseb. Elymus glaucus B u c k l . Festuca idahoensis Elmer. Festuca occidentalis Hook. Glyceria borealis (Nash) Batchelder Glyceria maxima (Hartm.) Holmb. subsp. grandis (S. Wats.) Hult J uncus bufonius L. Luzula parviflora (Ehrh.) Deav. Melica subulata (Griseb.) S c r i b n . Oryzopis asperifolia Michx. Phleum pratense L. Poa palustris L. Trisetum spicatum (L.) R i c h t e r 465 Bur-reed Bur-reed Twisted s t a l k Dandelion Foam flower White c l o v e r C a t - t a i l N e t t l e V a l e r i a n Green f a l s e h e l l e b o r e Brooklime American vetch Yellow v i o l e t Horned pond weed Wheatgrass Quackgrass Bentgrass Bentgrass H a i r grass F o x t a i l Chess B l u e j o i n t Northern reedgrass P l a i n s reedgrass Pinegrass Sedge Sedge Sedge Sedge Sedge Sedge Wood reed grass Blue w i l d - r y e Idaho fescus Western fescue Northern mannagrass American mannagrass Rush Wood rush A l a s k a oniongrass Mountain r i c e g r a s s Timothy Fowl bluegrass Spike t r i s e t u m FERNS AND FERN ALLIES: Athyrium filix-femina L. Roth Botvyohium multi.fi.dum (Omel.) Rupr. Dryopteris austriaca (Jacq.) Woynar. Gymnocarpium dvyopteris (L.) Newm. Lyoopodium annotinum L. Lyoopodium olavatum L. Lyoopodium complanatum L. Lyoopodium L., spp. Matteuccia struthiopteris (L.) Todar.o Ladyfern Leathery grape-fern Spiny wood-fern Oakfern S t i f f club-moss Running club-moss Ground-cedar Clubmoss O s t r i c h f e r n HORSETAILS: Equisetum. arvense L. LIVERWORTS: H o r s e t a i l Lophozia. Dum. spp. Marchantia polymorpha L. MOSSES: Brachytheoium B. S. G., spp. Calliergon ( S u l l . ) Kindb., spp. Dicranum Hedw., spp. Drepanocladus unoinatus (Hedw.) Warnst, Eurhynchium B.S., spp. Orthotrichum Hedw., spp. Pohlia Hedw., spp. Polytrichum Quniperinum Hedw. Ptilium crista-oastrensis (Hedw.) De Not. H a i r cap moss LICHENS; Alectoria Ach., spp. Cladonia ( H i l l ) Web., spp. Lob aria pulmonaria (L.) Hoffm. Parmelia Ach. , spp. Platismatia glauca (?) Usnea ( D i l l ) Adans., spp. Old man's beard Lungwort APPENDIX B SCIENTIFIC AND COMMON NAMES OF BIRD AND MAMMAL SPECIES MENTIONED FOR THE STUDY AREA 468 S c i e n t i f i c and common names were taken from C a r l et a l . (1959) f o r f i s h e s , from Godfrey (1966) f o r b i r d s , and from Cowan and Guiget (19 73) f o r mammals. Species are l i s t e d a l p h a b e t i c a l l y w i t h i n each c l a s s , based on t h e i r s c i e n t i f i c names. INSECTS: Choristoneura fumiferana (Clemens) FISHES: Onoorhynohus (Walbaum), spp. BIRDS: Bonasa vmbellus umbelloides (Douglas) Branta canadensis (Linnaeus) Falco sparverius sparverius Linnaeus Passerina ameona (Say) Sturnella negleota confluenta Rathbun Spruce budworm Salmon Ruffed grouse Canada goose Sparrowhawk L a z u l i bunting Western meadowlark MAMMALS: Aloes aloes andersoni Peterson Canis latvans inoolatus H a l l Canis lupus oolumbianus Goldman Castor canadensis sagittatus Benson Erethizon dorsatum nigresoens A l l e n Felis ooncolor missoulensis Goldman Lepus americanus pallidus Cowan Lynx canadensis canadensis K e r r Lynx rufus pallescens Merriam Martes americana abietinoides Gray Odocoileus hemionus hemionus (Rafinesque) Rangifer tarandus montanus Seton Ursus americanus cinnamomum (Audubon and Bachman) Ursus arctos horribilis Ord Moose Coyote Wolf American beaver Porcupine Cougar Varying hare Canada ly n x Bobcat Marten Mule deer Mountain caribou American b l a c k bear G r i z z l y bear APPENDIX C STATISTICAL DATA USED FOR THE INTRODUCTION (SECTION 1) 470 TABLE C - l Estimated Annual Economic Value of Minerals Produced i n the Omineca Mining D i s t r i c t , 1926-1974 Year Value ( i n IO 4 d o l l a r s ) Year ( i n Value 10h d o l l a r s ) 1926 25 1951 205 1927 16 1952 363 1928 31 1953 384 1929 23 1954 339 1930 6 1955 99 1931 3 1956 189 1932 1 1957 228 1933 1 1958 93 1934 9 1959 40 1935 10 1960 1,402 1936 13 1961 40 1937 14 1962 51 1938* 15 1963 32 1939 m** 1964 44 1940 m 1965 1,071 1941 155 1966 2,347 1942 32 7 1967 3,945 1943 536 1968 3,422 1944 141 1969 494 1945 14 19 70 4,131 1946 7 1971 2,869 1947 10 1972 3,552 1948 21 1973 9, 735 1949 79 1974 10,782 1950 165 *Data p r i o r to 1938 based on Omineca and Peace D i v i s i o n s . **Missing data. TABLE C-2 Number and T o t a l Province and Area of Farms f o r the P r i n c e , and Number of George Region, C a t t l e f o r the 1881 - 1971 ^ Year No. of P r o v i n c i a l farms Regional 2 Area of farms (km.'.) P r o v i n c i a l Regional No. of P r o v i n c i a l c a t t l e Regional 1881 2,743 1,786 80,451 1891 6,490 126,919 1901 6,501 6,060 125,002 1911 16,958 10,279 36 139,184 1921 21,973 11,576 5,824 219,058 101,289 1931 26,079 2,047 14,332 2,299 233,923 28,333 1941 26,394 2,305 16,323 2,495 333,873 42,479 1951 26,406 19,029 1956 24,748 18,368 1961 19,934 1,362 18,237 2,582 52,820 1966 19,085 1,406 3,655 77,445 1971 18,400 629 573,171 11,417 Note: Regional census area f o r 1881 - 1931 was the Cariboo e l e c t o r a l d i s t r i c t ; f o r 1941 - 1966, Census D i s t r i c t 8; and i n 1971, the F r a s e r - F t . George Regional D i s t r i c t . Area of the province i s 930,528 km ; the Cariboo e l e c t o r a l d i s t r i c t i s 81,417 km ; the r e g i o n a l Census D i s t r i c t (8) i s 186,440 km^; and the Fraser-Fort George Regional D i s t r i c t i s 51,196 km^. 472 Table C-3 Annual Cut of Timber ( A l l Species) and the Number of Sawmills Operating i n the P r i n c e George Forest D i s t r i c t , 1909 - 1975* - r No. of m i l l s Annual No. of m i l l s Annual Year operating c u t * * Year operating c u t * * 1909 1 m 1942 72 61 1910 3 m 1943 118 95 1911 m m 1944 141 63 1912 m m 1945 174 71 1913 m m 1946 210 91 1914 16 4 1947 329 114 1915 16 6 1948 345 146 1916 16 3 1949 384 130 1917 24 7 1950 447 158 1918 19 13 1951 551 212 1919 20 3 1952 604 243 1920 25 27 1953 689 243 1921 26 21 1954 700 221 1922 23 21 1955 730 300 1923 23 32 1956 687 321 1924 26 51 1957 704 318 1925 30 48 1958 669 326 1926 23 58 1959 648 413 1927 17 48 1960 621 424 1928 22 66 1961 564 416 1929 23 66 1962 543 459 1930 31 34 1963 493 531 1931 35 17 1964 450 574 1932 23 11 1965 368 596 1933 23 12 1966 299 623 1934 31 21 1967 258 674 1935 40 24 1968 213 776 1936 48 27 1969 943 1937 44 32 1970 1,062 1938 50 32 1971 1,153 1939 43 33 19 72 1,154 1940 61 43 1973 1,136 1941 88 53 1974 1975 1,100 957 *Data from annual reports of the B r i t i s h Columbia Forest S e r v i c e . **Annual cut expressed as 10 m . * * * S t a t i s t i c discontinued. 473 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 Year Area (km2) dis t u r b e d by: Year Area (km2) d i s t u r b e d by: w i l d f i r e l o g g i n g w i l d f i r e logging 1912 121 1944 1,787 1913 1 1945 728 1914 186 1946 655 1915 342 1947 475 1916 425 1948 1,503 1917 8 1949 501 1918 62 1950 3,120 ( 00 Table D-5 Data from P e l l e t Group Transects f o r Synoptic Surveys i n 1973 Study Transect - 1 -area l a b e l length P e l l e t groups* Age E l e v a - Sub-(no.) (m) W S Habitat (yr) t i o n (m) s t r a t e * * * Remarks Bowron B 1 335 1(8) - c o n i f e r f o r e s t 740 T pine (01) B 2 1,006 - - c l e a r c u t 2 740 T burned B 3 1,006 8 - c l e a r c u t 2 740 T unburned Eagle E 1 305 32(19) _ c o n i f e r f o r e s t M 710 L spruce-pine, ecotone (02) E 2 305 8(12) - burn 36 700 L at ecotone w i t h E l E 3 305 18(4) 4 p a r t i a l cutover 10(?) 710 L s e l e c t i v e cut E 4 305 4(5) - burn 36 700 L center of burn Grove G 1 305 3(2) 1 burn 12 765 T at ecotone w i t h G2 (03) G 2 305 3(3) - c o n i f e r f o r e s t M 765 T spruce-pine G 3 305 24(10) 1 burn 12 765 T at ecotone w i t h G4 G 4 305 4(5) 2 c o n i f e r f o r e s t M 765 T spruce-pine G 5 305 K D 1 burn 12 1,070 T exposed s i t e G 6 335 17(16) 1 burn 12 765 L burn center McKenzie M 1 305 1(16) 1 c l e a r c u t 3 910 T b urne d (04) M 2 305 2(1) - c l e a r c u t 3 910 T burned M 3 335 6(5) 1 c l e a r c u t 5 910 T unburned M 4 335 4 - c l e a r c u t 5 910 T M 5 305 42(19) - p a r t i a l cutover 11 910 T s e l e c t i v e cut M 6 305 18(17) 1 p a r t i a l cutover 11 910 T M 7 305 6(2) 1 p a r t i a l cutover 5(?) 