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Systematic Studies of Non-Fistulose Delphinium Taxa Common To

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Systematic Studies of Non-Fistulose Delphinium Taxa Common To Systematic studies of non-fistulose Delphinium taxa common to Montana by Paul Thompson Sawyer A thesis submitted to the Graduate Faculty in partial fulfillment of the requirements for the degree DOCTOR OF PHILOSOPHY in Botany Montana State University © Copyright by Paul Thompson Sawyer (1970) Abstract: About 1,800 herbarium specimens representing five different taxa, Delphinium bicolor var. bicolor, D. bicolor var. pilosum, D. menziesii var. nuttallianum, D. menziesii var. utahense and D. depauperatum have been examined. Species descriptions for these five taxa are provided along with a key which separates the species of Delphinium growing in Montana. Chromatograms of leaf extracts from 37 populations representing the five taxa mentioned above have been made and interpreted. Evidence from the chromatographic study indicates that the populations called D. nuttallianum var. pilosum by Hitchcock (1964) and D andersonii by Sawyer (1967) are members of the species D. bicolor. Based on morphological and habitat differences this taxon is called D. bicolor var. pilosum. The chromatographic patterns for the three species of Delphinium mentioned above indicate a close relationship of these species but each one has a unique and distinct pattern. Chromatographic evidence of a D. bicolor x D. menziesii var. utahense hybrid is presented but D. bicolor is retained as a separate species because it is believed that this taxon is evolving throughout the major extent of its range isolated from the other delphiniums. Two taxa within the species D. menziesii, D. menziesii var. nuttallianum and D. menziesii var; utahense are given varietal rankings based on morphological and chromatographic evidence. SYSTEMATIC STUDIES OF NON-'FISTULOSE DELPHINIUM TAXA COMMON TO MONTANA by PAUL THOMPSON SAWYER A thesis submitted to the Graduate Faculty in partial fulfillment of the requirements for the degree of ; DOCTOR OF PHILOSOPHY in Bo tany Approved; MONTANA STATE UNIVERSITY Bozeman, Montana August, 1970 V iii ACKNOWLEDGMENT The author expresses his most sincere appreciation to Dr. W. E. Booth, his major professor, for Dr. Booth's guidance and.patience throughout this investigation. Dr. Booth, Dr. J. H. Rumely and Mr. H •N. Metcalf deserve much more than my thanks for the time and effort they spent constructively criticizing this thesis during its prepar­ ation. Similarly, the author is indebted to Mr. T. D. McGlynn, Assistant Professor of the Humanities and Social Sciences Department at Montana Tech. The author expresses his deep gratitude to Dr. G. A. Strobel for his advice concerning chromatographic techniques and for the use of his laboratory facilities. Use of chromatographic cabinets belonging to Dr. P. D. Skaar is also appreciated. Fellow graduate students, L. L. Bahls, L. F. Jonas and R. D. Dorn, and Mrs. Darlene Harpster have earned my thanks for the roles they played collecting Delphinium materials. Dr. J . R. Hodgson, also a fellow student, whose line drawings appear in this thesis, has earned a debt of gratitude. My appreciation is expressed to the herbarium curators of the following institutions: Brigham Young University, University of Idaho University of Montana, Oregon State University, Washington State -University, Utah State University and the Plant Research Institute at Ottawa, Canada, for the loan of their Delphinium specimens. iv The Department of Botany and Microbiology at Montana State University deserves credit for numerous kinds of support offered to the author. J V TABLE OF CONTENTS Page VITA. '.......... ii ACKNOWLEDGMENTS............................. iii, TABLE OF CONTENTS . ............................................ v LIST OF TABLES ........................ vii LIST OF FIGURES . ............. viii ABSTRACT ......................................................... ix INTRODUCTION............... I REVIEW OF LITERATURE ................... ..*..... 3 Classification ......... ................. ..... 3 Cytogenetics................ 7 Evolution ............................................. 7 Species Concepts Applied to Delphinium ........... 9 MATERIALS AND METHODS ............. 11 Herbarium S t u d y .......... .................. ........ 11 Chromatographic S t u d y .............. 11 TAXA DESCRIPTIONS AND K E Y S ........................... 15 Description of the Genus Delphinium L.». ................. 15 Key to the Species of Delphinium in Montana . ............. 16 Species Descriptions ................... ■........ 18 D , b i c o l o r ................ .18 vi Page D. bicolor var. pilosum . 26 .D.. depauperatum ....... 30 D . menziesii . ........ 35 D . menziesii var. nuttallianum . 36 D. menziesii var. utahense » . 40 RESULTS ...... ........... 