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Interspecific Associations, Phenology, and Environment of Some Alpine Plant Communities on Lakeview Mountain, Southern British Columbia

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Interspecific Associations, Phenology, and Environment of Some Alpine Plant Communities on Lakeview Mountain, Southern British Columbia INTERSPECIFIC ASSOCIATIONS, PHENOLOGY, AND ENVIRONMENT OF SOME ALPINE PLANT COMMUNITIES ON LAKEVIEW MOUNTAIN, SOUTHERN BRITISH COLUMBIA by MARILYN JEAN RATCLIFFE B.Sc, The University. Of Victoria, 1979 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE FACULTY OF GRADUATE STUDIES Botany Department We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA July 1983 © Marilyn Jean Ratcliffe, 1983 J In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of BOTANY The University of British Columbia 1956 Main Mall Vancouver, Canada V6T 1Y3 Date JULY 29, 1983 DE-6 (3/81) i i Abstract Three major alpine plant communities were identified on Lakeview Mountain, Cathedral Provincial Park, using multivariate analysis of percentage cover data. Communities were dominated by Kobresia myosuroides, Carex scirpoidea (with one transitional area dominated by both Kobresia myosuroides and Carex scirpoidea), or by Carex scirpoidea and Carex capitata (with Salix nivalis as an additional dominant at one site). Community composition and distribution had little relationship with aspect or with the soils and microclimatic factors measured. Phenology was recorded for vascular species during the summer of 1980. Later flowering times were observed for a number of species in Kobresia myosuroides or Carex scirpoidea/Carex capitata dominated vegetation, and plants generally flowered earlier on southern aspects. Small scale patterns in the form of significant associations between species-pairs were detected in all communities, using a plotless point-line sampling technique. Patterns were abundant at this scale, with a total of 182 significant positive associations and 103 significant negative associations recorded between different species pairs. These interspecific associations varied considerably between sampled sites in the study area, with many occurring only once. Possible association-generating mechanisms have been discussed, and characteristics of the genotype, rather than the taxonomic species, have been suggested as critical in the formation of associations. A competitive hierarchy of dominant species has also been proposed, based on interspecific association and phenological data. Soils within the study area are classified as Alpine Dystric Brunisols, and are coarse textured, strongly acidic, low in available nutrients, and high in organic matter. Climate was relatively uniform over the study area during the 1980 growing season, as were microclimatic air and soil temperature profiles and air humidity profiles. Lower soil temperatures, however, occurred beneath Kobresia myosuroides dominated vegetation. iv Table of Contents Abstract ii List of Tables vi List of Figures vii Acknowledgements viii I. INTRODUCTION 1 THE ALPINE ZONE 2 LITERATURE REVIEW 4 OBJECTIVES 10 11 . STUDY AREA 11 LOCATION 11 LAND USE 13 GEOLOGY AND GEOMORPHOLOGY 15 GEOLOGY 15 GEOLOGICAL HISTORY 15 SOILS 16 CLIMATE 17 VEGETATION 18 ANIMALS 23 III. METHODS 24 VEGETATION 24 TRANSECT PLACEMENT 24 QUADRAT SAMPLING DESIGN 26 ANALYSES OF QUADRAT DATA 28 POINT-LINE SAMPLING 33 ANALYSIS OF POINT-LINE DATA 34 PHENOLOGY 38 NOMENCLATURE 39 SOILS 39 CLIMATE 41 IV. RESULTS 43 VEGETATION 43 COMMUNITY TYPES 43 1 . Transect A: 43 2. Transect B: 46 3. Transect C: 48 4. Transect D: 51 5. Transect E: ..........54 6. Transect F: 57 7. All Transects: ..60 PHENOLOGY 67 1. Constant Aspect: ...74 2. Constant Community Type: 75 INTERSPECIFIC ASSOCIATIONS 77 1 . Transect A: 78 2. Transect B: ..' 81 3. Transect C: 81 4. Transect D: .84 5. Transect E: 90 6. Transect F: 93 7. All Transects: 97 POSITIVE ASSOCIATIONS: 97 V NEGATIVE ASSOCIATIONS: 115 CONSTANCY OF INTERSPECIFIC ASSOCIATIONS 129 SPECIES ORDINATIONS 132 SOILS 135 MORPHOLOGY 135 PHYSICAL AND CHEMICAL PROPERTIES 135 CLIMATE 138 MESOCLIMATE 138 MICROCLIMATE 138 V. DISCUSSION 146 COMMUNITIES 146 PHENOLOGY 150 INTERSPECIFIC ASSOCIATIONS 152 POSSIBLE MECHANISMS GENERATING POSITIVE ASSOCIATION 154 1. Niche Differentiation: 154 2. Balanced Competitive Abilities: 158 3. Additional Mechanisms: 159 POSSIBLE MECHANISMS GENERATING NEGATIVE ASSOCIATION164 1. Morphology: 1 64 2. Abiotic Effects: 164 3. Competitive Exclusion: 165 GENOTYPIC RESPONSE 168 DOMINANT SPECIES 170 SOILS 175 CLIMATE 176 CONCLUSIONS 178 VI . SUMMARY 181 VEGETATION 181 SOILS 184 CLIMATE 184 VII. LITERATURE CITED 185 APPENDIX A - GEOLOGICAL HISTORY 213 APPENDIX B - SOILS 215 APPENDIX C - PERCENTAGE COVER DATA FOR SIX TRANSECTS ....217 APPENDIX D - PRINCIPAL COMPONENTS ANALYSIS OF SOIL DATA .224 vi List of Tables I. Selected Climatic Data for Weather Stations in Southern British Columbia (within the 1941-1970 period) . 