910 T s e l e c t i v e , seedblock M 8 305 6(2) 1 p a r t i a l cutover 5(?) 910 T M 9 305 4 1 c o n i f e r f o r e s t M 910 T spruce-pine M10 305 -(1) - c o n i f e r f o r e s t M 910 T M i l 305 -(4) - c l e a r c u t 1 910 T burn Ml 2 305 -(4) - c l e a r c u t 1 910 T CO Table D-5, Continued Study area (no.) Transect + l a b e l length P e l l e t groups* Age El e v a - Sub-(m) W S Habitat (yr) t i o n (m) s t r a t e * * * Remarks Ml 3 305 -(1) c l e a r c u t 1 910 T Ml 4 305 28(18) 2 p a r t i a l cutover 11 910 T M15 305 4(1) c o n i f e r f o r e s t M 910 T pine Ml 6 305 1(3) c o n i f e r f o r e s t 90 910 T immature pine Ml 7 610 1(2) road 3 910 T through Ml, M2 M18 305 4(2) - s k i d t r a i l 3 910 T through Ml, M2 M19 914 2(12) 1 road 5 910 T through M3, M4 M20 244 1(8) s k i d t r a i l 5 910 T through M3, M4 M21 610 3(13) road 5 910 T through M5, 6, 14 M22 610 1(7) road 11 910 T through M5, 6, 14 M23 671 25(34) road 11 910 T through M5, 6, 14 M24 605 10(80) road 11 910 T through M5, 6, 14 M25 305 14(9) s k i d t r a i l 5(?) 910 T through M7, M8 M26 305 5(11) 1 s k i d t r a i l 5(?) 910 T through M7, M8 S 1 305 3 mixed f o r e s t 680 B aspen-pine, same as Sl-1972 S 2 945 26(8) - c o n i f e r f o r e s t M 715 T pine-spruce, same as S2-1972 S 3 305 7 p a r t i a l cutover 15(?) 775 T cut and leave S 4 335 - mixed f o r e s t 535 RA spruce-cottonwood, r i v e r bottom S 5 305 8(1) p a r t i a l cutover 10(?) 670 RA spruce-cottonwood, r i v e r bottom S 6 1,602 72(21) road M 730 T mainly i n f o r e s t S 7 1,463 3(11) 6 c o n i f e r f o r e s t M 701 T-L pine-spruce S 8 1,372 3(9) 1 c o n i f e r f o r e s t M 701 T-L pine-spruce S 9 1,433 14(15) 2 c o n i f e r f o r e s t M 701 T-L pine-spruce S10 1,981 3(10) c o n i f e r f o r e s t M 730 T pine-spruce S l l 762 2(6) p a r t i a l cutover 670 L S12 259 -(3) c o n i f e r f o r e s t M 670 T Salmon (05) 4>-OO Table D-5, Continued S tudy Transect area l a b e l length P e l l e t groups* Age E l e v a - Sub-(no.) (m) W S Habitat (yr) t i o n (m) s t r a t e * * * Remarks S13 394 5(18) 2 p a r t i a l cutover 670 T s e l e c t i o n cut S14 93 2(2) - c o n i f e r f o r e s t M 670 T S15 124 K D - c o n i f e r f o r e s t M 670 T S16 373 3(5) - p a r t i a l cutover 730 T S17 311 0(1) - c o n i f e r f o r e s t M 730 T S18 311 4(13) - p a r t i a l cutover 730 T s e l e c t i o n cut S19 187 0(3) - c o n i f e r f o r e s t M 730 T S20 539 4(10) - p a r t i a l cutover 730 T s e l e c t i o n cut S21 610 6(8) - p a r t i a l cutover 730 T S22 701 34(12) -' c o n i f e r f o r e s t M 730 T S23 244 2(9) 2 p a r t i a l cutover 730 T unburned S24 305 7(3) 1 c o n i f e r f o r e s t M 730 T S25 305 7(8) 4 p a r t i a l cutover 900 T S26 305 7(10) 3 p a r t i a l cutover 900 T S27 914 25(19) - road 900 T through S25, S26 S28 610 19(29). - road 715 T through S3 *Counts are l a s t winter's groups (no. of older groups i n parentheses). **M = mature f o r e s t stand, older than 100 y r s . ***Substrate symbols are: T f o r drumlinized t i l l , L f o r l a c u s t r i n e , RA f o r recent a l l u v i u m , B f o r beach deposits. +A11 t r a n s e c t s were 3 m wide. 4>-00 ON 487 APPENDIX E CHARACTERISTICS OF SAMPLES COLLECTED FOR THE FOOD HABITS STUDY (SECTION 4) 488 Table E - l Rumen Samples: Date of K i l l , Sex, Age, and Loca t i o n of K i l l Date of k i l l Sex* Age* Lo c a t i o n of k i l l M.U.** 1971 Sept. M A Fyf e :.Lake 7-10 Oct. 6 M ' 8.5 Tzenzaicut Lake 5-13 Oct. U A Corkscrew Creek 7-12 Nov. 11 F 1.5 Barney Creek 7-16 Nov. 11 M 1.5 Barney Creek 7-16 Nov. 11 M 5.5 Barney Creek 7-16 Nov. 11 M 9.5 Salmon R i v e r (north side) 7-15 Nov. 14 M A Salmon R i v e r (north side) 7-15 Nov. 15 F 10.5 Purden Lake 7-7 Nov. 20 M A McGregor settlement 7-17 Nov. 25 M 0.5 Garvin Creek 7-15 Nov. 27 M A McGregor settlement 7-17 Nov. 28 M A McGregor settlement 7-17 Nov. U 0.5 Nechako Ri v e r (north side) 7-13 Nov. M A Nechako R i v e r (north side) 7-13 Nov. F A Nechako River (north side) 7-13 Nov. F 2.5 Salmon R i v e r (north side) 7-15 Nov. F 1.5 Salmon R i v e r (north side) 7-15 Nov. M A Barney Creek 7-16 Nov. F 8.5 Barney Creek 7-16 Nov. F 2.5 Barney Creek 7-16 Nov. U A Barney Creek (?) 7-16 Nov. M 9.5 McGregor settlement 7-17 Dec. U 0.5 Hansen Lake (north side) 6-5 1972 Jan. 18 M 2.5 Wansa Creek 7-7 Jan. M A Wansa Creek 7-7 Jan. U A Vama Varna Creek 7-7 Jan. U A Spey Creek 7-9 Feb. 2 U 0.5 Vama Vama Creek 7-7 Feb. 2 F 5.5 Vama Vama Creek 7-7 Feb. 2 F 0.5 Murray Creek 7-13 Feb. U U Not s p e c i f i e d 7-5 ? Mar. 2 M 0.5 Vanderhoof 7-13 Mar. 2 M A Mi l e 66, Hart Highway 7-16 Mar. 1-6 M 4.5 Baldy Hughes settlement 7-10 Mar. 6 F 0.5 Vanderhoof 7-13 Mar. M A Barney Creek at Fraser R i v e r 7-16 Apr. 19 F A Newlands 7-7 489 Table E - l , Continued Date of k i l l Sex* Age :* L o c a t i o n of k i l l M.U.** Oct. F A Eagl e t Lake 7-7 Oct. U A Eagl e t Lake 7-7 Nov. 7 F A Salmon R i v e r (north side) 7-15 Nov. F A McGregor settlement 7-17 Nov. F A Barney Creek 7-16 1973 Jan. 13 F A Vama Vama Creek 7-7 Jan. 19-26 M A Ke l l o g g Creek 7-12 Jan. 19-26 M 1. 5 K e l l o g g Creek 7-12 Jan. F A Vama Vama Creek 7-7 Jan. U A Willow R i v e r (?) 7-7 Jan. M A Chilako R i v e r 7-12 Feb. 4 F A Spey Creek 7-9 Feb. 8 M A Sl i m Creek 7-5 Feb. 26 F A Spey Creek 7-9 Feb. 26 M A Spey Creek 7-9 Feb. 26 M A Spey Creek 7-9 Mar. 1 F A Hart Highway (No. 97) 7-15 Sept. 15 U U Salmon R i v e r (north side) 7-15 Sept. 26 M A Salmon R i v e r (south side) 7-14 Sept. 27 M A Salmon R i v e r (south side) 7-14 Sept. 27 M A Salmon R i v e r (south side) 7-14 Oct. 14 U 0. 5 Salmon R i v e r (north side) 7-15 Oct. 23 U U Salmon R i v e r (north side) 7-15 Oct. F U Salmon R i v e r (north side) 7-15 *Abb r e v i a t i o n f o r sex: M - male, E - female, U - u n c l a s s i f i e d , and f o r age: A - adult 1.5 years or o l d e r , U - unaged. * * F i s h and W i l d l i f e Branch Management Unit. APPENDIX F SUCCESSION DATA (FOR SECTION 7) 491 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 MAJOR GROUP P l a n t species . S i t e number MF2 MF3 MF4 MF5 MF6 MF7 EVERGREEN TREES & SHRUBS: Abies lasiooarpa Picea glauca Pinus contovta DECIDUOUS TREES & SHRUBS: Acer glabrum Alnus spp. Amelanchier alnifolia Betula papyrifera Lonicera involucrata Populus tremuloides Ribes spp. Rosa spp. Rubus idaeus R. parviflorus Salix spp. Shepherdia canadensis Spiraea luoida S. douglasii Vaooinium oaespitosum V. membranaoeum V. myrtillus Vaooinium spp. Viburnum edule FORBS & DWARF SHRUBS: Achillea millefolium Aralia nudioaulis 3/16 4/10 2/8 1/20 5/18 6/2 1/2 16/30 11/14 t/2 2/8 t/4 1/6 2/14 3/18 6/30 1/2 t/6 13/64 1/6 3/6 18/74 1/8 11/46 1/12 2/4 t/8 2/12 1/4 t/4 2/2 t/2 9/46 t/2 5/12 t/2 5/26 8/36 12/66 t/2 10/52 t/6 2/30 t/2 4/14 t/2 4/14 5/26 2/8 5/10 4/36 2/24 t/2 4/48 7/40 4/38 5/38 3/18 492 Table F - l , Continued MAJOR GROUP S i t e number P l a n t species MF2 MF3 MF4 MF5 MF6 MF7 Arnica spp. 1/8 4/28 2/22 t/4 Aster spp. t/2 2/10 1/8 2/10 Chimaphila umbellata t/6 Clintonia uniflora t/10 4/32 t/4 6/28 Cornus canadensis 1/18 12/78 4/24 2/30 9/82 16/78 Disporum oreganum 2/12 t/2 t/2 5/12 Epildbium angustifolium 9/44 5/28 3/10 5/44 11/42 Galium spp. t/2 Geranium bicknellii 38/80 17/50 7/16 Hypochaeris radicata Linnaea borealis 7/32 13/56 . t/10 t/2 1/6 Mitella nuda Petasites frigidus Pyrola spp. 1/10 Rubus chamaemorus R. pedatus 2/42 Smilacina raoemosa 5/8 Streptopus amplexifolius Taraxacum spp. t/4 Tiarella unifoliata t/2 Trifolium repens Vicia spp. GRAMINOIDS: Agropyron repens Agrostis spp. 9/48 Calamagrostis t/2 20/34 1/4 Carex Spp. 3/30 8/16 9/38 1/10 Cinna latifolia t/2 Festuca spp. t/2 Oryzopsis asperifolia 493 Table F - l , Continued MAJOR GROUP S i t e number P l a n t species MF2 MF3 MF4 MF5 MF6 MF7 Poa palustris Unknown OTHER TAXA: Dryopteris austriaca Equisetum No. species recorded No. quadrats 19 33 23 50 50 50 5/18 t/4 16 31 23 50 50 50 *Species w i t h at l e a s t one case of canopy-cover > 5%. 