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A Systematic and Ecological Study of Astragalus Diaphanus (Fabaceae) Redacted for Privacy Abstract Approved: Kenton L

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A Systematic and Ecological Study of Astragalus Diaphanus (Fabaceae) Redacted for Privacy Abstract Approved: Kenton L AN ABSTRACT OF THE THESIS OF Carolyn E. Wright for the degree of Master of Science in Botany and Plant Pathologypresented on December 7, 1990. Title: A Systematic and Ecological Study of Astragalus diaphanus (Fabaceae) Redacted for Privacy Abstract approved: Kenton L. Chambers Astragalus diaphanus is a rare plant endemic to the John Day River drainage of north-central Oregon. This species has several interesting features, including the dimorphism of its fruit and its geographical isolation from the two nearest taxonomically related species, which occur in Colorado. This study addressed the species' distribution and habitat, the taxonomic relationships between the varieties of A. diaphanus, certain morphological comparisons among the species, possible reasons for the rarity of A. diaphanus, and the population biology of this taxon. Astragalus diaphanus was found to be more widespread in the John Day drainage than was previously known, but its range has shrunk due to habitat loss along the Columbia River. In this study, two varieties are recognized within a single species, based on striking morphological differences in pod forms which correspond to a break in geographical distribution. Other morphological characters are similar between the varieties. Flavonoid analysis and chromosome counts support this taxonomic treatment. Further study is needed to elucidate the relationships of A. diaphanus and its taxonomic relatives in Colorado. A low reproductive rate in A. diaphanus appears to be a potential problem, possibly contributing to its rarity. The species exhibits a combination of annual and biennial life- cycles. Many annual individuals of A. diaphanus perish without reproducing. This may be off-set by a large seed- bank, which is replenished sporadically by high production in robust biennials. There is not a need at the present time to provide legal protection to var. diaphanus, the more widespread and common taxon. However, var. diurnus has a very limited distribution along the South Fork of the John Day River and is represented by few populations. Many of these are marginally viable, and there are active threats to the existence of the latter variety. Astragalus diaphanus var. diurnus should be afforded legal protection as an endangered species. A Systematic and Ecological Study of Astragalus diaphanus (Fabaceae) by Carolyn E. Wright A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Master of Science Completed December 7, 1990 Commencement June 1991 APPROVED: (Th Redacted for Privacy Professor of Botany and Plant Pathology in charge of major Redacted for Privacy Head of department or Botany aryyrlantiPathology Redacted for Privacy Dean of Date thesis is presented December 7, 1990 Typed by researcher for Carolyn E. Wright ACKNOWLEDGEMENTS I would like to thank my committee members, Dr. K.L. Chambers, Dr. T.C. Moore, Dr. R. Frenkel, and Dr. D. Baisted, for their interest and careful reading of the thesis. Dr. Kenton L. Chambers provided funds for research and support. Fellow graduate students are always a source of friendship, inspiration, lively discussion and moral support. I'd like to thank Nancy Fredricks, Bob Meinke, Steve Shelly, Laura Morrison, Katie Marsden, Theresa Magee, Joe Antos, Mark Nelson, Tom Kaye, and Jacque Greenleaf. Credit is due to the following people for their contributions: Dr. John Miller assisted me with the flavonoid analysis. Betty Kerl conducted the soils analysis and Don Horneck assisted in interpreting the results. Dr. Paul Hammond identified butterflies, Dr. Paul Oman and Jim DiGiulio identified insects. Esther McEvoy illustrated seed pods. Curators at the following herbaria kindly loaned herbarium sheets for my use: Brigham Young University (BRY); California Academy of Sciences (CAS & DS); Intermountain Herbarium, Utah State University (UTC); New York Botanical Garden (NY); Oregon State University (OSC & WILLU); Rancho Santa Ana Botanic Garden (RSA & POM); Royal Botanic Gardens, Kew (K); University of California, Berkeley (UC); University of Colorado (COLO); University of Oregon (ORE); University of Washington (WTU); and Washington State University (WS). Finally, I would like to thank my husband, David Gross, for help with field work, photography, editing and much encouragement. TABLE OF CONTENTS INTRODUCTION 1 TAXONOMIC HISTORY 3 Overview of the Genus 3 Taxonomy and Synonymy of the Species 6 GEOGRAPHIC DISTRIBUTION 11 HABITAT OF ASTRAGALUS DIAPHANUS 17 Geology 17 Soils 22 Materials and Methods 22 Results and Discussion 22 Climate and Weather 27 Associated Species and Plant Communities 30 MORPHOLOGY 35 Comparative Species Descriptions 35 CHROMOSOMES 43 Materials and Methods 43 Results 43 Discussion 44 FLAVONOIDS 45 Materials and Methods 45 Structural Determination 48 Results 48 Discussion 54 PHYLOGENY 56 Primitive and Advanced Characteristics 56 Relationships of the Species 59 LIFE HISTORY OF ASTRAGALUS DIAPHANUS 60 Methods 60 First Season Phenology 61 Biennial Phenology 64 Reproductive Success 64 Herbivory, Insects, and Disease 67 Seed Germination and Greenhouse Studies 70 DISCUSSION AND CONCLUSIONS 73 Conservation 76 Conclusion 77 BIBLIOGRAPHY 78 APPENDICES 82 Collections Examined 82 Flavonoid Spectral Scans 88 LIST OF FIGURES Figure Page 1. Reported sites of Astragalus diaphanus. 