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FLORISTIC AFFINITIES of the SAN JOAQUIN ROADLESS AREA, IΝΥΟ NATIONAL FOREST, MONO COUNTY, CALIFORNIA by Helen M. Constantine

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FLORISTIC AFFINITIES of the SAN JOAQUIN ROADLESS AREA, IΝΥΟ NATIONAL FOREST, MONO COUNTY, CALIFORNIA by Helen M. Constantine FLORISTIC AFFINITIES OF THE SAN JOAQUIN ROADLESS AREA, IΝΥΟ NATIONAL FOREST, MONO COUNTY, CALIFORNIA by Helen M. Constantine-Shull A Thesis Presented to The Faculty of Humboldt State University In Partial Fulfillment of the Requirements for the Degree Master of Arts In Biology May, 2000 FLORISTIC AFFINITIES OF THE SAN JOAQUIN ROADLESS AREA, INΥΟ NATIONAL FOREST, MONO COUNTY, CALIFORNIA by Helen M. Constantine-Shul1 Approved by the Master's Thesis Committee: John O. Sawyer Jr., Major Professor Mike Mesler, Committee Member Ken Lang, Committee Member Robert Curry, Committee Member Milton Boyd, Graduate Coordinator Ronald A. Fritzsche, Dean for Research and Graduate Studies ABSTRACT I surveyed the flora of the San Joaquin Roadless Area, a 44km2 area on the east side of the central Sierra, and then analyzed its floristic affinities (origins) using Sorensen's similarity coefficient, cluster analysis, and Bray-Curtis ordination. The analyses compare the area's flora to 12 other California mountain floras, and suggest an origin for recolonizing taxa after severe disturbances by Holocene volcanic activity. I have described the historic and present climate, geology, physiography, and vegetation of the area for a better understanding of the origin and condition of the flora. The San Joaquin Roadless Area flora consists of 446 taxa, including two rare (CLAPS List 1Β) plants, Lupinus duranii Eastw. and Arabis pinzlae Rollins. The ranges of 3 taxa were extended. The flora is centered in geographic location, species composition, and proportion of phytogeographic elements in relation to other California mountain floras. Its species composition is most similar to Tuolumne Meadows (56%) and vicinity and least similar to the San Bernardino Mountains (30%). The average similarity between all 13 floras examined is 39%. The cluster analysis suggests three major groups of floras, 1) Sierran and Westside floras, 2) Great Basin and eastside floras, and 3) floras not similar to either group. My flora is included with other Sierran and Westside floras. The Bray-Curtis ordination shows a similar pattern, but with the three axes accounting for only 29% of the variation. Ordinations on subsets of the floras (1: Floras of similar size, 2: Floras close to the San Joaquin Roadless Area, and 3: Sierran floras) also show a similar pattern, with the three axes accounting for >70% of the variation between the floras. The phytogeographic affinities of the entire San Joaquin Roadless Area are generally consistent with those of the entire Sierran flora. However, within each affinity category they differ. The San Joaquin Roadless Area contains a higher percentage of Old iii Cordilleran taxa, and lower percentages of Circumboreal, Northern Hemisphere, Lowland California, and Great Basin taxa. A cluster analysis of the percentages in each affinity category shows a similar pattern to that of the other analyses. Conflicting extremes of wet and dry environments, due to weather patterns and volcanic soils, have come together in the San Joaquin Roadless Area to create a moderate environment that supports a newly immigrated, centrally aligned flora. The availability of favorable conditions has influenced the relative success of colonizing taxa with different origins and adaptations. The proportions of taxa from each region of origin are reflected in the dominating moisture regimes of the area. My results confirm the Mammoth Gap as an effective migration corridor for Lowland California taxa to the east side of the Sierra. Despite destructive volcanic disturbances, there is a possibility of alpine refugia where some of the first colonizers of the study area survived. A Sierran origin for the rare plant, Arabis pinzlae is suggested. iv ACKNOWLEDGEMENTS This thesis would not have been possible without the wonderful and generous support of many people. I would like to thank everyone involved for helping me make this project happen. My major professor, John Sawyer, encouraged me through the years with unending patience and perceptive guidance. I will always cherish John's inspiring classes and our journeys to the mountains and deserts. His extensive knowledge and engaging, enthusiastic discussions helped me to better understand the plants and their setting. My committee members, Mike Mesler, Ken Lang, and Robert Curry provided valuable comments and feedback on the thesis. I am fortunate to have taken Dr. Mesler's plant taxonomy class and taught his laboratory. These gave me a strong foundation for identifying my specimens. Dr. Curry of UC Santa Cruz—CSU Monterey Bay contributed his first hand knowledge of the area and its geology and traveled to the study area to hike with me. I am especially grateful to Dr. G. Ledyard Stebbins who shared his knowledge on the origins of Sierran plants and determined the phytogeographic affinity of each taxon on my species