494 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 MAJOR GROUP P l a n t species S i t e number MF8 MF9 MF10 MF11 MF12 MF13 EVERGREEN TREES & SHRUBS: Abies lasiocarpa 20/42 Picea glauca Pinus contorta 1/8 DECIDUOUS TREES & SHRUBS: Acer glabrum 1/4 Alnus spp. 4/6 Amelanchier alnifolia ' Betula papyrifera t/2 Loniaera involucrata 2/6 1/2 Populus tremuloides. Ribes spp. 1/12 Rosa spp. 8/24 1/4 Rubus idaeus 6/22 R. parvifloras 7/28 Salix spp. Shepherdia canadensis Spiraea lucida 6/32 3/20 S. douglasii Vaccinium caespitosum V. membranaaewn V. myrtillus Vaccinium spp. 3/34 10/46 Viburnum edule 2/10 1/6 FORBS & DWARF SHRUBS: Achillea millefolium Aralia nudicaulis 3/34 3/10 t/2 1/10 t/2 2/6 1/18 t/4 1/20 1/6 7/50 8/28 4/8 6/24 16/36 1/6 1/14 2/4 1/2 3/8 2/6 3/12 3/14 t/4 8/38 2/14 1/2 1/6 1/6 6/28 6/36 t/2 1/18 2/12 9/60 1/10 2/6 1/2 t/2 7/22 15/26 2/20 4/30 1/10 2/10 2/12 2/8 7/32 1/4 1/8 9/56 7/30 4/32 495 Table F-2, Continued MAJOR GROUP S i t e Number P l a n t species MF8 MF9 MF10 MF11 MF12 MF13 Arnica spp. t/6 Aster spp. 3/14 6/20 4/30 9/70 Chimaphila umbellata t/8 1/6 1/10 Clintonia uniflora t/4 6/10 2/14 1/12 1/6 Cornus canadensis 12/66 7/32 21/88 12/58 9/80 8/60 Disporum oreganum t/2 Epilobium angustifolium 28/92 15/66 2/18 6/34 12/68 Galium spp. 1/6 \u00E2\u0080\u00A2t/2 t/6 13/42 Geranium bicknellii t/2 t/2 t/2 Hypoohaeris radioata Linnaea borealis 1/6 1/6 4/4 9/38 1/8 Mitella nuda Petasites frigidus 3/58 17/88 Pyrola spp. 1/8 1/6 3/28 t/16 6/32 Rubus ohamaemorus 3/20 7/34 R. pedatus t/4 \u00E2\u0080\u00A2 1/6 1/8 1/6 Smilaoina racemosa 3/14 1/6. t/2 Strep topus amplexifolius Taraxacum spp. t/2 t/2 1/16 8/38 Tiarella unifoliata t/2 t/6 Trifolium repens Vicia spp. 5/20 GRAMINOIDS: Agropyron repens 7/28 Agrostis spp. t/2 Calamagrostis spp. t/2 2/6 3/18 Carex spp. 1/6 1/8 18/64 4/20 Cinna latifolia Festuca spp. 3/10 Oryzopsis asperifolia Table F-2, Continued 496 MAJOR GROUP P l a n t species S i t e Number MF8 MF9 MF10 MF11 MF12 MF13 Poa palustris Unknown OTHER TAXA: Bryopteris austriaaa Equisetum spp. No. species recorded No. quadrats 2/10 34 50 19 50 33 50 4/18 1/10 34 50 30 50 41 50 *Species w i t h at l e a s t one case of canopy-cover > 5%. 497 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 MAJOR GROUP S i t e number P l a n t species MF14 MF15 MF16 MF17 MF18 MF19 EVERGREEN TREES & SHRUBS: Abies lasiocarpa 23/62 13/34 10/24 2/2 Picea glauca t/1 1/4 10/24 2/10 Pinus contorta 2/2 1/4 1/4 DECIDUOUS TREES & SHRUBS: Acer glabrum Alnus spp. 21/40 Amelanchier alnifolia 1/4 8/28 2/10 t/2 Betula papyrifera 10/52 2/6 Loniaera involucrata 3/10 t/4 6/30 4/24 1/14 2/2 Populus tremuloides 7/14 1/4 2/16 t/2 t/2 Ribes spp. t/2 Rosa spp. 2/16 6/54 2/18 7/50 15/66 11/46 Rubus idaeus t/4 t/4 2/14 R. parviflorus t/4 t/2 3/12 2/14 t/4 1/6 Salix spp. 1/2 t/4 37/90 t/4 Shepherdia canadensis Spiraea lucida 4/22 1/10 11/48 2/8 3/14 11/54 S. douglasii t/2 1/8 6/12 17/44 Vaccinium caespitosum 1/12 11/14 1/8 ' 2/20 V. membranaaeum 1/4 10/36 4/20 V. myrtillus Vaccinium spp. 1/6 Viburnum t/2 1/12 4/18 FORBS & DWARF SHRUBS: Achillea millefolium 1/10 1/2 2/16 Aralia nudicaulis t/6 t/6 11/46 1/10 t/2 498 Table F-\u00E2\u0080\u00A23, Continued MAJOR GROUP S i t e number P l a n t species MF14 MF15 MF16 MF17 MF18 MF19 Arnica spp. 2/14 Aster spp. 5/22 14/76 5/24 7/44 1/4 3/20 Chimaphila umbellata 1/20 Clintonia uniflora t/2 7/46 1/10 4/14 Cornus canadensis 30/90 9/78 16/74 4/38 11/66 9/46 Disporum oreganum 2/10 Epilobium angustifolium t/4 7/50 7/62 22/68 Galium spp. 5/28 1/6 2/16 t/4 Geranium bicknellii 5/34 Hypochaeris radicata 1/2 Linnaea borealis 4/30 t/6 2/30 t/6 t/6 14/44 Mitella nuda Petasites frigidus 10/56 6/46 2/24 10/72 Pyrola spp. 1/16 t/8 t/6 1/6 t/2 1/30 Rubus ohamaemorus k i l l 8/52 3/22 4/28 1/8 R. pedatus 1/6 7/28 9/28 t/2 1/12 Smilacina racemosa 1/4 t/2 t/2 Streptopus amplexifolius Taraxacum spp. 2/18 2/18 Tiarella unifoliata Trifolium repens 1/4 Vioia spp.. t/8 t/2 GRAMINOIDS: Agropyron repens t/2 Agrostis spp. Calamagrostis spp. 2/14 2/6 t/4 17/36 3/8 9/20 Carex spp. 3/20 t/2 2/6 Cinna latifolia 6/14 13/44 1/12 Festuca spp. 2/8 8/18 7/16 1/4 Oryzopsis asperifolia Table F-3, Continued MAJOR GROUP S i t e number P l a n t species MF14 MF15 MF16 MF17 MF18 MF19 Poa \u00E2\u0080\u00A2palustris Unknown 2/2 t/2 t/2 OTHER TAXA: Dryopteris austriaca Equisetum spp. t/8 t/2 No. species recorded 34 43 47 41 31 44 No. quadrats 50 50 50 50 50 50 *Species w i t h at l e a s t one case of canopy-cover > 5%. 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 MAJOR GROUP S i t e number P l a n t species MF20 MF21 MF22 EVERGREEN TREES & SHRUBS: Abies lasiocarpa t/2 1/2 Picea glauca 3/10 4/24 Pinus contorta t/2 25/46 DECIDUOUS TREES & SHRUBS: Acer glabrum Alnus spp. 2/4 Amelanchier alnifolia 1/2 2/6 Betula papyrifera 3/6 2/14 Lonicera involucrata 4/10 4/18 t/2 Populus tremuloides t/4 1/4 Ribes spp. t/4 Rosa spp. 1/10 2/14 10/50 Rubus idaeus t/2 t/2 R. parviflorus 1/8 Salix spp. 7/16 9/32 Shepherdia canadensis Spiraea lucida 4/12 7/52 S. douglasii Vaccinium caespitosum t/2 V. membranaceum t/2 2/20 V. myrtillus Vaccinium spp. 4/18 Viburnum edule 5/26 t/2 FORBS & DWARF SHRUBS: Achillea millefolium t/2 1/14 Aralia nudicaulis 18/62 501 Table F-4, Continued MAJOR GROUP S i t e number P l a n t species MF20 MF21 MF22 Arnica spp. Aster spp. 7/36 Chimaphila umbellata Clintonia uniflora 2/2 1/10 Cornus canadensis 13/68 12/58 Disporum oreganum Epilobium angustifolium 8/50 2/18 Galium spp. 1/10 t/4 Geranium bicknellii t/2 Hypochaeris radicata 6/54 Linnaea borealis \u00E2\u0080\u00A2 t/2 18/56 14/52 Mitella nuda Vetasites frigidus 2/28 Pyrola spp. 4/28 t/6 Rubus chamaemorus t/2 t/2 R. pedatus t/2 Smilacina racemosa 1/6 Streptopus amplexifolius Taraxacum spp. 4/30 Tiarella unifoliata Trifolium repens 8/32 Vicia spp. GRAMINOIDS: Agropyron repens t/4 Agrostis spp. 10/34 Calamagrostis spp. 7/32 3/10 Carex spp. 2/10 6/20 Cinna latifolia Festuca spp. 2/16 Oryzopsis asperifolia 502 Table F-4, Continued MAJOR GROUP S i t e number P l a n t species MF20 MF21 MF22 Poa \u00E2\u0080\u00A2palustris Unknown OTHER TAXA: Dryopteris austriaoa Equisetum spp. No. species recorded No. quadrats 6/12 56 50 37 50 35 50 *Species w i t h at l e a s t one case of canopy-cover > 5%. 503 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 MAJOR GROUP P l a n t species S i t e number SRI SR2 SR3 SR4 SR5 SR6 EVERGREEN TREES & SHRUBS: Abies lasiooarpa t/2 Pioea glauoa 4/8 Pinus contorta DECIDUOUS TREES & SHRUBS: Acer glabrum Alnus spp. Amelanchier alnifolia Betula papyrifera Lonioera involucrata Populus tremuloides Ribes spp. Rosa spp. Rubus idaeus R. parviflorus Salix spp. Shepherdia canadensis Spiraea lucida S. douglasii Vaooinium oaespitoswn V. membranaoeum t/6 V. 'myrtillus Vaooinium spp. Viburnum edule 3/8 FORBS & DWARF SHRUBS: Achillea millefolium Aralia nudicaulis t/2 1/4 1/8 4/12 t/2 16/50 1/2 8/22 1/2 3/5 t/2 2/22 t/2 11/45 16/20 1/3 2/13 1/3 9/43 1/3 10/46 7/32 3/20 12/66 26/90 12/56 8/26 t/2 t/4 9/30 1/4 1/6 4/18 2/8 8/36 2/4 2/12 3/16 14/32 1/2 2/4 3/6 1/2 7/22 4/26 t/2 t/2 5/24 1/12 3/22 1/12 13/26 6/16 1/4 3/22 t/4 11/22 3/28 13/54 9/38 1/4 7/52 2/13 16/66 5/26 504 Table F-\u00E2\u0080\u00A25, Continued MAJOR GROUP S i t e number P l a n t species SR1 SR2 SR3 SR4 SR5 SR6 Arnica spp. > t/5 Aster spp. 2/8 Chimaphila umbeltata t/2 4/32 Clintonia uniflora 1/8 10/70 1/14 2/28 5/38 Cornus canadensis 9/78 21/100 14/80 4/40 10/72 11/76 Disporum oreganum 2/5 Epilobium angustifolium 1/6 t/5 2/6 t/2 1/6 t/2 Galium spp. 2/10 t/6 t/2 1/16 Geranium bicknellii Hypochaeris radicata Linnaea borealis 6/32 13/47 11/76 3/18 2/18 14/48 Mitella nuda 1/6 1/7 5/18 2/20 Petasites frigidus 1/16 5/40 3/18 Pyrola spp. t/2 1/6 2/4 Rubus chamaemorus 6/40 4/24 19/88 R. pedatus 1/6 Smilacina raoemosa 3/12 Streptopus amplexifolius 5/32 1/6 6/40 2/6 Taraxacum spp. Tiarella unifoliata 1/10 t/2 Trifolium repens Vicia spp. GRAMINOIDS: Agropyron repens Agrostis spp. Calamagrostis spp. Carex spp. t/2 Cinna latifolia Festuoa spp. 4/12 2/10 t/2 3/14 Oryzopsis asperifolia 1/8 t/5 4/18 1/8 505 Table F-5, Continued MAJOR GROUP S i t e number P l a n t species SRI SR2 SR3 SR4 SR5 SR6 Poa palustris Unknown OTHER TAXA: Dryopteris austriaoa Equisetum spp. No. species recorded No. quadrats t/2 1/6 27 50 6/2 35 50 23 50 3/10 4/22 37 50 16/58 2/6 28 50 27 50 *Species w i t h at l e a s t one case of canopy-cover > 5%. 