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Calochortus Howellii: Ecoloav of a Rare Serpentine

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Calochortus Howellii: Ecoloav of a Rare Serpentine AN ABSTRACT OF THE THESIS OF Nancy Ann Fredricks for the degree ofMaster of Science in Botany and Plant Pathology presented on April 28. 1986 Title: Calochortus howellii: Ecoloav of a Rare Serpentine Endemic. and Comparison with the New Species. C. umPouaensis (Liliaceae). Redacted for Privacy Abstract approved: Kenton L. Chambers The habitats and population ecology of a rare plant species of southwest Oregon, Calochortus bowellii Watson (Liliaceae) are characterized and a related new species, C. umpquaensis Fredricks, is described from a very limited area in Douglas County, Oregon. These two species differ in the size, pigmentation, and pubescence of their petals, size and position of their capsules, and shape and testa morphology of their seeds. Based on preliminary flavonoid analysis and chromosome karyotypes, C. umpquaensis is placed with C. howellii in the subsection Nitidi of the section Eucalochortus. Morphological similarities unique to the species pair suggest a close relationship between C. howellii and C. umpquaensis. Calochartus umpquaensis typically occurs on serpentine- derived soils of grassland-forest ecotones. Its known distribution is limited to several sites within an area less than 5 km2 along the Little River, a tributary of the North Umpqua River. Because of its extremely restricted range and vulnerability to extinction, measures are recommended to assure its protection. calochortus howellii is endemic to the upper Illinois River area of Josephine County, Oregon. It occurs on serpentine-derived soils high in nickel and chromium, with low calcium to magnesium ratios. Most populations are located on southeast to southwest facing slopes on benches within an elevational range of 350 to 540 m. Seed germination rates in the laboratory after storage at 5 C were near 100 percent. Based on the high incidence of insect visitation and high seed set, pollination did not appear to limit reproductive output of the species. Floral patterns detectable under ultraviolet light may serve to attract and guide potential pollinators. Insect visitors were observed and identified. Predispersal seed predation, losses from the seed bank, and high juvenile mortality may contribute to low recruitment in natural populations. A long-term demographic study of Calochortus howellii was initiated in 1983. Data were collected for 375 individuals over three field seasons, 1983-1985. Of those plants tagged, 67 percent bloomed at least once during the three year period. Only 8 percent flowered in all three years. The highest incidence of flowering occurred in 1983, corresponding to the year of highest precipitation. Due to bulb dormancy, more years of data are necessary to ascertain population density and to formulate a population model that reflects the population accurately. Hypotheses of flowering periodicity and effects of precipitation on flowering and breaking of dormancy are proposed for future testing. Calochortus howellii: Ecology of a Rare Serpentine Endemic, and Comparison with the New Species, Q. umpquaensis (Liliaceae) by Nancy Ann Fredricks A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Master of Science Completed April 28, 1986 Commencement June 1986 APPROVED: Redacted for Privacy Professor of Botany in Charge of Major Redacted for Privacy Chairman of the Department of Botany and Plant Pathology Redacted for Privacy Dean of Grad SchoolG Date thesis is presented April 211_ 1986 Typed by Nancy Ann Fredricks ACKNOWLEDGEMENTS I would first like to express my deep appreciation to Dr. Kenton Chambers for his support, guidance, and encouragement throughout this project. I am also grateful to the other members of my committee, Dr. Robert Frenkel, Dr. Ed Guerrant, and Dr. David Hibbs for their constructive comments and careful reading of the manuscripts. Special thanks are extended to others who also reviewed manuscripts in various stages of completion. Comments and suggestions from Dr. Joe Antos, Dr. Larry Hufford, Teresa Magee and Dr. John Miller were most valuable. Additional discussions with fellow graduate students including Mark Chatfield, Dr. Dave Dalton, Dee Lusk, Katie Marsden, Bob Meinke, Steve Shelly, and Carolyn Wright provided insights into various aspects of the study. Others who have given of their time include Russ Holmes, BLM Botanist for the Roseburg District, who spent a day in the field and provided new localities of C.. umpquaensis. Bob Meinke, Steve Shelly, Dr. John Miller, and Edgar and Ruth Ross collected material for flavonoid study. Gary Fredricks assisted with field work and photography. Dr. Jadwiga Mole-Bajer translated a pertinent review paper. Cary Shufelt provided computer