Range Ecology of the Porcupine Caribou Herd in Canada Don £. Kussel 1
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Range Ecology of the Porcupine Caribou Herd in Canada Don £. Kussel 1 Art M. Martell Wendy A. C. Nixon Research, Management and Husbandry of Reindeer and other Northern Ungulates Special Issue No. 8,1993 Rangifer Published by: Nordic Council for Reindeer Research (NOR). Editor: Sven Skjenneberg Address: Postboks 378, N-9401 Harstad, Norway Telephone: +47 (0)82 64 172 Telefax: +47 (0)82 66 280 Editorial board Karstein Bye, The Reindeer Husbandry Administration, N-9500 Alta, Norway Öje E. Danell, The Swedish University of Agriculture, Institute of Forest Improvement, Bo: 7007, S-750 07 Uppsala, Sweden Raimo Hissa, University of Oulu, Department of Zoology, Zoophysiological Laboratory, Lin namaa, SF-90570 Oulu, Finland Christian Krogell, Forestry- and Agriculture Ministry, Hunting and Fishing Department, Estnäs gatan 7 F, Helsingfors, Finland Peter Hansen, Greenland Home Rule Authority, Department of Nature Management, Bo; 309, DK-3900 Nuuk, Greenland Nicholas J. C. Tyler, University of Tromsø, Department of Arctic Biology, Breivika, N-9037 Tromsø Norway Robert G. White, University of Alaska Fairbanks, Institute of Arctic Biology, Fairbanks, Alaskj 99775, U.S.A. ISSN 0801-6399 A/S Harstad Tidendes trykkeri Special Issue No. 8 RANGIFER Range Ecology of the Porcupine Caribou Herd in Canada Don E.Russell ArtM. Martell Wendy A. C. Nixon Published by Nordic Council for Reindeer Research (NOR) Harstad, Norway, 1995 RANGIFER Vol. 13 1993 Special Issue No. 8 Contents Page List of tables 8 List of figures 10 Acknowlegements 13 Executive summary 14 Introduction 14 Winter 14 Characteristics of the range 14 Food habits 15 Activity 15 Cratering dynamics 15 Late spring 15 Characteristics of the range 15 Habitat selection 1° Food habits 16 Activity 17 Late spring range use by bulls 17 Summer 17 Movements and distribution 18 Insects and weather 18 Climate and weather patterns 18 Caribou response to insects 18 Activity, food and habitat • 18 Energetic implications 19 CHAPTER 1. Introduction 21 CHAPTER 2. Winter 27 Introduction 27 Characteristics of the winter range 28 Study area 28 Climate 28 Rangifer, Special Issue No. 8, 1993 3 Terrain 28 Surficial geology 29 Historical winter distribution 30 Methods 34 Snow • 34 Lichen biomass 35 Fire history 36 Results • 36 Snow 36 Lichen biomass 40 Fire history 42 Discussion 42 Snow 42 Lichen biomass 43 Fire history 46 Summary : 47 Food habits 47 Methods 47 Results and discussion 48 Effect of analysis at different times 48 Differences among years 48 Differences over winter 49 Seasonal diets 49 Fecal sample versus ingested diet 49 Conclusions 51 Winter activity 51 Methods 51 Results 53 Band size 53 Activity pattern 53 Activity budgets 54 Proportional use of active periods 57 Discussion 57 Effect of daylength on percent lying 57 Activity budgets 59 Range quality 60 Cratering dynamics 61 Methods 61 Results and discussion 63 Conclusions 70 Conclusions from winter 70 CHAPTER 3. Late spring 73 Calving grounds 73 Introduction 73 Study area 74 Porcupine caribou calving distribution 75 Characteristics of the range 78 Methods 78 4 Rangifer, Special Issue No. 8, 1993 Weather 78 Biomass 79 Tussock density 79 Phenology 79 Nutrients 79 Results and discussion 79 Weather • • • • 79 Phenology • 80 Biomass 81 Tussock density 82 Nutrients 86 Summary 88 Habitat selection 88 Methods 88 Results and discussion • 89 Food habits 91 Methods 91 Results 91 Effect of analysis at different times 91 Differences between periods 91 Differences among years 91 Fecal sample versus ingested diet 92 Discussion 92 Caribou activity 95 Methods 95 Results 95 Band Size 95 Movement rate 95 Active and bedded cycles 95 Activity budgets 97 Focal animal observations 97 Nursing 97 Discussion 99 Summary discussion 101 Spring range of bulls 103 Introduction 103 Study area 103 Methods 103 Distribution 103 Activity 103 Phenology 103 Habitat selection 103 Food habits 104 Data analysis 104 Results and discussion 105 Distribution • • 105 Activity 105 Phenology 107 Habitat selection •. 107 RangiSer, Special Issue No. 8 1993 5 Food habits 108 Summary and conclusions 109 Comparison of bulk and cows 110 CHAPTER 4. Summer 113 Introduction 113 Study area . : 113 Summer movements 114 Historical 114 Herd movements - 1984, 1985 116 Methods 116 Results 117 Discussion 118 Activity of dipterans in relation to date and weather 120 Introduction 120 Methods 121 Weather 121 Mosquito sampling 121 Oestrid flies 121 Result 122 Mosquito sweeps 122 Discussion 123 Influence of season and weather on Dipterans 123 Climate and weather patterns 126 Introduction 126 Methods 126 Results and discussion 126 Data summaries 126 Average daytime air temperatures 127 Average hourly air temperatures 127 Average hourly wind speeds 128 Correlations between permanent and temporary sites 128 Regional weather versus insects 129 Response of groups and individual caribou to insect harassment 131 Introduction • 131 Methods 131 Results 