506 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 MAJOR GROUP P l a n t species S i t e number SR7 SR8 SR9 SR10 SR11 SR12 EVERGREEN TREES & SHRUBS: Abies lasiocarpa Picea glauca Pinus contorta DECIDUOUS TREES & SHRUBS: Acer glabrum Alnus spp. Amelanchier alnifolia Be tula papyrifera Loniaera involucrata Populus tremuloides Ribes spp. Rosa spp. Rubus idaeus R. parviflorus Salix spp. Shepherdia canadensis Spiraea lucida S. douglasii Vaccinium aaespitosum V. membranaceum V. myrtillus Vaccinium spp. Viburnum edule FORBS & DWARF SHRUBS: Achillea millefolium Aralia nudicaulis 7/16 9/16 11/26 5/16 13/36 6/16 3/10 1/8 1/4 9/48 1/4 1/4 2/12 t/2 7/32 t/2 2/12 5/24 3/14 2/8 t/2 10/26 7/38 10/38 7/32 t/2 t/4 1/2 3/14 1/4 11/28 5/14 2/14 3/22 t/8 2/8 8/40 1/8 6/36 19/62 3/16 5/30 1/10 1/6 8/46 8/42 t/6 2 5/12 1/4 2/14 1/8 4/18 6/34 13/46 5/28 1/6 3/8 5/12 1/2 8/24 2/6 4/24 3/12 8/36 1/4 1/4 1/4 3/16 507 Table F-6, Continued MAJOR GROUP S i t e number P l a n t species SR7 SR8 SR9 SR10 SR11 SR12 Arnica spp.. 9/50 2/18 1/10 t/2 t/6 Aster spp. 6/28 t/4 2/12 Chimaphila umbellata Clintonia uniflora 7/42 8/62 2/16 1/8 6/50 Cornus canadensis 12/62 19/88 9/58 12/58 8/48 5/45 Disporum oreganum Epilobium angustifolium 2/18 1/4 t/4 Galium spp. 4/24 1/12- 3/22 t/4 2/16 Geranium bicknellii Hypchaeris radicata Linnaea borealis 2/12 3/12 2/18 6/22 1/10 2/20 Mitella nuda 6/44 t/4 5/30 6/30 2/20 Petasites frigidus 6/46 6/48 t/4 3/18 1/10 3/32 Pyrola spp. 2/16 2/12 t/2 1/14 Rubus chamaemorus 7/42 3/22 3/6 2/16 R. pedatus 5/22 5/20 3/16 2/10 Smilacina racemosa Streptopus amplexifolius 6/24 1/4 1/6 10/30 7/36 3/14 Taraxacum spp. Tiarella unifoliata t/2 15/56 1/8 1/6 Trifolium repens Vicia spp. GRAMINOIDS: Agropyron repens 1/6 Agrostis spp. t/2 Calamagrostis spp. Carex spp. 1/4 Cinna latifolia Festuca spp. 5/24 2/10 3/16 t/4 Oryzopsis asperifolia 12/42 6/28 508 Table F-6, Continued MAJOR GROUP P l a n t species S i t e number SR7 SR8 SR9 SR10 SR11 SR12 Poa palustris Unknown OTHER TAXA: Dryopteris austviaca 17/78 Equisetum spp. 8/36 No. species recorded 33 No. quadrats 50 30 50 7/34 6/34 35 50 12/52 11/46 35 50 2/18 27 50 3/28 9/52 37 50 *Species w i t h a t l e a s t one case of canopy-cover > 5%. 509 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 MAJOR GROUP P l a n t species S i t e number SRI 3 SR14 SR15 SR16 SR17 SR18 EVERGREEN TREES & SHRUBS: Abies lasiooarpa Picea glauoa Pinus oontorta DECIDUOUS TREES & SHRUBS: Acer glabrum Alnus spp. Amelanchier alnifolia Betula papyrifera Loniceva involucrata , Populus tremuloides Ribes\u00E2\u0080\u00A2 spp. Rosa spp. Rubus idaeus R. parviflorus Salix spp. Shepherdia canadensis Spiraea lucida S. douglasii Vaooinium oaespitosum V. membranaoeum V. myrtillus Vaooinium spp. Viburnum edule FORBS & DWARF SHRUBS: Achillea millefolium Aralia nudicaulis 15/46 22/50 1/12 5/16 15/36 t/2 1/10 1/2 2/8 11/38 3/18 1/8 13/52 5/18 t/2 t/2 3/10 1/8 1/4 1/6 1/8 4/12 4/8 3/12 21/40 1/4 3/26 1/16 3/16 t/2 1/2 5/16 7/30 14/42 17/68 18/52 t/4 2/4 t/4 2/12 1/10 1/4 1/6 5/28 t/2 1/4 1/4 t/2 1/4 2/18 8/24 2/2 10/36 5/18 12/66 13/54 3/20 11/42 23/80 12/32 2/16 4/28 12/60 13/56 4/28 1/10 510 Table F-7, Continued MAJOR GROUP P l a n t species S i t e number SR13 SR14 SR15 SR16 SR17 SR18 Arnica spp. t/6 t/6 2/12 Aster spp. t/2 1/6 2/14. 1/10 Chimaphila umbellata 1/4 7/32 1/12 Clintonia uniflora 1/8 3/42 7/54 11/60 2/22 2/24 Cornus canadensis 10/62 20/88 11/66 16/58 9/68 24/90 Disporum oreganum 11/52 4/20 Epilobium angustifolium t/4 1/6 Galium spp. 1/8 2/18 Geranium bicknellii Rypochaeris radicata Linnaea borealis 6/52 15/46 11/44 t/2 3/24 11/40 Mitella nuda 2/14 t/8 Petasites frigidus 1/8 8/48 Pyrola spp. t/2 2/20 2/16 2/20 t/2 Rubus chamaemorus R. pedatus 2/8 2/12 t/2 Smilacina racemosa 1/6 1/4 Streptopus amplexifolius 5/30 1/4 13/60 8/22 Taraxacum spp. Tiarella unifoliata 2/14 1/2 1/4 2/8 Trifolium repens Vicia spp. GRAMINOIDS: Agropyron repens Agrostis spp. Calamagrostis spp. 3/6 1/6 Carex spp. Cinna latifolia Festuca spp. 4/10 Oryzopsis asperifolia 1/4 6/16 5/18 Table F-7, Continued MAJOR GROUP S i t e number P l a n t species SR13 SR14 SR15 SR16 SR17 SR18 Poa palustris Unknown OTHER TAXA: Dryopteris austriaca 8/38 2/10 1/8 21/62 t/2 3/14 Equisetum spp. 4/28 t/2 4/20 No. species recorded 30 27 30 33 23 33 No. quadrats 50 50 50 50 50 50 *Species w i t h at l e a s t one case of canopy-cover > 5%. 512 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 MAJOR GROUP P l a n t species S i t e number SR19 SR20 SR21 SR22 SR23 EVERGREEN TREES & SHRUBS: Abies lasiooarpa Pioea glauoa Pinus contort a DECIDUOUS TREES & SHRUBS: Acer glabrum Alnus spp. Amelanohier alnifolia Betula papyrifera Lonioera involucrata Populus tremuloides Ribes spp. Rosa spp. Rubus idaeus R. parviflorus Salix spp. Shepherdia' canadensis Spiraea lucida S. douglasii Vaooinium oaespitosum V. membranaceum V. myrtillus Vaooinium spp. Viburnum edule FORBS & DWARF SHRUBS: Achillea millefolium Aralia nudioaulis 10/34 11/16 t/4 22/54 25/55 3/6 1/4 t/2 5/18 6/38 t/2 2/16 2/4 1/4 3/14 t/2 t/2 1/4 1/2 9/42 6/28 1/8 3/14 7/36 t/2 10/36 14/42 1/8 1/6 2/6 2/12 t/2 5/18 4/24 7/38 16/60 1/6 4/28 14/46 9/34 3/14 t/4 2/14 t/2 1/i 4/14 13/64 1/6 14/56 513 Table F-8, Continued MAJOR GROUP P l a n t species SR19 S i t e number SR20 SR21 SR22 SR23 Arnica spp. Aster spp. Chimaphila umbellata Clintonia uniflora Cornus canadensis Disporum oreganum Epilobium angustifolium Galium spp. Geranium bicknellii Rypochaeris radicata Linnaea borealis Mltella nuda Petasites frigidus Pyrola spp. Rubus chamaemorus R. pedatus Smilacina racemosa Streptopus amplexifolius Taraxacum spp. Tiarella unifoliata Trifolium repens Vicia spp. GRAMINOIDS: Agropyron repens Agrostis spp. Calamagrostis spp. Carex spp. Cinna latifolia Festuca spp. Oryzopsis asperifolia 3/12 4/24 11/64 t/2 1/6 3/22 3/26 1/12 6/50 6/32 4/26 t/8 7/24 t/2 5/50 22/88 5/32 2/14 t/2 14/34 1/6 1/4 t/2 1/4 10/46 1/4 13/74 1/12 5/32 5/28 1/6 8/60 9/40 3/10 2/10 4/28 11/66 2/8 2/12 2/22 7/44 3/12 t/2 4/28 3/18 8/56 8/72 1/6 2/26 t/4 2/16 2/18 1/8 3/24 7/38 4/28 2/6 3/8 10/16 6/10 514 Table F-8, Continued MAJOR GROUP S i t e number P l a n t species SR19 SR20 SR21 SR22 SR23 Poa palustris Unknown 33/60 11/44 10/50 5/22 5/24 26 29 27 50 50 50 OTHER TAXA: Dryopteris austriaoa 5/32 2/18 Equisetum spp. ' t/4 No. species recorded 32 36 No. quadrats 50 50 *Species w i t h at l e a s t one case of canopy-cover > 5%. 