19 II. Transect Characteristics 24 III. Methodology for Physical and Chemical Soil Analyses 40 IV. Associated Species 77 V. Positive Associations for 11 Sample Groups 98 VI. Negative Associations for 11 Sample Groups 116 VII. Constancy of Positive Associations 130 VIII. Constancy of Negative Associations 131 IX. Species Pairs Associated Both Positively and Negatively in Different Vegetation Groups 133 X. Physical and Chemical Properties from Soil Profiles within 11 Vegetation Groups 136 XI. Mesoclimatic Data For Three Weather Stations, 1: S84°W, 2481 m; 2: S2°E, 2402 m; and 3: N29°E, 2475 m. 139 vii List of Figures 1. Location of Study Site in Cathedral Provincial Park, British Columbia ..12 2. Position of Transects (A-F) in the Study Site 25 3. Quadrat Sampling Design 27 4. Transect A Multivariate Analyses 44 5. Transect B Multivariate Analyses 47 6. Transect C Multivariate Analyses 50 7. Transect D Multivariate Analyses 53 8. Transect E Multivariate Analyses 56 9. Transect F Multivariate Analyses 58 10. Centered PCA - All Data Sets 62 11. Centered and Standardized PCA - All Data Sets 63 12. RA - All Data Sets 64 13. Cluster Analysis - All Data Sets 66 14. Transect A Phenology 69 15. Transect B Phenology 69 16. Transect C Phenology 70 17. Transect D Phenology 71 18. Transect E Phenology 72 19. Transect F Phenology 73 20. Group A Positive and Negative Associations 80 21. Group B Positive and Negative Associations 82 22. Group C1 Positive and Negative Associations 83 23. Group C2 Positive and Negative Associations 85 24. Group D1 Positive and Negative Associations 87 25. Group D2 Positive and Negative Associations 88 26. Group D3 Positive and Negative Associations 89 27. Group E1 Positive and Negative Associations 91 28". Group E2 Positive and Negative Associations ....92 29. Group F1 Positive and Negative Associations 94 30. Group F2 Positive Associations 95 31. Group F2 Negative Associations 96 32. Microclimatic data for 11 vegetation groups during summer 1980 140 vi i i Acknowledgement I wish to thank my supervisor, Dr. Roy Turkington, for his assistance, enthusiasm, and guidance throughout the course of this study. Thanks are also extended to my commitee members, Drs. G. Bradfield and J. Maze, for providing valuable suggestions and critically reviewing the manuscript. I am grateful to Drs. Piet de Jong and Malcom Greig for statistical consultation, and to Dr. T. Ballard for advice regarding soil sampling and analysis and for providing laboratory facilities. The computer program for analysis of interspecific associations was written by Dave Zitton. I also thank Barry Wong and John Emanuael for computing consultation. Aid in the identification and verification of plant species was provided by Dr. G.W. Douglas and Terry Mcintosh. Immeasurable appreciation is extended to Dr. G.W. Douglas for providing advice, inspiration, and untiring interest during this research, particularly in the first field season. I also thank George A. Douglas, Mr. A.G. Ratcliffe, Melanie Madill, and Angela Chen for field and data manipulation assistance, and Tom Fleet and Karl Gehringer of Cathedral Lakes Resort for their interest in the study ix and for their friendship. Finally, very special thanks are due to Dr. K. Wilf Nicholls. His assistance with final figures and, most importantly, his unfailing patience, support, and understanding, made the completion of this thesis possible. Financial assistance through a National Science and Engineering Research Council of Canada postgraduate scholarship is gratefully acknowledged. 1 I . INTRODUCTION Alpine vegetation ecology has been essentially limited to the consideration of community patterns in relation to abiotic factors, such as time of snow release, microclimate, and physical and chemical properties of the soil. Savile (1960) has stated that in severe environments such as the alpine, competition is unimportant compared to physical factors, "allowing essentially random occurrence of plants without distinct associations and frequent coexistence of related species that have extremely similar requirements". This opinion is supported by Bliss (1962). Whittaker (1975) argued along similar lines and considered evolution to be affected more strongly by selection for survival in relation to problems of the physical environment and less strongly by selection involving interaction and competition with other species.
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