15 2. Distribution of Columbia River basalts in the John Day River drainage. 19 3. Basalts near Service Creek and Johnson Creek. 21 4. Seedlings of Astragalus diaphanus in spring and fall with limited red coloration. 25 5. Astragalus diaphanus biennials showing conspicuous red coloration. 26 6. Pods of Astragalus diaphanus var. diaphanus. 37 7. Pods of Astragalus diaphanus var. diurnus. 39 8. Pods of Astragalus sparsiflorus var. sparsiflorus and var. majusculus. 40 9. Two-dimensional thin-layer chromatography spot maps of Astragalus diaphanus var. diaphanus. 49 10. Two-dimensional thin-layer chromatography spot maps of Astragalus diaphanus var. diurnus. 50 11. Annual phenology of Astragalus diaphanus. 62 12. Biennial phenology of Astragalus diaphanus. 65 LIST OF TABLES Table Page 1. Historical collections of Astragalus diaphanus. 11 2. Current locations of Astragalus diaphanus by 12 county in Oregon. 3. Selected soil characteristics at three sites of A. diaphanus. 23 4. Precipitation data for selected weather stations, 1981-1985. 28 5. Soil and air temperature measurements taken at three selected locations of Astragalus diaphanus. 29 6. Associated species of Astragalus diaphanus var. diaphanus and var. diurnus. 31 7. Two-dimensional thin-layer chromatography Rf values and spot colors for Astragalus diaphanus. 51 8. Flavonoids isolated from Astragalus diaphanus var. diaphanus and var. diurnus. 53 9. Spectral values obtained for flavonoids isolated from Astragalus diaphanus. 53 10. Primitive and advanced characters of subsect. Sparsiflori. 57 11. Site information and number of individuals tracked.61 12. Insects found with Astragalus diaphanus. 69 13. Germination rates of untreated seeds from selected sites of Astragalus diaphanus. 71 A SYSTEMATIC AND ECOLOGICAL STUDY OF ASTRAGALUS DIAPHANUS (FABACEAE) INTRODUCTION The subject of my study is the plant species Astragalus diaphanus Douglas ex Hooker, a member of the family Fabaceae, which is endemic to the John Day drainage of north-central Oregon. The species is of interest to taxonomists because of the unusual morphological variability of its fruits, which range in shape from sickle-like and flattened with three angles, to broadly rounded and inflated. Of further interest is the species' isolated geographical occurrence, its two nearest taxonomic relatives being found east and west of the main Rocky Mountain chain in Colorado. Astragalus diaphanus has been considered for possible listing as a threatened or endangered species for Oregon. The purpose of my study was to determine the natural range, habitats, and taxonomic relationships of the races differing in fruit shape within A. diaphanus, as well as to compare the taxon morphologically with its two related Colorado species, A. sparsiflorus and A. wetherillii. I also addressed possible reasons for the rarity of A. diaphanus and gathered preliminary data on its population biology. 2 The study involved field searches for populations of the species, descriptions of its habitats and associated species, and collection of specimens for morphological and flavonoid analyses. Plants were grown in the greenhouse from seeds collected in natural populations. Chromosome counts were made of three of the five taxa in the study. Field studies addressed the annual versus biennial life history strategy of the species, as well as some measures of reproductive success. Finally, threats to the future existence of the species were assessed, to help evaluate its rarity and the need for listing under state and federal endangered species laws. 3 TAXONOMIC HISTORY Overview of the Genus Astragalus L. is one of the largest plant genera in the world, represented by an estimated 1600 species (by some counts up to 3000 species)(Isely 1983). Principal centers of distribution include western North America, the Andes of South America, central Asia, Iran and Turkey (Barneby 1964). It is the largest genus of flowering plants in North America, with about 375 species (Barneby 1989). Within the Fabaceae (Leguminosae), the genus is placed in the subfamily Papilionoideae, the tribe Galegeae (Bronn) Torrey and Gray, and the subtribe Astragalinae (Adans.) Benth. (Polhill and Raven 1981). An excellent
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