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Regional Gradient Analysis and Spatial Pattern of Woody Plant Communities in Oregon

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Regional Gradient Analysis and Spatial Pattern of Woody Plant Communities in Oregon AN ABSTRACT OF THE THESIS OF Janet L. Ohmann for the degree of Doctor of Philosophy in Forest Science presented on November 29. 1995. Title: Regional Gradient Analysis and Spatial Pattern of Woody Plant Communities in Oregon. Abstract approved: Thomas A. Spies Knowledge of regional-scale patterns of plant community structure and controlling factors is largely qualitative and based on numerous local studies. Data from a subsample of 10,000 field plots were used to quantify and map compositional gradients of woody plant communities across Oregon forests. Canonical correspondence analyses explained 9-14% of the total species variation at three geographic scales. Climate contributed most to total variance explained (44-58%), followed by geology (6-16%), disturbance (7-13%), and topography (4- 8%). The dominant compositional gradient was associated with climate, from the lower elevation, moderate, maritime climate of the coast to the higher elevation, drier, continental climate of the interior. The second canonical axis followed a gradient from the warm, dry growing seasons of the interior valleys and eastern Cascades, to cooler, wetter mountainous areas. Only those parent materials that present extreme growing conditions, primarily ultramafic soils, were significant at the regional level. Ecological relationships differed substantially among subregions within the state. Except for topography, which contributed more to explained variation at finer geographic scales, ecological differences among subregions overshadowed effects of geographic extent. Topographic effects were stronger and beta diversity greater in eastern Oregon, where moisture is more limiting for plant growth, and weakest near the coast where climate is more favorable. The secondary importance of topography, disturbance, and substrate can be attributed to their influence on relative abundances of species within a local area, rather than on species presence or absence within broader regions. Community structure varied at a finer spatial scale in eastern than in western Oregon, and species turnover along gradients was greater for shrubs than trees. Amount of unexplained variation in the species data was high but not atypical of gradient analyses. Yet spatial structure in the species data that was uncorrelated with explanatory factors suggests potential to improve the canonical correspondence analysis models, particularly in eastern Oregon. Study findings have implications for considering biological diversity in regional conservation strategies, in planning for global climate change, and in the design of regional inventory and monitoring programs. Regional Gradient Analysis and Spatial Pattern of Woody Plant Communities in Oregon by Janet L. Ohmann A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy Completed November 29, 1995 Commencement June 1996 Doctor of Philosophy thesis of Janet L. Ohmann presented on November 29. 1995 APPROVED: Major Professor, representing Forest Science Head of Department of Forest Science Dean of Graduate School I understand that my thesis will become part of the permanent collection of Oregon State University libraries. My signature below authorizes release of my thesis to any reader upon request. Janet L. Ohmann, Author ACKNOWLEDGEMENTS Completing a study of this size and complexity would have been impossible without the financial, technical, and moral support of numerous individuals and organizations. Lisa Lackey and Paul Hardwick gave freely of their time and expertise in applications of geographic information systems (GIS) technology. George Lienkaemper also provided several GIS coverages and technical support. Barbara Marks, David Wallin, and John Kimball were collaborators in executing the temperature model on our particular input dataset. I'm indebted to Brad Smith for generously providing several analytical tools, namely his "SOLARPDX," "ECOAID," and "ECOTOOLS" software programs, as well as a copy of the Eastside Ecology Database and the use of his southwest Oregon plot dataset. Denis White provided the GIS coverage with documentation for the EMAP hexagons. The efforts and expertise of countless individuals went into collecting, editing, and managing data from the thousands of field plots I incorporated into my ana'ysis. Numerous professionals of the Inventory and Analysis Program have toiled over decades to make possible the invaluable plot dataset for nonfederal lands. Ecologists and others in the Region 6 Area Ecology Program generously provided data, ecological consultation, and technical assistance. They included (alphabetically): Tom Atzet, Nancy Diaz, Fred HaIl, Miles