133 Group dynamics 133 Annoyance responses of individual caribou 133 Caribou annoyance responses in relation to date and weather 134 Relationship between position in group and insect harassment 134 Discussion 134 Caribou group dynamics 134 Behavioral responses of individual caribou 135 Position in a group 135 Activity budgets, food and habitat selection 135 Introduction 135 Methods 136 Results 136 Rangifer, Special Issue No. 8, 1993 Movement rates of caribou • Activity budgets and feeding intensity 138 Diurnal activity • 1^ Habitat use ^ Food habits I4* Discussion ^ Movement of groups and individuals 142 Activity 142 Feeding intensity and activity • 143 Conclusions *4^ CHAPTER 5. Energetic implications 147 Introduction *47 Methods 147 Results and discussion 149 References 157 Appendix A 162 Appendix B I64 Appendix C 167 Rangifer, Special Issue No. 8, 1993 7 LIST OF TABLES Table 1.1. The 15 periods in the annual life cycle of the Porcupine Caribou Herd. Table 2.1. Mean daily temperatures at two locations on the Dempster Highway, 1979-82. Table 2.2. Snow depths (cm) by snow regions in the winter range of the Porcupine Caribou Herd, winter 1979-80 to 1981-82. Table 2.3. Snow density (g/cm3) by snow region in the winter range of the Porcupine Caribou Herd, winter 1979-80 to 1981-82. Table 2.4. Snow hardness (kgx 10"2) by snow regions in the winter range of the Porcupine Caribou Herd, winter 1979-80 to 1981-82. Table 2.5. Average biomass of divot samples of each lichen species associated with each cover class (g/m2). Table 2.6. Regression of log real versus log estimated biomass by species. Table 2.7. Summary of lichen biomass (g/m2; mean±SD) by species or species group for the five geo• graphic regions of the Dempster Higway. Table 2.8. The proportional area, mean biomass and proportional biomass of the six range types within the five physiographic regions along the Dempster Highway. Table 2.9. Summary of stand age by community type. Table 2.10. Snow parameters from various Rangifer winter range studies. Table 2.11. Lichen biomass measurements from various Rangifer studies. Table 2.12. Average percentages and coefficients of variation (CV) of discerned plant fragments in repli• cates of caribou fecal samples that were analysed at one time (0) and replicates that were analy• sed at different times (D). Table 2.13. Average percentages ( ± SE) of discerned plant fragments in fecal samples collected from the range of the Porcupine Caribou Herd, 1979-82. Table 2.14. Average percentages of actual intake (mean ± SD (range]) and average percentages of discerned plant fragments in fecal samples from feeding trials on captive caribou and reindeer at the University of Alaska in 1982 by L. Duquette. Table 2.15. Length of winter lying and active periods and active/lying cycle for the Porcupine Caribou Herd, 1979-82. Table 2.16. Activity budgets (% time ± estimated 95% confidence limits) of caribou in north-central Yukon in winter, 1979-82. Table 2.17. Proportional use of active periods (% time ± estimated 95% confidence limits) by caribou in north-central Yukon in winter, 1979-82. Table 2.18. Crater measurements and regional snow parameters (SE in brackets). Table 2.19. Breakdown of cratering observations for the Porcupine Herd in winter and spring, 1979-82. Table 2.20. Summary of cratering activities (time in seconds) for the Porcupine Caribou Herd during the three winters 1979-82 (n mean SE). Table 2.21. Results of multiple regression relating caribou cratering activities to crater characteristics and regional snow parameters for the Porcupine Caribou Herd, 1979-82. Table 2.22. Comparison of mean values for snow parameters and caribou cratering characteristics for the Porcupine Caribou Herd, 1979-82. Table 2.23. Energetic parameters determined from caribou cratering activités on the range of the Por• cupine Caribou Herd. Table 3.1. New green biomass (g/m2) on the Yukon calving grounds of the Porcupine Caribou Herd, 1980. Table 3.2. Percent coverage of range types on the calving grounds of the Porcupine Caribou Herd as determined from aerial photographs. Total area mapped (ha) in parenthesis. 8 Rangifer, Special Issue No. 8, 1993 Table 3.3. New green biomass (g/m2) in different range types on the calving grounds of the Porcu• pine Caribou Herd in northern Yukon, 1980. Table 3.4. Summary of tussock variables for the eight stands sampled on the calving grounds of the Porcupine Caribou Herd. Table 3.5. Correlation of variables measured during estimation of tussock density to the total number of flowers per tussock. Table 3.6. Results of stepwise multiple regression and r2 at inclusion of independent variable in the prediction of number of flowers. Table 3.7. Estimate of tussock density, probable limit of error and Pielou's Index of Nonrandomness for the eight stands sampled. Table 3.8. Fiber, ash and nutrient content (% dry weight) of key species collected in 1980. Table 3.9. Percent of animals observed in the eight range types on the Yukon calving grounds of the Porcupine Caribou Herd, 1979-81.