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University Microfilms International 300 N INFORMATION TO USERS This was produced from a copy of a document sent to us for microfilming. While the most advanced technological means to photograph and reproduce this document have been used, the quality is heavily dependent upon the quality of the material submitted. The following explanation of techniques is provided to help you understand markings or notations which may appear on this reproduction. 1.The sign or "target” for pages apparently lacking from the document photographed is "Missing Page(s)”. If it was possible to obtain the missing page(s) or section, they are spliced into the film along with adjacent pages. This may have necessitated cutting through an image and duplicating adjacent pages to assure you of complete continuity. 2. When an image on the film is obliterated with a round black mark it is an indication that the film inspector noticed either blurred copy because of movement during exposure, or duplicate copy. Unless we meaht to delete copyrighted materials that should not have been filmed, you will find a good image of the page in the adjacent frame. If copyrighted materials were deleted you will find a target note listing the pages in the adjacent frame. 3. When a map, drawing or chart, etc., is part of the material being photo­ graphed the photographer has followed a definite method in "sectioning” the material. It is customary to begin filming at the upper left hand corner of a large sheet and to continue from left to right in equal sections with small overlaps. If necessary, sectioning is continued again—beginning below the first row and continuing on until complete. 4. For any illustrations that cannot be reproduced satisfactorily by xerography, photographic prints can be purchased at additional cost and tipped into your xerographic copy. Requests can be made to our Dissertations Customer Services Department. 5. Some pages in any document may have indistinct print. In J l cases we have filmed the best available copy. University Microfilms International 300 N. ZEEB RD„ ANN ARBOR, Ml 48106 8214135 Roberts, Marvin Lee SYSTEMATIC STUDIES OF NORTH AMERICAN BIDENS SECTION BIDENS (COMPOSITAE) The Ohio State University Ph.D. 1982 University Microfilms International 300 N. Zeeb Road, Ann Arbor, M I 48106 Copyright 1982 by Roberts, Marvin Lee All Rights Reserved PLEASE NOTE: In all cases this material has been filmed in the best possible way from the available copy. Problems encountered with this document have been identified here with a check markV . 1. Glossy photographs or pages 2. Colored illustrations, paper or print,_____ 3. Photographs with dark background J A. Illustrations are poor copy_____ 5. Pages with black marks, not original copy_____ 6. Print shows through as there is text on both sides of page_____ 7. Indistinct, broken or small print on several pages_____ 8. Print exceeds margin requirements_____ 9. Tightly bound copy with print lost in spine_____ 10. Computer printout pages with indistinct print_____ 11. Page(s)__________ lacking when material received, and not available from school or author. 12. Page(s)__________ seem to be missing in numbering only as text follows. 13. Two pages numbered___________. Text follows. 14. Curling and wrinkled pages_____ 15. Other______________________________________________________________ __ University Microfilms International SYSTEMATIC STUDIES OF NORTH AMERICAN BIDENS SECTION BIDENS (COMPOSITAE) DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of the Ohio State University B y Marvin L. Roberts• B.S., M.S. The Ohio State University 1982 Reading Committeei Approved By Dr. Daniel J. Crawford Dr. Tod F. Stuessy Dr. Ralph E. Boerner otany ACKNOWLEDGMENTS t I wish to thank the numerous people who have assisted in the completion of this dissertation. Par­ ticular gratitude is extended to Dr. Daniel J . Crawford for continuous support and advice since the hegining of the project. Dr s. Tod F. Stuessy and Ralph E. Boemer, of my reading committee*deserve thanks for their patience and many criticisms which improved this dissertation. I also thank Dr. Ronald L. Stuckey for advice, information, and criticism regarding many aspects of this project. The line drawings were provided "by Ms. Cheryl Kimerline. Field studies were supported by National Science Foundation Doctoral Dissertation Improvement Grant DEB-78-20212. I wish to thank the curators of the herbaria who loaned specimens for this study (CU, DAO, F, GH, MTJB, NEBC, PH) and to others who have provided material. Special thanks are extended to Dr. A. E. Schuyler of PH for materials and information. VITA August 7. 