515 Table F-9 2 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 Age of s e r a i stage (yr.) Species 1 5 10 25 45 75 110 135 150 200 Abies lasiooarpa 111 31 199 305 Aoer glabrum 21 t 3 Alnus spp. 1 4 38 Amelanchier alnifolia 1 13 8 Betula papyrifera 14 672 Cornus stolonifera 3 1 Lonioera involucrata 13 2 13 5 Picea glauoa 8 388 71 13 Pinus oontorta ( 5 90 Populus tremuloides 1 88 t P. balsamifera Pseudotsuga menziesii 3 1 Ribes spp. 23 29 7 15 12 6 Rosa spp. 1 11 2 2 5 t 1 1 1 Rubus idaeus 5 t R. parviflorus t 1 1 t 2 Salix spp. 2 48 2466 Sambucus racemosa 6 Shepherdia canadensis 20 Sorbus spp. 7 2 t 1 Spiraea lucida 6 t t t t 1 t S. douglasii 20 24 10 11 Vaooinium spp. 2 1 10 2 t Viburnum edule 7 2 t 2 T o t a l phytomass 13 120 160 3616 102 77 194 56 226 349 No. s i t e s 3 2 1 1 3 1 4 2 1 3 516 Table F-10 2 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 Age of s e r a i stage (yr) Species 1 5 10 25 45 110 150 Abies lasiooarpa 17 16 5 Acer glabrum Alnus spp. Amelanohier alnifolia 10 2 11 Betula papyrifera 34 14 Cornus stolonifera 1 1 Lonioera involucrata 1 9 5 2 2 5 Picea glauoa 28 1 Pinus contorta 17 Populus tremuloides 23 28 81 211 P. balsamifera 3 Pseudotsuga menziesii Ribes spp. 17 9 Rosa spp. 5 1 1 1 2 Rubus idaeus R. parviflorus Salix spp. 2 165 2215 77 t Sambucus racemosa Shepherdia canadensis Sorbus spp. 3 Spiraea luoida t 2 8 S. douglasii 32 18 45 Vaooinium spp. t Viburnum edule 5 5 2 T o t a l phytomass 24 88 348 2461 83 41 92 No. s i t e s 1 2 2 1 1 2 3 517 Table F - l l Phytomass 2 (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 Substrat es T i l l L a c u s t r i n e Species 135 150 200 150 Abies lasiocarpa 196 268 90 Acer glabrum 10 32 1 Alnus spp. Amelanchier alnifolia Be tula papyrifera t 2 3 Cornus stolonifera Lonicera involucrata 1 13 1 Picea glauca 2 Pinus contorta Populus tremuloides 7 10 88 P. balsamifera t Pseudotsuga menziesii Ribes spp. 3 1 Rosa spp. 1 9 Rubus idaeus R. parviflorus t 1 Salix spp. Sambucus racemosa Shepherdia canadensis Sorbus spp. 1 3 1 Spiraea lucida 1 1 5 S. douglasii 16 Vaccinium spp. 11 8 4 Viburnum edule 2 8 16 2 T o t a l phytomass 46 276 302 197 No. s i t e s 1 2 1 3 Table F-12 2 .2 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 Species Age of s e r a i stage (yr) 10 25 45 75 110 135 150 200 Abies lasiooarpa Acer glabrum Alnus spp. Amelanchier alnifolia Betula papyrifera Cornus stolonifera Lonioera involucrata Picea glauoa Pinus oontorta Populus tremuloides P. balsamifera Pseudotsuga menziesii Ribes spp. Rosa spp. Rubus idaeus R. parviflorus Salix spp. Sambuous racemosa 0.03 0.63 0.02 0.18 0.25 0.06 0.22 5.11 0.10 3.75 0.75 0.09 1.34 0.97 0.93 0.34 1.79 2.62 0.01 0.03 0.07 0.11 0.51 0.11 0.03 0.03 0.21 0.09 0.14 0.001 0.01 0.02 0.002 0.01 0.04 0.01 0.20 0.03 0.04 0.09 0.01 0.02 0.13 0.01 0.007 0.03 0.92 17.01 0.01 0.01 0.03 0.01 0.02 0.02 0.03 0.02 0.01 0.08 Table F-12, Continued Age of s e r a i stage (yr) Species 1 5 10 25 45 75 110 135 150 200 Shepherdia canadensis 0. 21 Sorbus spp. 0. .17 0. 05 0. .01 0. 02 Spiraea lucida 0. .01 0.01 0. ,002 0.01 0. 01 0. ,03 0. 01 S. douglasii 0. .03 0.05 0. 02 0. ,02 Vaccinium spp. 0. .05 0. 02 0. ,19 0. ,03 0. 002 Viburnum edule 0. ,18 0. 06 0. ,01 0. 04 T o t a l b a s a l area 0.19 2. ,13 2.59 29.89 0. .97 0.54 1. 92 0. 62 1. 92 3. 05 No. s i t e s 3 ; I 1, i : 3 1 4 2 1 3 i h-1 520 Table F-13 2 2 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 Age of s e r a i stage (yr) Species 1 5 10 25 45 110 150 Abies lasiooarpa .25 .22 .06 Acer glabrum Alnus spp. Amelanchier alnifolia .15 .25 .16 Betula papyrifera .63 .26 Cornus stolonifera .03 .01 Lonioera involucrata .01 .15 .01 .04 .05 .08 Picea glauoa .36 .02 Pinus contort a .23 Populus tremuloides .55 .52 1.11 2.47 P. balsamifera .06 Pseudotsuga menziesii Ribes spp. .03 .01 Rosa spp. .01 .03 .03 .06 Rubus idaeus R. parviflorus \u00E2\u0080\u00A2 .01 Salix spp. .05 3.14 7.29 .65 .01 Sambucus racemosa Shepherdia canadensis Sorbus spp. .05 .004 .07 Spiraea lucida .01 .01 .03 .21 S. douglasii .05 .01 .08 Vaooinium spp. Viburnum edule .01 .01 .06 T o t a l B a s a l area .57 1.07 5.83 16.82 .78 .40 .82 No. s i t e s 1 2 2 1 1 2 3 521 Table F-14 2 2 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 T i l l L a c u s t r i n e Species 135 150 200 150 Abies lasiocarpa 1. 73 2.28 0. 76 Acer glabrum 0.31 0.97 0.04 Alnus spp. Amelanchier alnifolia Be tula papyrifera 0.003 0.04 0.06 Cornus stolonifera Loniaera involucrata 0.02 0.22 0.02 Picea glauca 0.03 Pinus contorta Populus tremuloides 0.13 0.19 0.96 P. balsamifera 0.003 Pseudotsuga menziesii Ribes spp. 0.01 Rosa spp. 0.02 0.13 0.03 0.001 Rubus idaeus R. parviflorus 0.01 0.03 Salix spp. Sambucus racemosa Shepherdia canadensis Sorbus spp. 0.04 0.08 0.03 Spiraea lucida 0.01 0.01 0.12 S. douglasii 0.02 Vaccinium spp. 0.22 0.16 0.07 Viburnum edule 0.06 0.15 0.38 0.06 T o t a l Basal area No. s i t e s 0.69 , 1 3.45 2 3.02 1 2.15 3 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 Species Age of s e r a i stage (yr) 10 25 45 75 110 135 150 200 Abies lasiooarpa Acer glabrum Alnus spp. Amelanchier alnifolia Betula papyrifera Cornus stolonifera Lonioera involucrata Picea glauoa Pinus oontorta Populus tremuloides P. balsamifera Pseudotsuga menziesii Ribes spp. Rosa spp. Rubus idaeus R. parviflorus Salix spp. Sambucus racemosa 129 60* 79+ 70 104 103 115* 125 70+ 85+ 54 45* 71(4) 43* 181 107 68 144* 189 374(4) 67(4) 192(4) 175* 467 39 66 63 121+ 469(4) 62 43 103(9) 111* 239* 91* 71+ 77* 69(9) 115* 72 49 49 51* 53+ 82+ 70 67 51(4) 49\" 121+ 45* 57 34 160+ 89* 44 56+ 41(4) On r-o Table F-15, Continued Age of s e r a i stage (yr) Species 1 5 10 25 45 75 110 135 150 200 Shepherdia canadensis 152+ Sorbus spp. 98 72+ 69 83* Spiraea lucida 64 45 49* 45* 41(6) 59(5) 49* S. douglasii 75* 103 69 60 Vaooinium spp. 59 47* 54(6) 76 32 Viburnum edule 78+ 75(6) 73 75* Mean height 73 85 101 361 78 77 84 62 90 86 No. s t a t i o n s 5 10 4 5 10 4 19 9 5 12 *n = 2. +n = 3. 524 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 Age of s e r a i stage (yr) Species 1 5 10 25 45 110 150 Abies lasiocarpa 59+ 63+ 64 Acer glabrum Alnus spp. Amelanchier alnifolia 86(5) 83* 95(4) Be tula papyrifera 138+ 173+ Cornus stolonifera 85 59* 64 Loniaera involucrata 70(4) 67* 62+ 63(5) 69(4) Piaea glauca 69(5) 82 Pinus contorta 224 Populus tremuloides 91(4) 117(4) 189(5) 325* P. balsamifera 99 Pseudotsuga menziesii Ribes spp. 51 58 Rosa spp. 68(7) 74 51* 48(4) 52(6) Rubus idaeus R. parviflorus 32 Salix spp. 98 112(5) 252(5) 335 112 Sambucus racemosa Shepherdia canadensis Sorbus spp. 83(4) Spiraea lucida 62 61* 44+ 72(11) S. douglasii 67+ 65+ 73+ Vaccinium spp. \u00E2\u0080\u00A2 43 Viburnum edule - 65 66 70(4) Mean height 89 82 118 252 93 57 76 No. s t a t i o n s 4 10 9 5 5 9 15 *n = 2. +n = 3. 525 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 T i l l L a c u s t r i n e Species 135 150 200 150 Abies lasiooarpa 206+ 153* 180+ Acer glabrum 95* 102(5) 84 Alnus spp. > Amelanchier alnifolia Betula papyrifera 54 124 160 Cornus stolonifera Lonioera involucrata 76 76 65 Pioea glauoa Pinus contorta Populus tremuloides 188* 148 399* P. balsamifera 51 Pseudotsuga menziesii > Ribes spp. 40 Rosa spp. 79 90+ 75* 36 Rubus idaeus R. parviflorus 30 48* Salix spp. Sambucus racemosa Shepherdia canadensis Sorbus spp. 95 102 103 Spiraea luoida 54+ 53* 102+ S. douglasii 55 Vaooinium spp. 53* 56+ 51(5) Viburnum edule 70* 85* 81* 77(4) Mean height 65 91 118 104 No. s t a t i o n s 4 10 5 13 *n = 2. +n = 3. 526 Table F-18 Number of Stems Samples i n the Shrub Layer, by Species, i n a Mesic Sub-Boreal Forest Sere on T i l l Substrates Age of s e r a i stage (yr) Species 1 5 10 25 45 75 110 135 150 200 Abies lasiooarpa 32 7 6 13 Acer glabrum 22 1 26 Alnus spp. 46 2 11 Amelanchier alnifolia 15 2 8 3 1 Betula papyrifera 17 19 Cornus stolonifera 3 8 Lonioera involucrata 27 7 7 10 40 Picea glauoa 6 9 13 3 Pinus contort a 3 11 Populus tremuloides 2 1 1 1 P. balsamifera 1 Psuedotsuga menziesii Ribes spp. 10 3 3 1 2 4 Rosa spp. 16 41 8 5 24 1 8 3 14 Rubus idaeus 64 7 R. parviflorus 2 10 4 2 25 Salix spp. 1 1 33 32 Sambucus racemosa 6 Shepherdia canadensis - 11 Sorbus spp. 9 12 1 2 Spiraea lucida 4 45 2 2 2 17 7 4 S. douglasii 18 3 9 3 Vaooinium spp. 5 7 24 1 2 Viburnum edule 22 20 1 T o t a l no. stems 154 215 68 74 68 27 184 45 18 108 S i t e s : MF 2,5, 8, 22 21 4 1,9 7 6 10 SR 1,3 4 9,11 22, 23 5 6, 10 527 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 Age of s e r a i stage (yr) Species 1 5 10 25 45 110 150 Abies lasiocarpa 15 21 4 Acev glabrum Alnus spp. Amelanchier alnifolia 26 4 24 Betula papyrifera 30 3 Cornus stolonifera 5 3 Lonicera involucrata 2 21 31 3 4 19 12 Picea glauca 21 1 Pinus contorta 1 Populus tremuloides 46 29 34 4 P. balsamifera 1 Pseudotsuga menziesii Ribes spp. 2 2 Rosa spp. 23 5 5 9 31 Rubus idaeus R. parviflorus 1 Salix spp. 5 223 68 2 1 Sambucus racemosa Shepherdia canadensis Sorbus spp. 18 Spiraea lucida 3 2 5 50 S. douglasii 24 9 32 Vaccinium spp. 2 Viburnum edule 8 1 2 14 T o t a l no. stems 48 136 368 82 14 71 182 S i t e s : MF 12 15,17 13,18 20 14 16 SR 12 18 8,21 528 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 T i l l L a c u s t r i n e Species 135 150 200 150 Abies lasiooarpa 2 9 4 Acer glabrum 7 33 Alnus spp. 14 Amelanchier alnifolia Betula papyrifera 1 8 Cornus stolonifera Lonioera involucrata 1 21 Picea glauoa 1 2 Pinus contorta Populus tremuloides 2 4 5 P. balsamifera 1 Pseudotsuga menziesii Ribes spp. 1 Rosa spp. 1 16 2 1 Rubus idaeus R. parviflorus 2 2 Salix spp. Sambucus racemosa Shepherdia canadensis Sorbus spp. 2 2 2 Spiraea lucida 5 3 14 S. douglasii 5 Vaooinium spp. 16 20 12 Viburnum edule 5 14 19 13 T o t a l no. stems 37 117 63 58 S i t e s : MF 3 11 SR 2 13 14,17,20 529 Table F-21 Oven-Dried Weights of Components of Major Shrub Species i n Sub-Boreal Forests Species Sample s i t e type ] Oven-dried wts (g) leaves current twigs o l d twigs t o t a l Abies lasiocarpa G2 3 0.9 0.9 1.8 G4 2 2.0 1.0 1.0 4.0 M4 1 0.9 0.8 0.8 2.5 2 1.5 0.5 0.5 2.5 Acer glabrum Alnus spp. M4 18.0 2.0 162.0 182.0 Amelanchier alnifolia Mahonia nervosa Gl 3 0.8 0.6 25.0 26.4 - 3 0.7 0.4 12.0 13.1 3 7.0 0.5 39.0 46.5* 3 4.0 0.6 96.0 100.6* 3 0.5 0.5 25.0 25.9* G2 3 81.4 12.2 309.3 407.0 G3 1 3.0 0. 