Hemstrom, Bill Hopkins, Bill Johnson, Charlie Johnson, Cindy McCain, Lisa McCnmmon, and Lenny Volland. Thanks especially to committee chair and colleague, Tom Spies, for his guidance, insightful suggestions, and patience throughout all phases of my study. Members of my graduate corrimittee, Bill McCornb, Ron Neilson, Reed Noss, and Steve Sharrow, also provided helpful suggestions on thesis content. My funding came from the Inventory and Economics Research, Development, and Application Program and the Ecosystem Processes Research Program of the Pacific Northwest Research Station, USDA Forest Service. In particular, I'm grateful to Dan Oswald for launching me on this endeavor, and for continued support and patience through its completion. Last but not least, I thank my friends for sticking it out with me over the years of my project. TABLE OF CONTENTS Page, INTRODUCTION I Current Knowledge of Regional Patterns in Forest Communities 4 Research Objectives and Hypotheses 9 METHODS 12 Study Area 12 Physiography, Geology, and Soils 12 Climate 20 Vegetation 24 Disturbance and Landscape Patterns 27 Paleoecology 31 Vegetation Data 32 Explanatory Variables 34 Field-Recorded Attributes 34 Temperature 36 Precipitation 37 Solar Radiation 38 Geology 38 Spatial Position 39 Gradient Analyses 39 CCA with Variance Partitioning 40 Stepwise CCA 41 Indirect Gradient Analyses 44 Mapping Geographic Patterns of Dominant Gradients 45 Community Classification 45 RESULTS 46 Variance Partitioning Analyses 46 Importance of Spatial Position and Environment 46 Importance of Local and Regional Explanatory Variables 48 TABLE OF CONTENTS (Continued) Paae Stepwise CCA 50 Gradient Structure 50 Dominant Gradients at the State Level 50 Dominant Gradients at Half-State and Subregion Levels 72 Positions of Plant Growth Forms Along CCA Gradients 83 Spatial Patterns in Fit of the CCA Model 87 Comparison of Direct and Indirect Gradient Analyses 95 Comparison of Tree and Shrub Strata 96 Community Patterns 98 Ecotones in Oregon Plant Communities 104 Ecological Differences Between Land Ownerships 108 DISCUSSION 112 Determinants of Vegetation on a Site 112 Spatial Structure in the Species and Environment Data 112 Temperature and Moisture 114 Topography 117 Overstory Canopy Cover 119 Disturbance 120 Geology 126 Within-Region Variation in Patterns of Community StrlJcture and Environment 126 Sampling and Scale Effects 128 Sample Size and Rare Species 129 Effects of Geographic Scale 131 Resolution of the Species Data 134 Limitations and Sources of Unexplained Variation 135 TABLE OF CONTENTS (Continued) Page Summary of Research Hypotheses 137 Objective One 137 Objective Two 140 IMPLICATIONS FOR CONSERVATION AND RESOURCE MANAGEMENT 141 Regional Biodiversity Conservation Strategies 141 Planning for Global Climate Change 144 Monitoring Biodiversity in Forest Ecosystems 145 CONCLUSIONS 147 BIBLIOGRAPHY 150 APPENDICES 161 LIST OF FIGURES Ficiure Paqe Map of Oregon showing major cities, rivers, and physiographic 3 features, subregion boundaries, and latitude/longitude. Tentative distribution of some of the major forest zones within an 8 environmental field based on moisture and temperature indices (Emmingham 1982) (adapted from Franklin and Dyrness 1973, p. 50). Maps of selected explanatory variables, a. Elevation (m); 17 b. Mean annual precipitation (mm); c. Mean growing-season (May-September) precipitation (mm); d. Seasonal variability in precipitation (coefficient of variation of wettest and driest months (usually December and July), mm); e. Percent of mean annual precipitation that falls in June-August; f. Mean monthly temperature (°C) during the growing-season (May-September); g. Seasonal variability in temperature (coefficient of variation of mean monthly temperature (°C rescaled to a 0-255 scale) of the coldest and warmest months (usually December and August). Precipitation and temperature maps are based on Daly et al. (1994) and Marks (1990). Geologic types of Oregon. 21 Federal land ownerships in Oregon. 26 Stand age and history of clearcutting on field plots. 30 Relations between sample size (number of plots) and number of 47 species, total inertia (TI), and total variation explained (WE) for CCAs of all woody species. Samples are the five subregions, two half-states, and the entire State. Biplots from CCA of all woody species, showing species with 55 5% constancy. a. Oregon, axes I and 2; b. Oregon, axes 2 and 3; c. Western Oregon, axes I and 2; d. Eastern Oregon, axes I and 2; e. Coast subregion, axes I and 2; f. Western Cascades subregion, axes I and 2; g. Klamath subregion, axes I and 2; h. Eastern Cascades subregion, axes I and 2; i. Blue Mountains subregion, axes I and 2. Maps of plot scores (linear combinations) from canonical 64 correspondence analysis (CCA) of all woody species, Oregon. a. axis 1; b. axis 2; c. axis 3;
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