48 DISCUSSION................... .. 58 SUMMARY ................... .......... 66 APPENDIX ......................... .. 69 LITERATURE CITED . ................ 82 vii LIST OF TABLES Page Table I. Names reported in the literature for five Delphinium taxa. Names appearing on the same horizontal line apply to the same taxon.. 6 IIt Delphinium material used in the chromato­ graphic study........ T, , ............. 12 III. Characteristics of components obtained by chromatographing leaf extracts from.Delphinium populations . ..... .. .. .. ...... .. •• ... 49 IV. Distribution of leaf extract components from Delphinium taxa. Compounds were- distinguished using a modified paper chromatography technique of Alston and Turner (1963). X indicates the presence of a particular spot on the chromato­ gram ..... ... ... .. ... ... 54 ■V. Paired affinity index values of the chromato­ graphic data for six taxa of Delphinium. A high value between two taxa indicates, a closer relationship of those taxa than a lower value . 57 viii LIST OF FIGURES Page Figure 1. Delphinium bicolor, var. bicolor ........... .. 22 2. Delphinium bicolor var. pilosum ................... 28 3. Delphinium depauperatum. ............... 33 .4. Delphinium menziesii var. nuttallianum ........ 39 . 5. Delphinium menziesii var.. utahense........... 44 6. Distribution maps for five taxa of Delphinium occurring in Montana ................. 47 7. Diagrammatic chromatograms typical of D. bicolor var. bicolor,. D . bicolor var. pilosum, D. menziesii var. utahense and D. menziesii var. nuttallianum...................... .. &1 8. Diagrammatic chromatograms typical of a D. bicolor x D. menziesii population from Ajax Mountain and D. depauperatum ......... 53 9. Plate of D. bicolor var. bicolor .. ., ,. .. ...... 71 10. Plate of D. bicolor var, pilosum ................... 73 11. Photograph of the hoIotype of D . nuttallianum yar.. pilosum ........... .......... 75 12. Plate of D menziesii var.. nuttallianum, ...... 77 13. Plate of D. menziesii var. utahense ........ 79 14. Plate of D . depauperatum 81 ix ABSTRACT About 1,800 herbarium specimens representing five different taxa, Delphinium bicolor var. bicolor, D. bicolor var. pilosum, D. menziesii var. nuttallianum, D^. menziesii var. utahense and D. depauperaturn have been examined. Species descriptions for these five taxa are provided along with a key which separates the species of Delphinium growing in Montana. Chromatograms of leaf extracts from 37 populations representing the five taxa mentioned above have been made and interpreted. Evidence from the chromatographic study indicates that the populations called D. nuttallianum var. pilosum by Hitchcock (1964) and I). andersonii by Sawyer (1967) are members of the species D. bicolor. Based on morphological and habitat differences this taxon is called D. bicolor var. pilosum. The chromatographic patterns for the three species of Delphinium mentioned above indicate a close relationship of these species but each one has a unique and distinct pattern. Chromatographic evidence of a D. bicolor x D. menziesii var. utahense hybrid is presented but ]). bicolor is retained as a separate species because it is believed that this taxon is evolving throughout the major extent of its range isolated from the other delphiniums. Two taxa within the species D. menziesii, D. menziesii var. nuttallianum and D. menziesii var; utahense are given varietal rank­ ings based on morphological and chromatographic evidence. INTRODUCTION Many morphological studies including those of Ewan (1945), Taylor (1960), Hitchcock (1964), Sawyer (1967) and Sutherland (1967) have led to various interpretations of certain taxa within the genus Delphinium. With the exception of preliminary breeding and transplant data provid­ ed by Sutherland (1967) there are no other systematic data supporting these interpretations. Five non-fistulose taxa of questionable status occur in Montana. Hence it seemed that investigations other than those of a morphological nature could be undertaken profitably to provide evidence which may lead to a better understanding of their relationships. Paper chromatographic studies involving flavonoid compounds have been used by taxonomists to investigate the following relationships: (I) interspecific hybrid populations such as those of Baptisia (Alston and Turner, 1963), Asplenium (Smith and Levin, 1963) and Vernonia (Hunter, 1967); (2) separation of species as in the case of Dicentra (Fahselt and Ownbey, 1968); (3) distinguishing diploid and tetraploid species of Tragopogon (Brehm and Ownbey, 1965); and (4) following the introgressing populations of Iris (Carter and Brehm, 1969). Although Sutherland (1967) reported that he was unable to separate species of Delphinium using Alston and Turner's (1963) paper chromatographic techniques he indicated that it might be useful
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