list. Bruce Bingham of the U.S. Forest Service provided valuable computer assistance with access to the Paradox database and matrix format conversion. My thanks and appreciation go out to Connie Millar of the U.S. Forest Service for securing funding for this project through the Sierra Nevada Ecosystem Project. Her guidance and knowledge on the paleobotany of the area provided me with insight as to the origins of the area's flora. The Biology Graduate Student Association also provided financial assistance for thesis binding. Stew Winchester of Diablo Valley College opened my eyes to the world of plant ecology and encouraged me to continue my studies at the graduate level. Dr. Dennis Walker clearly and thoroughly introduced me to botany and gave me the chance to teach and apply v my art to the science. Sally Miller of Friends of the Ingo was my inspiration for studying the San Joaquin Roadless Area. She also provided many unpublished local references. The contributions of many others helped make this project a success. Teresa Sholars of College of the Redwoods, Ft. Bragg and James Morefield of the Nevada Natural Heritage Program confirmed my rare plant identifications. Gordon Leppig provided plant presses and helped with identifications in the HSU Herbarium. Botanical professionals Dean Taylor and Mark Bagley offered their guidance and information on plants and literature resources. Greg Reis of the Mono Lake Committee introduced me to the world wide web and its many climate data resources. Richard Perloff and Kathleen Nelson of the U.S. Forest Service issued me collecting permits. My peers, Margaret Willits, Ann Francis, Stassia Samuels, and Julie Evans among others shared valuable knowledge and observations with me. Special thanks go out to all of my friends on the Eastside and at the Mono Lake Committee for their encouragement, companionship, and places to stay while I was doing my field work. Tommy Kashirsky provided a warm, dry base camp with protection from mosquitoes and a large table on which to sort and press my specimens. Finally, I would like to thank my family whose loving support made all this possible. My Mother, my fearless and dedicated field partner, kept me company, learned botany, and added several new plants to my species list. I am forever thankful to my husband, Richard, for his patience and moral support throughout this project — in the midst of starting our marriage and our own business. He helped me in the field when he could get away, and did everything when I was gone or writing for weeks at a time. He always reminded me of the significance of my work, and encouraged me when the finished thesis seemed so far away. His parents, Marian and Blaine Shull, helped me keep this project a top priority. I am thankful for their encouragement and support throughout the years. And thanks, always, to the mountains. vi This thesis is dedicated to Dr. G. Ledyard Stebbins 1906-2000 for his inspiration and contributions to the sciences of evolutionary botany and phytogeography. TABLE OF CONTENTS Page ABSTRACT iii ACKNOWLEDGEMENTS v LIST OF TABLES xi LIST OF FIGURES xiii INTRODUCTION 1 Purpose 2 The study area 4 Location and boundaries 4 Physiography 4 Elevational zones 7 Geology 11 Recent volcanics 14 Glacial history 16 Soils 17 Climate 18 Regional 18 Local 19 Weather data 23 Human impacts 28 Research and botanical collecting history 30 Vegetation 31 Paleovegetation 31 Current vegetation 33 viii TABLE OF CONTENTS (CONTINUED) METHODS 44 Field methods 44 Specimen identification methods 46 Analysis methods 49 Phytogeographic affinities of the taxa 49 Comparison of my flora to other floras 52 Phytogeographic affinities of the floras 53 RESULTS 57 Floristic analysis 57 Rare and uncommon taxa 57 Range extensions 59 Alien taxa 60 Other noteworthy taxa 60 Noteworthy occurrences on the limestone 61 Noteworthy absences 62 Statistical analysis 62 Comparison of all floras 62 Comparison of floras of similar size 67 Comparison of floras close to the San Joaquin Roadless Area 70 Comparison of Sierran floras 73 Floristic affinities of the San Joaquin Roadless Area 76 DISCUSSION 83 Floristic affinities of the San Joaquin Roadless Area 83 Species composition analyses 83 Analyses of phytogeographic affinities 85 ix TABLE OF CONTENTS (CONTINUED) Mammoth Gap as a migration corridor 87 Origins of the San Joaquin Roadless Area flora 87 The origin of Arabis pinzlae 88 CONCLUSIONS 92 LITERATURE CITED 93 APPENDIX A. Annotated species list of the San Joaquin Roadless Area 106 APPENDIX B. Environmental variables 138 APPENDIX C. Similarity matrices 139 x LIST OF TABLES Page Table 1. Summary of average annual weather data for stations near the San Joaquin Roadless Area 25 Table 2. Locations of important collecting areas 47 Table 3. Phytogeographic elements as defined by Stebbins (1982, 1997 Pers. comm.) 51 Table 4. California mountain floras used in the statistical analyses 55 Table 5. Subsets of the thirteen floras used in the analyses 56 Table 6. Coefficients of determination for the correlations between ordination distances and distances in the original n-dimensional space from the analysis of all floras 66 Table 7.
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