7 13.0 16.7 1 0.2 0.1 5.0 5.3 1 0.3 0.1 0.6 1.0 1 11.0 3.0 41.1 55.1 Ml 1 48.0 7.0 58.0 113.0 1 29.0 1.0 35.0 55.0 M3 1 30.0 2.0 28.0 60.0 Ml 2 8.0 4.0 0.0 12.0 M3 1 52.0 14.0 26.0 82.0 M4 1 4.0 0.9 21.0 25.9 2 14.0 2.0 134.0 150.0 G l 3 28.0 0.5 77.0 105.5 3 32.0 0.5 98.0 130.5 3 18.0 2.0 45.0 65.0 G4 1 17.0 4.0 37.0 58.0 1 28.0 6.5 143.0 177.5 1 4.0 2.0 6.0 12.0 1 6.0 2.0 22.0 30.0 1 1.0 0.9 2.0 12.0 1 2.5 0.9 1.0 4.4 2 38.0 22.0 102.0 162.0 3 22.5 4.9 27.4 120.4 3 9.1 1.6 10. 7 87.7 3 38.0 6.1 44.1 196.1 3 \u00E2\u0080\u00A222.0 5.0 27.0 124.0 M3 1 2.0 0.3 0.1 2.4 530 Table F-21, Continued Oven-dried wts (g) Species Sample s i t e type 1 leaves current twigs o l d twigs t o t a l Rosa spp. Rubus idaeus Rubus parviflorus Salix spp. M3 2 39.0 12.0 15.0 66.0 1 4.0 0.4 0.8 5.2 1 6.0 m 3.0 9.0** M4 1 4.0 1.0 4.0 9.0 1 1.0 0.2 0.7 1.9 1 2.0 1.0 6.0 9.0 1 2.5 1.0 2.5 6.0 1 4.0 0.9 7.0 11.9 1 m 0.5 14.0 m Ml . 2 8.0 m 2.0 10.0** G2 3 1.7 0.8 0.8 3.3 G3 3 1.0 m 0.3 1. 3** 1 0.2 0.1 0.1 0.4 0.1 m 0.1 0.2** 2 16.5 1.5 9.0 27.0 G4 2 7.0 1.4 2.5 10.9 1 1.0 1.0 0.8 2. 8 3 7.0 0.9 2.0 9.9 Ml 2 5.0 m 1.0 6.0** 1 1.0 0.7 0.7 2.4 M3 1 2.0 m 1.0 3.0** M4 1 9.5 m 6.0 15.5** 2 15.0 m 6.0 21.0** Gl 3 19.0 0.6 66.0 85.6 3 15.0 0.6 46.0 61.6 3 43.0 4.0 132.0 179.0 3 27.0 0.4 87.0 114.4 3 13.0 0.3 m m G4 1 85.0 11.5 269.0 365.5 1 118.5 31.0 329.0 478.5 1 65.0 21.0 222.0 308.0 1 95.0 21.0 260.0 376.0 1 37.5 8.0 114.0 159.5 1 . 71.0 23.0 197.0 291.0 1 72.5 11.5 199.0 283.0 4 24.0 137.0 514.0 675.0 4 50.0 5.0 172.0 227.0 4 36.0 7.0 119.0 162.0 4 18.0 3.0 57.5 78.5 4 9.0 3.3 30.0 42.3 G4 2 42.0 26.5 131.0 199.5 531 Table F-21, Continued Oven-dried wts (g) S ample current o l d Species s i t e t y p e 1 leaves twigs twigs t o t a l Shepherdia canadensis M2 1 1 M4 1 Spiraea lucida G3 1 1 2 1 1 1 G4 2 Ml 1 1 1 1 1 1 1 1 1 1 M2 1 1 1 1 1 1 1 1 M3 1 1 2 M4 1 1 1 1 1 1 1 1 1 30.5 10.9 210.0 251.4 38.5 20.5 250.5 309.5 27.0 10.0 49.5 86.5 2.0 0.2 2.0 4.2 3.0 0.4 1.0 4.4 24.0 10.0 26.0 60.0 5.0 0.1 1.0 \u00E2\u0080\u00A2 6.1 0.5 0.2 0.5 1.2 48.0 0.0 82.0 130.0 0.7 0.8 0.7 2.2 115.0 2.0 46.0 163.0 135.0 3.0 59.0 197.0 26.0 2.0 10.0 38.0 13.0 0.9 1.0 14.9 7.0 0.9 3.0 10.9 15.0 1.0 5.0 21.0 25.0 m 8.0 33.0** 1.0 m 0.9 1. 9** 24.0 m 10,0 34.0** 4.0 m 1.5 6.3** 2.0 0.5 3.0 5.5 0.2 0.1 0.3 0.6 3.0 0.5 16.0 19.5 2.0 0.4 8.0 10.4 0.9 0.4 0.5 1.8 5.0 0.4 9.0 14.4 0.6 0.4 2.0 3.0 3.0 0.5 9.5 13.0 1.0 0.1 0.2 1.3 1.0 m 0.6 1.6** 53.0 m 19.0 72.0** 4.5 1.4 13.0 18.9 2.0 1.0 2.0 5.0 0.9 0.9 0.8 2.6 2.0 0.8 2.0 4.8 3.5 1.0 6.0 10.5 3.0 0.7 2.0 5.7 2.0 0.3 0.6 2.9 3.0 1.0 4.0 8.0 4.0 0.8 7.0 11.8 532 Table F-21, Continued Oven-dried wts (g) Sample current o l d Species s i t e t y p e 1 leaves twigs twigs t o t a l Vaooinium spp. G2 G3 Ml M2 M3 M4 Viburnum edule G3 3 0.9 0.8 0.9 2.6 2 2.0 m 0.3 2.3** 1 0.8 m 0.1 0. 2** 1 0.4 m 0.1 0.5** 1 1.0 m 1.0 2. 0** 1 1.0 0. 7 0.8 2.5 1 1.0 m 0.9 1.9** 1 1.0 1.0 1.0 3.0 1 1.0 0.9 0.9 2.8 1 2.0 1.0 1.5 4.5 1 2.0 0.9 1.3 3.2 1 1.2 0.9 1.1 3.2 1 7.0 0.2 47.0 56.0 1 2.0 0.2 12.0 14.2 1 4.0 0.8 27.0 31.8 1 3.0 0.4 2.0 5.4 1 5.0 0.3 39.0 44.3 1 6.0 0.4 50.0 56.4 1 4.5 0.9 31.5 36.9 1 6.0 0.9 2.6 9.5 1 5.0 1.0 27.0 33.0 1 5.5 0.5 0.0 6.0 1 0.6 0.3 0.0 0.9 1 3.0 1.0 0.9 4.9 1 0.4 0.4 0.0 0.8 1 7.0 3.0 25.0 35.0 1 8.0 2.0 35.0 45.0 1 6.0 1.5 46.0 53.5 1 4.0 2.0 40.5 46.5 1 6.0 2.0 50.0 58.0 1 6.0 2.0 28.0 36.0 1 7.0 3.0 35.0 45.0 1 0.5 0.9 1.0 2.4 1 5.0 2.0 47.0 54.0 1 7.0 5.0 41.0 53.0 1 0.4 0.1 2.0 2.5 1 0.3 0.1 4.0 4.4 1 0.4 0.1 0.9 1.4 1 0.1 0.0 0.7 0.8* 2 4.0 1.5 8.0 13.5 *These samples a l s o had ca t k i n s whose dry weights were 0.6, 13.0 and Table F-21, Continued 0.5 g, r e s p e c t i v e l y . **Twigs not separated i n t o current and o l d e r growth. Sample type codes: 1 = p l a n t s from 0.5- x 2.0- m, 2 = p l a n t s from balance of 5- x 5- m perquadrat not i n quadrats, 3 = s i n g l e p l a n t , 4 = p l a n t s from a l - x 1- m p l o t . 2 For separations of evergreens, leaves i n c l u d e d annual plus o l d e r needles. For some samples current and o l d twigs were not separated (shown by * * ) . APPENDIX G DATA FOR NUTRIENT CONTENTS OF SAMPLED PLANT SPECIES (SECTION 8) 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 Abies lasiooarpa Amelandhier alnifolia Month Sample* E 3 \" E 4 G 3 G 4 S 5 E l E 2 Apr t 5.5 5.4 May t 1 c 6.3 7.1 6.5 6.7 5.7 Jun t 1 c 14.1 10.1 12.2 10.3 11.1 J u l t 1 Aug t 1 8.4 7.8 7.5 8.7 6.5 Sep t 5.5 6.4 7.7** 5.6 4.3 1 7.1 8.9 8.2 8.7 6.3 c 8.0 7.3 7.7 5.9 Oct 6.3** 7.0 5.7 Nov c 7.5** Dec c 6.3 7.1 6.6 7.1 5.5 Jan c 6.5 6.8 6.8 6.5 6.4 Feb c 6.3 6.4 6.6 6.7 6.4 Mar c 5.7 6.9 6.5** 6.5 6.2 Apr c 5.9 6.0 5.7 6.4 5.8 * t - twig, 1 - l e a f , c - combined twig and l e a f . **Duplicate sample. 536 Table G - l , Continued Betuta papyrifera Month Sample* E 1 E 2 G 1 G 2 S' 2 Apr t 7.1 7.0 May t 1 c 5.6 25.4 5.2 23.0 Jun t 1 c 14.4 10.4** 15.2 9.6 17.8 J u l t 1 c 9.0 14.6 Aug t 1 c 4.0 14.3** 4.0 13.2 4.6 14.6 4.4 Sep t 1 c 5.2 10.0 4.0** 10.2 4.8 11.0 5.0 10.1** Oct t 5.4 5.2 Nov t 5.0 5.4 Dec t 4.8 5.0 6.2 5.2 Jan t 5.4 5.2 5.6 6.0** Feb t 5.0 5.2 5.4 5.4 Mar t 5.2 5.2 5.4 Apr t 6.0 4.6 5.2 5.6 * t - twig, 1 - l e a f , c - combined twig and l e a f . \"'Duplicate sample. 537 Table G - l , Continued Cornus stolonifera Lobaria vulmonaria Month Sample* E 1 E 2 G 4 S 1 E 3 G 4 Apr t 4.9 4.5 5.0 14.8 * May t 1 c 16.0 Jun t 1 c 12.5 14.0 9.8 16.9 13.8 14.7 10.2 J u l t 1 c Aug t 1 c 3.6 10.7 7.6 3. 7 10.4 7.5 4.2 12.5 8.9 4. 5** 13.7 7.5 Sep t 1 c 4.5 5.6 5.1 4.7 7.0 5.9 3.7 11.0 7.7 4.8 9.8 7.6 11.4 Oct t 5.1 5.3 5.5 Nov t 11.4 Dec t 5.5 4.7 4.5 4.7 12.4 Jan t 5.0 4.3 4.4 5.5** 12.4 Feb t 5.4 4.2 4.8 5.2 11.8 Mar t 4.8 4.5** 4.9 4.5 10.0 Apr t 4.2 4.3** 4.9 12.4 * t - twig, 1 - l e a f , c - combined twig and l e a f . **Duplicate sample. 538 Table G - l , Continued Populus tremuloides Populus balsamifera Month Sample* B l E 2 G 2 B l G l G 3 Apr t 7.1 5.6 5.6 9.1 May t 1 c 11.5** 12.6 Jun t 7.0 1 c 14.0 J u l t 8.1 6.6 1 13.4 c 10.9 10.5 Aug t 5.6 5.9 1 13.4 10.3 c 9.7 8.7 9.2** 7.6 Sep t 6.1** 5.5 3.7 4.1 1 10.0 8.9 6.0 4.9 c 8.1 7.7 5.3 4.7 Oct t 5.7** Nov t 7.1** 5.9** 4.7 4.4 Dec t 5.4** 4.4 Jan t 5.7 5.7 4.8 4.7 Feb t 5.5 6.3 4.7 5.0 Mar t 5.7 6.5 5.0 4.7 Apr t 6.0 6.6** 4.4 5.6 * t - twig, 1 - l e a f , c -**Duplicate sample. combined twig and l e a f . 5 3 9 Table G - l , Continued Salix spp. Month Sample* B l B 2 B 3 E - l E 2 G l G 2 Apr May Jun J u l Aug Sep Oct Nov Dec Jan Feb Mar Apr 5.7 5.3 5.3 t 1 c t 1 c t 1 c t 1 c t 1 c t t t t t t t 6.9** 10.1** 13.8 4.6 13.6 9.1 4.6 8.1 6.5 5.9 5.7 4.6 4.7 5.2 10.4 20.5 3.3 12.5 7.9 3.6 6.7 5.4 6.2 5.7 5.1 5.8 4.7 5.6 9. 7** 15.0** 15.1** 12.3** 10.1** 6.0 8.0 7.4 5. 9** 6.1 6.4 6.2 5.2 5.2 14.4 4.0 12.4** 8.7 5.3 7.5 6.4 6.0 5.4 4.5 6.7 4.2 13.0 4.4 9.9 7.5 5.5 9.6** 7.6 5.6 5.2 5.4 5.0 4.1 4.3 12.1 11.1 8.4 4.5 6.9 6.1 5.4 5.4 5.0 4.9 5.6 5.0 6.0 7.6 12.6** 10.1 3.9** 11.4 7.6 5.4 8.7 7.2 5.7 6.0 5.8 4.6 5.3 5.0 * t - twig, 1 - l e a f , c -**Duplicate sample. combined twig and l e a f . 