1948 Born- Fostoria, Ohio 1971 B.S., The Ohio State University, Columbus, 1972-75 Assistant Curator, Department of Botany, The Ohio State University 1976-77 Graduate Research Associate, Department of Botany, The University of Wyoming, Laramie. 1977 M. S. in Botany, The University of Wyomings Laramie 1977 Graduate Teaching Assistant, Department of Botany, The Ohio State University, Columbus 1981 Visiting Lecturer, Department of Botany, Rutgers University, New Brunswick PUBLICATIONS Roberts, M. L. 1972. Butomus umbellatus in the Mississippi Watershed. Castanea 37« 83 -85”. Roberts, M. L. 1972. Wolffia in the bladders of Utriculariai an "herbivorous" plant? Mich. Bot. 11« 67-69. Roberts, M. L. & J. Youger. 1974. Aquatic and terrestrial plants of the Paint Creek Lake Project Area, pp. 14-8-193, 466-516. In C. E. Herdendorf and D. H. Stansberry, eds. Final Report Environmental Analysis of the Paint Creek Lak§ Project, Ohio. Department of Army, Corps of Engineers, Huntington District. • • • 111 Stuckey, R. L. & M. L. Roberts. 1974. Additions to the Bibliotheca Rafinesquiana. Taxon 23 * 365-372 . Stuckey, R. L. & M. L. Roberts. 1974. Bibliography of Theses and Dissertations on Ohio Floristics and Vegetation in Ohio Colleges and Universities.- Ohio Biol. Surv. Inform. Circ. No. 7. 92 pp. Roberts, M. L. 1976. IOPB Chromosome Number Reports LIII. Taxon 25* 484. Roberts, M. L. & D. J. Crawford. 1977* An elution chamber for the extraction of flavonoid compounds from paper chromatograms. Phytochem. Bull. 101 6-7. Stuckey, R. L. & M. L. Roberts. 1977- Rare and endangered aquatic vascular plants of Ohio* an annotated list of the imperiled species. Sida 7* 21-41. Roberts, M. L. 1980. The flavonoids of Meealodonta beckii (Compositae) and isolation of 2',3-dihydroxy-4- methoxy-4'-glucosyl chalcone. Biochem. Syst. & Ecol. 81 115-118. Roberts, M. L., R. L. Stuckey, and R. S. Mitchell, 1981. Hvdrocharis morsus-ranae (Hydrocharitaceae) 1 New to the United States. Rhodora 83 * 147-148. FIELD OF STUDY Major Field* Systematic Botany Studies in the systematics and phytochemistry of the genus Bidens (Compositae). Professor Daniel J. Crawford. TABLE OF CONTENTS Page ACKNOWLEDGMENTS.................................. ii VITA............................................... ill LIST OF TABLES................................... V LIST OF FIGURES....... vii CHAPTER I. THE IDENTITY OF BIDENS HETERODOXA AND B. INFIRMA. AND STATUS OF BIDENS SECTIONS PLATYCARPAEA AND HETERODONTA.............. 1 Introduction............................ 2 Materials and Methods................... 4 Morphology........... 5 Taxonomic History of Bidens heterodoxa... 11 Disposition of Bidens infirma and the varieties of Bidens heterodoxa......... 14 Status of Bidens sections Heterodonta and Platvcaroaea...................... 22 Conclusions........................ 24 Literature Cited......................... 31 II. FLAVONOIDS OF AMERICAN BIDENS SECTION BIDENS (COMPOSITAE)....................... 33 Introduction............................ 34 Materials and Methods................... 38 Results« Compound Identities............ 41 Results: Compound Destribution.......... 48 Discussion and Conclusions ........ 53 Literature Cited....... 84 v CHAPTER Page III. ALLOZYME VARIATION IN BIDENS DISCPIPEA. 87 Introduction. ....... ................... 88 Materials and Methods. .............. 90 Resultsi Morphology and Reproduction.... Results! Genetic Variation............ Discussion. .... ...................... 100 Conclusions............................. 103 Literature Cited. ........... 108 IV. VARIATION AMONG THE POPULATIONS OF BIDENS BIDENTPIPES (COMPOSITAEi COREOPSIDINAE).... 111 Introduction...................... 112 Methods and Materials............. 114 Results................. ........ 117 Discussion........................ 127 Literature Cited.................. 140 V. CYTOLOGY, BIOLOGY, AND SYSTEMATICS OF MEGALODONTA BECKII (COMPOSITAE)..... 142 Introduction......... 143 Materials and Methods. 144 Results and Discussion 145 Summary.............. References........... m APPENDIX A. POPULATIONS OF BIDENS EXAMINED FOR FLAVONOIDS............... 167 LIST OF TABLES Le CHAPTER I Page 1. Varieties of species of Bidens sections Platvcarpaea and Heterodonta (sensu Sherff, 1937) exhibiting extensive parallel awn variation.................................... 25 2. Distribution of disc corolla and correlated fruit characters in selected species of Bidens section Bidens..................... CHAPTER II 1. Taxonomy of Bidens section Bidens according to Sherff (1937, 1955. 1965). with explanatory notes. ...................... 57 2. Morphological characteristics of species groups in Bidens section Bidens. Cytolo'gical data compiled from published sources and my own determinations........... 58 3. Tentative identities of flavonoid glycosides detected in Bidens sect. Bidens*............ 59 Distribution
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