assistance. Dr. Don Armstrong permitted use of his laboratory for a number of experiments and Mark Chatfield and Katie Marsden provided technical assistance and a physiological perspective. Curators of the following herbaria kindly provided loan specimens: Museum of Natural History, University of Oregon (ORE); California Academy of Sciences/Dudley Herbarium (CAS); University of California, Berkeley (UC); Humboldt State University (HSU); Rancho Santa Ana Botanic Garden (RSA), Marion Ownbey Herbarium, Washington State University (WS); University of Washington Herbarium (WTU); New York Botanical Garden (NY); and Missouri Botanical Garden Herbarium (MO). Appreciation is extended to Dr. Thomas Moore and the Department of Botany for financial support through a teaching assistantship. Support, research, and travel funds were provided by the Herbarium at Oregon State University through Dr. Kenton Chambers. Finally, I would like to acknowledge the support and encouragement of my family and friends. Most of all, I thank Gary Fredricks for his unfailing support, patience, and confidence in me throughout this project and the past nine years. TABLE OF CONTENTS Chapter Page I. General Introduction 1 II. Morphological Comparison of calochortus 7 bowellii and a New Species from Southwestern Oregon, Q. umpquaensis Abstract 7 Introduction 8 Materials and Methods 10 Collection and Analysis of Morphological Data 10 Chromosome Karyotypes 11 Flavonoid Analysis 12 Results and Discussion 13 Morphological Comparison 13 Seeds 13 Seedlings 22 Mature Plants 23 Chromosome Karyotypes 36 Flavonoid Analysis 39 Description of Q. umpquaensis 45 Discussion of the "Type Specimen" Problem of Q. howellii 46 Conclusions 49 Contributions 51 Literature Cited 52 III. The Ecology of Calochortushowellii 54 Abstract 54 Introduction 55 Materials and Methods 59 Field Methods 60 Associated Species 60 Soils 60 Insect Visitors 63 Seed Set 64 Germination 64 Results 64 Associated Species 64 Soil Analysis 67 Pollination 69 Predation 73 Seed Set 74 Germination 74 Discussion 74 Conclusions 83 Literature Cited 85 TABLE OF CONTENTS (continued) IV. Population Dynamics of Calochortug howellii 89 Abstract 89 Introduction 90 Materials and Methods 93 Meteorological Data 93 Collection of Field Data 94 Analysis of Field Data 94 Results 95 Meterological Data 95 Vegetative Emergence 96 Flowering Frequency 100 Morphometric Analysis 100 Discussion 105 Effect of Precipitation on Flowering, Emergence and Phenology 105 Morphometrics of the Population and the Individual 107 Evidence for Bulb Dormancy 108 Conclusions and Future Proposals 110 Literature Cited 113 Bibliography 114 Appendix 120 LIST OF FIGURES Figure eagg. 1 Distribution of C. howellii and C. 9 umpquaensis with serpentine areas delineated. 2 Scanning electron micrographs of C. 16 howellii seeds at 22% and C. umpquaensis seeds at 30X. 3 Scanning electron micrographs of distal 18 and proximal ends of seed at 140X. 4 Scanning electron micrographs of C. 20 howellii (22%) and C. umpquaensis (30X) seeds, and of seed surfaces near median ridge at 150X. 5 Photographs of flowers of C. umpquaensis 24 and C. howellii at lx. 6 Photographs of petals of C. umpquaensis 26 at 1% and C. howellii at 1.5X. 7 Scanning electron micrographs of petal 29 glands of C. howellii and C. umpquaensis at 20X and 551. 8 Scanning electron micrographs of 31 branched petal gland processes, trichomes and papillae at 150X and 55X. 9 Petals of C. howellii photographed 33 under natural and ultraviolet light. 10 Karyotypes of C. umpquaensis and C. 38 howellii. 11 Two-dimensional chromatograms from 41 flavonoid analysis of eight species of Calochortus. 12 Distribution of C. howellii. 62 13 Two-dimensional polar ordination of sites 66 using Sorensen's index of similarity. 14 Insect visitors to C. howellii. 71 15 Frequency histogram of leaf width of C. 104 howellii individuals emerging in plots in 1985. LIST OF TABLES Table Page 1 Subgeneric classification of 2 Calochortus after Ownbey (1940). 2 Comparison of morphological features 15 of C. howellii and C_umpquaensis. 3 Average Fir values from two-dimension- 40 al TLC of petal extracts from eight calochortus species. 4 Verified localities of C. howellii. 61 5 Sorensen similarity matrix of asso- 65 ciated species of Q. howellii from five locations. 6 Results of soil analysis for six 68 sites in southwestern Oregon. 7 Insect visitors to Q. howellii and 70 percent occupancy of flowers by each species on Mariposa Meadow in July 1983. 8 Numbers of seed and unfilled ovules 75 per capsule of Q. howellii collected at Selma site. 9 Seasonal and monthly precipitation 97 at Cave Junction Weather Station. 10 Density of tagged individuals on 98 Mariposa Meadow. 11 Numbers of individuals of C. howellii 99 per plot. 12 Numbers of tagged individuals bloom- 101 ing during 1983, 1984, and
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