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A Systematic Study of Lathyrus Vestitus Nutt. Ex T

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A Systematic Study of Lathyrus Vestitus Nutt. Ex T AN ABSTRACT OF THE THESIS OF Steven Leo Broich for the degree of Doctor of Philosophy in Botany and Plant Pathologypresented on 4 March 1983 Title: A SYSTEMATIC STUDY OF LATHYRUS VESTITUS NUTT. EX T. & G. (FABACEAE) AND ALLIED SPECIES OF THE PACIFIC COAST Abstract approved: Redacted for Privacy Kenton L.Chdbers Eight species of Lathvrus (Fabaceae, Faboideae) endemic to the Pacific Coast of North America were the subject of a systematic investigation. Taxa studied included Lathvrus vestituq Nutt. ex T. & G., L. .1aetiflorus Greene, Lo iepsonii Greene, L. splendens Kellogg, L. polv-chvllus Nutt. ex T. & G., L. holochlorus (Piper) C. L. Hitchcock, L. delnorticus C. L. Hitchcock and L. sulphureus Brewer. Taximetric studies, including hierarchical and non-hierarchical cluster analyses followed by discriminant analyses, revealed the existence of one new species. Lathvrus holochlorus, Lo delnorticus, L. sulphureus, L. Polviohvllus, L. iepsonii and L. splendens were found to be morphologically distinct while extensive morphological intergradation was found between J. vestitus and L. laetiflorus. Principal Components analysis was used to document two clinal trends within the L. vestitus-laetiflorus complex. Flower size increases and floral shape changes from north to south along the Pacific Coast; pubescence increases clinally from north to south and from the coast inland. The anatomy of stems and leaflets was examined by tissue clearing methods. Vascular anatomy of nodes and internodes was found to conform to patterns described previously for European species of Lathvrus. Epidermal cell shape and stomatal frequency on leaflets varies greatly within species and these data were not useful for taxonomic distinctions. The diploid chromosome number for all species studied was found to be 2n = 14; only small differences in karyotypes could be noted. Preliminary studies of breeding systems in Lathvrus yestitus, L. Dolvphvllus and L. holochlorus revealed these perennial herba- ceous species to be self-incompatible, obligate outcrossers depen- dent primarily upon large bees for pollination. The new species, named Lathvrus alandulosus, is described; plants referable to Lo vestitus, and L. laetiflorus are reorganized into five subspecies under the former name.A key and descriptions for all species of Lathvrus, native and introduced, found west of the Sierra-Cascade Crest in Washington, Oregon and California is presented. A Systematic Study of Lathvrus vestitus Nutt. ex T. & G. (Fabaceae) and allied species of the Pacific Coast by Steven L. Broich A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy Completed 4 March 1983 Commencement June 1983 APPROVED: Redacted for Privacy Professor of Botany in charge of major Redacted for Privacy Chairman of Department of Botany and Plant Pathology Redacted for Privacy Dean of Gradu School (f Date thesis is presented Typed by the Author Steven Leo Broich To Jolene and Ann and Doug AO(NOWLEDGENE/4TS I wish to thank all who have helped make this research possible. Kenton Chambers suggested the problem and freely shared his knowledge and experience throughout all stages of the project. Its completion would not have been possible without this support. Taximetric studies were based partially on personal computer programs provided by David McIntire and J. A. Keniston. Additional help and advise was given by Eric Beals and especially by ArVydas Grybauskas. Mary Allcott, W. Roderick and especially James R. Griffin graciously responded to requests for seed and thereby provided valu- able germplasm for the cytogeneticstudies. Thanks also to the curators and directors of the herbaria who made large, long term loans of specimens essential to this study; to William Chilcote and Joe Antos for their livelyecological dis- cussions; to Andrew Moldenke for sharing his knowledgeof Lathyrus pollination mechanisms; to Thomas Nelson for a valuable orientation to northern California and the Lathyruq growing there; to Mark Nelson for Lathyrus collections from Curry County, Oregon; and to all graduate students, friends and acquaintances who have combined to make living in Oregon a richly rewarding scientific and cultural experience. I also wish to express my deep gratitude to Jolene: any success I might claim is really ours together. Funds for this project were provided by the Oregon State Univer- sity Herbarium and by the National Science Foundation. TABLE OF CONTENTS Page- I. INTRODUCTION 1 II. TAXONOMIC HISTORY 3 III. MATERIALS AND METHODS 7 IV. RESULTS Taxithetric Studies 15 Anatomical Studies 45 Karyotype Studies 47 Reproductive Biology 58 Ecology and Distribution 65 V. EVOLUTIONARY RELATIONSHIPS 75 VI. THE SPECIES OF LATHYRU OF THE PACIFIC COAST Introduction 80 A Key to the Species of Lathvruz of the Pacific Coast 82 Descriptions of Taxa 87 VII. BIBLIOGRAPHY 115 VIII. APPENDICES Introduction 121 Appendix A: The Specimens of LATH 122 Appendix B: The Specimens of LATHTKO 129 Appendix C: Additional Specimens Examined 136 LIST OF FIGURES Figure page, 1 Relationships among the North American taxa of Lathvrus as presented by C. L. Hitchcock (1952). 