540 Table G - l , Continued Vaooinium Salix spp. Sorbus spp. membranaceum Month Sample* S 2 S 3 S 5 E l E 2 E l Apr t 6.6 5.6 May t 1 c 10.5 6.0 Jun t 1 c 10.1 13.0 J u l t 1 c 7.6 15.5 11.2 6.2 12.1 9.6 13.1 Aug t 1 c 4.2 12.1 8.7 4.1 10.8 8.0 4.2 11.9** 8.6 4.7 9.7 Sep t 1 c 5.0 10.1** 4.1 8.2 5.9 6.1 11.0 8.9 6.2 Oct t 5.2 5.3 6.2 5.1 Nov t Dec t 5.2 5.7 5.5 Jan t 6.0** 4.8 5.1** 5.8 5.8 Feb t 5.4 5.4 5.7** 5.4 5.4 Mar t 5.4 5.0** 4.9 4. 7 5.6** Apr t 5.6 4.6 5. 1** * t - twig, 1 - l e a f , c - combined twig and l e a f . 5.4 11.2 **Duplicate sample. 541 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 P r i n c e George Study Area, A p r i l 19 72 to A p r i l 1973 Abies lasiocarpa Be tula papyrifera Month Sample* E 4 G 4 S 5 B 1 B 3 E 2 Apr. t 8.8 9.7 12.7 May t 1 c 10.3 9.8 8.2 6.9 Jun t 1 c 10.2 10.1 J u l t 1 c 9.2 8.8 10.4 Aug t 1 c 9.0 11.1 9.6 8.9 10.2 10.7 7.4 Sep t 1 c 8.4 11.4 6.9 9.2 11.0 12.0 8.5 10.0 14.3 7.9 Oct t 11.1 8.6 12.2 Nov t 10.8 8.9 13.5 Dec t 9.3 13.4 9.5 Jan t 9.5 9.5 9.4 9.6 9.7 Feb t 11.3 10.8 Mar t 8.4 9.8 9.5 9.1 10.6 Apr t 8.8 4.2 7.0 9.4 11.2 8.7 * t - twig, 1 - l e a f , c - combined twig and l e a f . 0 542 Table G-2, Continued c c Betula papyrifera Cornus stolonifera Month Sample* G l G 2 S 2 \u00E2\u0080\u00A2 E 2 G 4 S I Apr t 7.9 May t 10.6 11.3 1 7.9 9.1 Jun t 10.9 12.1 5.2 1 8.8 6.3 4.0 5.6 9.3 J u l t 9.7 1 5.4 Aug t 11.4 11.1 10.7 6.9 9.3 1 7.3 6.1 4.3 3.0 6.7 c Sep t 10.7 10.8 10.9 11.1 3.3 9.3 1 5.0 7.2 6.1 4.2 4.0 c Oct t 8.9 9.3 11.6 Nov t 10.2 10.9 7.1 Dec t 10.9 11.5 11.2 Jan t 11.0 10.6 9.2 9.0 9.7 ' 10.8 Feb t 8.9 10.7 11.8 Mar t 10.5 10.7 7.8 7.5 12.7 Apr t 8.9 10.7 10.8 8.9 8.0 12.3 * t - twig, 1 - l e a f , c - combined twig and l e a f . 543 Table G-2, Continued Populus tvemuloides Salix spp. Month Sample* E 2 G 2 B l B 3 E 2 Apr t 8.5 9.7 9.8 11.1 May t 9.1 11.3 10.3 1 c Jun t 11.3 1 c 10.1 J u l t 1 8.7 c 7.6 Aug t 11.7 11.9 1 8.0 5.8 6.4 9.0 c Sep t 9.4 10.0 7.7 9.3 1 8.6 6.4 5.9 12.4 c Oct t 8.2 13.6 Nov t 10.4 12.4 10.9 Dec t Jan t 7.9 9.3 10.7 11.7 12.1 Feb t ' 12.2 Mar t 9.0 8.8 11.5 12.2 Apr t 8.1 8.9 11.8 14.1 * t - twig, 1 - l e a f , c - combined twig and l e a f . 544 Table G-2, Continued Salix spp. Sorbus spp. Month Sample* G l G 2 S 3 S 5 E 2 Apr May Jun J u l Aug Sep Oct Nov Dec Jan Feb Mar Apr 8.4 t 1 c t 1 c t 1 c t 1 c t 1 c t t t t t t t 9.5 13.1 11.0 8.6 6.7 5.6 4.3 9.8 6.8 8.4 11.4 5.8 11.4 10.5 11.1 13.1 13.0 5.6 6.3 11.5 4.9 9.4 5.7 10.0 3.9 12. 7 6.7 11.3 11.4 11.8 10.2 12.5 12.3 12.4 11.6 13.1 11.9 4.6 6.4 5.5 10.4 11.1 8.6 * t - twig, 1 - l e a f , c - combined twig and l e a f . APPENDIX H CLIMATIC DATA USED FOR SECTION 546 Table H - l Penetrance Values (1-11 Scale) f o r Snow Hardness Estimates Across the South-Facing Ecotone at the Grove Area, 1973 S i t e Penetrance values (4 r e p l i c a t e s each date) l o c a t i o n S t a t i o n Feb . 8 Mar 6 Apt . 4 76 m i n 1 11. 10, 2, 11 4, 10, 6, 5 5, 3, 11, 3 f o r e s t 2 11, 11, 11, 11 4, 11, 11, 11 5, 11, 11, 11 3 11, 5, 4, 7 6, 7, 11, 11 10, 4, 7, 6 4 5, 11, 11, 4 11, 11, 11, 11 t * t , t , t 5 11, 11, 4, 4 11, 11, 11, 11 t , t , t , t 15 m i n 1 3, 7, 11, 3 4, 7, 3, 4 t , t , t , t f o r e s t 2 11, 6, 11, 11 11, 11, 11, 11 t , t , t , t 3 11, 7, 8, 9 4, 10, 11, 11 t , t , t , t 4 11, 5, 11, 11 5, 5, 5, 7 7, 3, 11, 11 5 11, 3, 11, 11 4, 6, 5, 11 5, 5, 9, 4 at ecotone 1 8, 7, 4, 8 4, 5, 4, 7- 4, 7, 6, 4 2 11, 11, 5, 11 5, 3, 4, 2 4, 2, 5, 4 3 4, 4, 6, 6 3, 3, 4, 2 2, 3, 1, 7 4 5, 3, 4, 6 4, 3, 3, 5 3, 5, 4, 2 5 4, 3, 3, 5 6, 4, 5, 11 3, 4, 5, 6 15 m i n 1 8, 4, 2, 2 3, 7, 11, 8 2, t , 1, 2 open 2 4, 8, 4, 5 5, 7, 4, 7 10, 3, 1, 6 3 11, 4, 5, 4 4, 4, 5, 4 2, 2, 4, t 4 4, 9, 7, 2 10, 4, 10, 5 2, 2, 1, 1 5 4, 8, 3, 2 5, 10, 7, 8 2, 2, 3, 3 76 m i n 1 2, 5, 2, 9 4, 5, 11, 11 2, 2, 1, 5 open 2 2, 4, 6, 11 7, 5, 3, 7 11, 5, 3, 3 3 6, 10, 10, 4 5, 4, 7, 11 2, 2, t , 4 4 3, 3, 7, 7 6, 3, 4, 4 t , t , 6, t 5 6, 4, 4, 3 4, 7, 5, 4 3, 7, 2, 1 * t - snow present but no reading given by penetrometer. 547 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 Distance i n t o f o r e s t Forest Distance i n t o open A l l s t a t i o n Month 76m 15m edge 15m 76m means TEMPERATURE (C) Nov -2 -2 -2 -7 -1 -3 Dec -11 -12 -10 -17 -10 -13 Jan -10 -10 -5 -17 -7 -10 Feb -5 -7 -7 -15 -3 -7 Mar 1 0 0 -5 2 -1 Apr 4 4 4 -1 5 3 May 10 6* 8 3 8 7 Means -2 -3 -2 -8 -1 -3 RELATIVE HUMIDITY (%) Nov m** m m m m m Dec 84 83 84 79 63* 79 Jan 75 74 75 75 59* 72 Feb 76 69 77 71 56* 70 Mar 63 69 71 65 52* 64 Apr 58 61 66 59 39 56 May 55 66 70 60 55 61 Means 68 71 74 68 54 67 *Estimated as described i n s e c t i o n 9.2.3. **Missing. 548 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 Distance i n t o f o r e s t Distance i n t o open Buckhorn Month 76m 15m 15m 76m s i t e (1070m) Oct ( 8 ) * 13 9 Nov (42) 8 4 Dec (24) 16 8 9 33 Jan (34) 22 12 24 64 164(37)* Feb (26) 16 9 12 39 104(23) Mar (27) 23 16 16 46 115(55) Apr (28) 31 27 22 66 J u l (91) 31 33 16 47 *No. of day sampled. Table H-4 Snow Depths Ecotone at the (cm) Grove Across the Study Area West-, 19 72 Facing -73' Winter Date Distance i n t o f o r e s t Forest Distance i n t o open sampled 122m 76m 15m edge 15m 76m 122m Nov 1 0* 0 0 0 0 0 0 Dec 5 10 5 8 10 15 15 15 Jan 5 33 53 28 74 89 61 86 Feb 8 25 10 20 69 51 53 53 Mar 6 28 . 8 18 46 58 61 58 Apr 4 23 3 0 30 46 56 48 Apr 30 0 0 0 0 0 0 0 *n = 1 f o r each f i g u r e . APPENDIX I GLOSSARY OF TERMS AND ABBREVIATIONS USED IN THE TEXT 550 Most of the d e f i n i t i o n s are taken from Daubenmire (1959) , Odum (1971) , and the S o c i e t y f o r Range Management, Range term glossary committee (1974). Allometry: The r e l a t i v e growth of a p a r t i n r e l a t i o n to an e n t i r e organism. Basal area: The c r o s s - s e c t i o n a l area of a t r e e at 1.4 m above the ground, u s u a l l y expressed as the sum of the b a s a l areas of t r e e s i n a f o r e s t i n square f e e t per acre or square meters per hectare. Browse: N, that p a r t of l e a f and twig growth of shrubs, woody vines and t r e e s a v a i l a b l e f o r animal consumption. V, the act of e a t i n g browse. Burn: Canopy coverage Cl e a r c u t : Climax: Community; Crown clo s u r e A land area over which a w i l d f i r e has passed, t y p i c a l l y r e c e n t l y but i s a l s o a p p l i e d to burned-over f o r e s t e d areas that have f a i l e d to regenerate to t r e e s . The percentage of ground covered when a polygon drawn about the e x t r e m i t i e s of the undisturbed canopy of each p l a n t i s p r o j e c t e d upon the ground, and a l l such p r o j e c t i o n s on a given area summed ( i n f l o r e s c e n c e s excluded). N, the area i n a f o r e s t remaining a f t e r the f e l l i n g of t r e e s . In a commerical c l e a r c u t , only merchantable trees are f e l l e d ; i n a b i o l o g i c a l c l e a r c u t , a l l t r e e s are f e l l e d . V, the f e l l i n g of trees i n a s p e c i f i e d area. The t h e o r e t i c a l l y t e r m i n a l , s t a b i l i z e d stage of a sere capable of perpetuation under the p r e v a i l i n g c l i m a t i c and edaphic c o n d i t i o n s ; a community i n dynamic e q u i l i b r i u m which w i l l not p r e d i c t a b l y change i t s composition or physiognomy. Any assemblage of p l a n t s and animals l i v i n g i n a p r e s c r i b e d area or p h y s i c a l h a b i t a t , t y p i c a l l y i n a common s p a t i a l arrangement. The p r o p o r t i o n of the a v a i l a b l e space i n a s p e c i f i e d t r e e or shrub l a y e r t h a t i s occupied by trees or shrubs. 551 Crown cover: Cunit: Cutover: DBH: The canopy formed by the tree l a y e r i n a f o r e s t . 