4 2 Diagramatic representation of flower measurements in- cluded in LATH and LATHTWO. 10 3 Cluster Analysis of LATHA: Single Linkage 17 4 Cluster Analysis of LATHA: Complete Linkage 17 5 Cluster Analysis of LATHA: UPGMA 18 6 Cluster Analysis of LATHB: Single Linkage 18 7 Cluster Analysis of LATHB: Complete Linkage 19 8 Cluster Analysis of LATHE: UPGMA 19 9 The ordination of specimens of LATH by scores of the first two canonical variables resulting from stepwise discriminant analysis. 27 10 Ordination of specimens of LATHA by scores of the first two canonical variables resulting from stepwise discrim- inant analysis. 29 11 Ordination of specimens of LATHE by scores of the first two canonical variables resulting from stepwise discrim- inant analysis. 29 12 Ordination of specimens of LATHTWO by scores of the first two canonical variables resulting from stepwise discriminant analysis. 35 13 Ordination of specimens in LATHTWO and an additional 12 specimens of L. splendens by scores of the first two canonical variables resulting from stepwise discriminant analysis. 37 14 Geographical distribution of scores of the first principal component in central California. 41 15 Geographical distribution of scores of the first principal component in southern California. 42 16 Geographical distribution of scores of the second principal component in central California. 43 List of Figures (Continued) Figure Page 17 Geographical distribution of scores of the second principal component in southern California. 44 18 Chromosomes of Pacific Coast T,athvru. 52 19 Chromosomes of Pacific Coast Lathvrus. 54 20 Geographic distribution of fertile and semi-sterile herbarium specimens sampled from the LATHTWO collection. 57 21 Dissection of a flower of 10. delnorticus C. L. Hitchc. illustrating the typical floral morphology for Pacific Coast Lathvrus. 59 22 Geographic distribution of the Pacific Coast Lathvrus allied to L. vestitus Nutt. ex T. & G. 67 23 Geographic distribution of Lo vestitus Nutt. ex T. & G. subspecies in southern California. 70 24 Geographic distribution of Lathvrus vestitus, L. lap- sonii and possible hybrid collections in central California. 71 25 Phenological comparison of L. vestitus and L. lepsonij, in the areas in which they are sympatric. 72 LIST OF TABLES Table Lag2. 1 Traits included in LATH and LATHTWO. 8 2 Means and standard deviations for all metric traits for the taxa in LATH. 22 3 Initial F-values and traits selected in the stepwise discriminant analyses of LATH, LATHA and LATHB. 24 4 Mean Mahalanobis'D2 values within taxa and the D2 distances between taxa from the discriminant analysis of LATH. 25 5 Means and standard deviations for all traits of the groups hypothesized to exist in LATHTWO. 32 6 Mean Mahalanobis' D2 values within taxa and D2 distances between taxa from the discriminant analysis of LATHTWO. 33 7 Data for the first three eigenvectors resulting from a principal components analysis of the correlation matrix from LATHTWO. 39 8 A summary of literature concerning chromosome counts for species of Lathyrus endemic to North America. 48 9 A catalogue of chromosome counts made in the present study. 50 10Mean seed weight in mg, standard deviations and ranges for seed collections of Pacific Coast Lathyrus species. 64 A SYSTEMATIC STUDY OF LATHYRUS VESTITUS NUTT. ex T. & G. (FABACEAE) AND ALLIED SPECIES OF THE PACIFIC COAST I. INTRODUCTION Lathvrus L. is a genus of approximatly 150 species of temperate papilionoid legumes (Fabaceae, Faboideae). It is a member of the tribe Vicieae (Adans.) DC. which also includes Vicia (ca. 140 species), Lens Mill. (5 species, including the domesticated lentil), Pisum L. (2 species, including the domesticated pea) and Vavilovia A. Fedorov (monotypic) (Kupicha 1977, 1981).No clear distinction exists between LathvruQ and Vicia; transitional species, found in Lathvrus sect. Orobus (Hassler 1973) and Vicia sect. Vicilla (Kupicha 1976), can be separated only on the basis of certain arbitrary character- istics (Kupicha 1981). The center of diversity for both Lathvrus and Vicia is in the Mediterranean and Irano-turanian region (Kupicha 1981). A complete subgeneric taxonomy of Lathyrus has never been pub- lished. European species were organized into sections by Acherson and Graebner (1910) and variations on this scheme have been used since then (Fedchenko 1948; Davis 1970; Czfranova1971).
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