100.cubic f e e t of timber. A s p e c i f i e d area i n a f o r e s t from which some or a l l of the trees.have been removed, u s u a l l y a p p l i e s to merchantable t r e e s . The diameter of a t r e e at 1.4 m above ground l e v e l . D e c o r t i c a t e : To remove the bark from. D i g e s t i b i l i t y : The p r o p o r t i o n of a forage taken i n t o the d i g e s t i v e t r a c t that i s absorbed i n t o the body. Ecotone: F l o r i s t i c s Forage: Forb: Forest type: Frequency of occurrence: A t r a n s i t i o n between two or more d i f f e r e n t communities; a j u n c t i o n zone t h a t i s narrower than the a d j o i n i n g communities and that has features of neighbouring communities as w e l l as c h a r a c t e r i s t i c s of i t s own. A q u a n t i t a t i v e assessment of the kinds of p l a n t species growing i n a defined area or community. N, a l l browse, herbaceous and lichenous food that i s a v a i l a b l e to a defined group of animals. V, the act of e a t i n g forage. Any herbaceous p l a n t other than those i n the Gramineae, Cyperaceae, and Juncaceae f a m i l i e s . A f o r e s t stand e s s e n t i a l l y s i m i l a r throughout as regards f l o r i s t i c composition, physiognomy and e c o l o g i c a l s t r u c t u r e . A s t a t i s t i c a l expression of the presence and absence of i n d i v i d u a l s of a species i n a s e r i e s of sample p l o t s , uniform i n shape and area, l o c a t e d . i n one or more stands of a kind. Constancy i s the number of stands i n which a species i s recorded compared t o the t o t a l number of stands sampled of one k i n d . Graminoid: A herb belonging to the Gramineae, Cyperaceae or Juncaceae f a m i l i e s . 552 H a b i t a t : The sum t o t a l of environmental c o n d i t i o n s of a s p e c i f i c place where an animal or p l a n t l i v e s ; the s p a t i a l niche. Habitat The occurrence of animals i n h a b i t a t s more s e l e c t i o n : f r e q u e n t l y than expected by chance; the behavior of animals that r e s u l t s i n them choosing to occupy h a b i t a t s . Habitat-type: A c o l l e c t i v e term f o r those u n i t s of the landscape c h a r a c t e r i z e d by measurable d i f f e r e n c e s i n v e g e t a t i o n , or by d i f f e r e n c e s i n p o s i t i o n on the landscape: d i s t i n g u i s h e d from the d e f i n i t i o n of a land u n i t that i s capable of producing a c e r t a i n (predictable) k i n d of climax vegetation. For example, r i p a r i a n h a b i t a t type, a burn h a b i t a t - t y p e , a mature f o r e s t h a b i t a t - t y p e . Habitat use: Occupation of a h a b i t a t . Herbs: Any f l o w e r i n g p l a n t except those t h a t develop p e r s i s t e n t woody stems above ground. I n t e r c e p t i o n : The process by which p r e c i p i t a t i o n i s r e t a i n e d by leaves, branches, boles and other organs of p l a n t s before the moisture reaches the ground. I n t e r c e p t i o n Intercepted moisture by p l a n t s that i s l o s t l o s s : through evaporation. L i g n i n : The complex non-carbohydrate c o n s t i t u e n t of the w a l l s of c e r t a i n c e l l s , e s p e c i a l l y i n woody t i s s u e . M i c r o c l i m a t e : L o c a l combinations of atmospheric f a c t o r s which d i f f e r from the p r e v a i l i n g macroclimate due to v a r i a t i o n s i n p l a n t cover, topography, slope p o s i t i o n , and pr o x i m i t y to l a k e s ; the climate of the microhabitat. Mixed f o r e s t : A f o r e s t composed of two or more tree sp e c i e s , one of which i s deciduous and at l e a s t two of which each comprises at l e a s t 2 0 per cent of the t r e e s present. P a r t i a l l o gging: A logging method i n which only a p o r t i o n of the merchantable trees are cut. 553 Phytomass: Primary p r o d u c t i v i t y : Puckerbrush: The t o t a l q u a n t i t y at a given time of l i v i n g p l a n t s of one or more species per u n i t of space (species phytomass), or of a l l the species i n a community (community phytomass). The rate at which r a d i a n t energy i s stored by photosynthetic and chemosynthetic a c t i v i t y of producer organisms i n the form of organic substances which can be used as food m a t e r i a l s . Gross primary p r o d u c t i v i t y i s the t o t a l rate of photosynthesis i n c l u d i n g organic matter used i n r e s p i r a t i o n . Net primary p r o d u c t i v i t y i s the net rate of storage i n excess of organic matter u t i l i z e d f o r r e s p i r a t i o n during the p e r i o d of measurement. Used i n t h i s t h e s i s t o r e f e r to above ground production. Thickets of a l d e r , w i l l o w , aspen, maple or other s i m i l a r species; t y p i c a l l y l e s s than nine m t a l l . Quamaniq: S e l e c t i v e logging: Sere: S e r a i stage: An Eskimo term used to describe the depression i n snow underneath t r e e s , due to i n t e r c e p t i o n . The f e l l i n g system of removing only c e r t a i n trees such as the l a r g e s t ones i n a f o r e s t . Two commonly used v a r i a n t s are \"mark s e l e c t i o n \" where trees f o r f e l l i n g are i n d i v i d u a l l y marked, and \"diameter l i m i t s e l e c t i o n \" where minimum harvestable diameters are s p e c i f i e d f o r merchantable species. The e n t i r e sequence of communities that replace one another on a given area. One of the r e l a t i v e l y t r a n s i t o r y stages or communities t h a t i n t o t a l make up the sere. Shrub: Any f l o w e r i n g p l a n t that has p e r s i s t e n t , above ground, woody stems and t h a t t y p i c a l l y produces at l e a s t s e v e r a l b a s a l shoots i n s t e a d of a s i n g l e b o l e , and t h a t has a low growth form. Snow hardness: See snow penetrance, Snow penetrance: The force r e q u i r e d to break through a snow c r u s t l a y e r of a defined t h i c k n e s s . 554 S o i l A group of s o i l s e r i e s that occur on s i m i l a r a s s o c i a t i o n : parent m a t e r i a l s and tha t form a n a t u r a l p a t t e r n as a r e s u l t of v a r i a b l e drainage or some other f a c t o r s ; a group of defined and named taxonomic s o i l u n i t s , r e g u l a r l y g e o g r a p h i c a l l y a s s o c i a t e d i n a defined p r o p o r t i o n a l p a t t e r n . Stand: An e f f e c t i v e aggregation of one or more s p e c i f i c p l a n t species w i t h more or l e s s u n i f o r m i t y i n physiognomy, composition and h a b i t a t c o n d i t i o n s . Standing The t o t a l amount of the biomass (phytomass) of crop: organisms of one or more species w i t h i n a defined area. Succession: The non-phenological, d i r e c t i o n a l change i n communities. Swamp, open A land area.containing excessive water much of and brush: the year and covered w i t h vegetation. Open swamps (synonymous w i t h marsh) lack w e l l developed t r e e and shrub l a y e r s , brush swamps have a more or l e s s continuous l a y e r of shrubs. Tree: Any f l o w e r i n g p l a n t that t y p i c a l l y has a s i n g l e p e r s i s t e n t woody stem and commonly more than 3 m t a l l . V a c c i n i a : A general term used to describe members of the genus Vaaainium. Vegetation A p l a n t community wi t h d i s t i n g u i s h a b l e type: c h a r a c t e r i s t i c s , of any s i z e , rank, or stage of succession. Wind run: The average v e l o c i t y of wind during a s p e c i f i e d time p e r i o d and at a s p e c i f i e d height above the ground. "@en . "Thesis/Dissertation"@en . "10.14288/1.0094631"@en . "eng"@en . "Plant Science"@en . "Vancouver : University of British Columbia Library"@en . "University of British Columbia"@en . "For non-commercial purposes only, such as research, private study and education. Additional conditions apply, see Terms of Use https://open.library.ubc.ca/terms_of_use."@en . "Graduate"@en . "Habitat selection and use in winter by moose in sub-boreal forests of north-central British Columbia, and relationships to forestry"@en . "Text"@en . "http://hdl.handle.net/2429/21531"@en .