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Information to Users INFORMATION TO USERS This manuscript has been reproduced from the microfilm master. UMI films the text directly from the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter face, while others may be from any type of computer printer. The quality of this reproduction is dependent upon the quality of the copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandard margins, and improper alignment can adversely affect reproduction. In the unlikely event that the author did not send UMI a complete manuscript and there are missing pages, these will be noted. Also, if unauthorized copyright material had to be removed, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, charts) are reproduced by sectioning the original, beginning at the upper left-hand corner and continuing from left to right in equal sections with small overlaps. Each original is also photographed in one exposure and is included in reduced form at the back of the book. Photographs included in the original manuscript have been reproduced xerographically in this copy. Higher quality 6" x 9" black and white photographic prints are available for any photographs or illustrations appearing in this copy for an additional charge. Contact UMI directly to order. University Microfilms International A Bell & Howell Information Company 300 North Zeeb Road. Ann Arbor. Ml 48106-1346 USA 313/761-4700 800/521-0600 Order Number 9505151 Systematics of Cltbadium L. (Asteraceae, Heliantheae) Arriagada, Jorge Eugenio, Ph.D. The Ohio State University, 1994 UMI 300 N. ZeebRd. Ann Arbor, MI 48106 SYSTEMATICS OF CLIBADIUM L. (ASTERACEAE, HELIANTHEAE) DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy in the Graduate School of The Ohio State University By Jorge E . Arriagada ****** The Ohio State University 1994 Dissertation Committee Approved by Ralph E. J. Boerner Daniel J. Crawford Tod F. Stuessy Advisor^ Department of Plkfrt Biology Dedicated to my wife Violeta and my daughters Paula and Andrea ii ACKNOWLEDGMENTS Numerous people must be acknowledged for their assistance in helping me complete this dissertation. First and foremost I would like to thank my advisor, Professor Tod F. Stuessy, for his invaluable help and guidance, constant encouragement and continuous support throught all phases of my graduate study. Appreciation is also owed to Professor Daniel Crawford for his suggestions on the final version of this thesis and for his interest in and support of all aspects of my professional development, and to Professor Ralph Boerner, chairman of the Department of Plant Biology, for his advice and critical review of the manuscript. Many institutions have been instrumental in helping make my dissertation successful. Financial support for field trips was provided by the National Science Foundation (INT- 9017 695), The Garden Club of America, The Tinker Foundation, The Organization of American States, The Department of Plant Biology, OSU, and The Ohio State University Janice C. Beatley Herbarium Award. Numerous friends and colleges have provided help and assistance during my field trips. Special thanks go to Peter Dobbeler for sharing with me his expertise on Costa Rican vegetation. I thank Santiago Diaz-Piedrahita, an outstanding Colombian synantherologist, who shared with me his botanical and historical expertise on Colombian Compositae. I also thank Jaime Aguirre, Primo Chavarria, Jorge Gomez Laurito, Hazel Hernandez, Luis J. Poveda, Jorge Rojas, Pablo Sanchez Vindas, Fausto Sarmiento, Cristina Velez-Nauer, and other people too numerous to mention. Special thanks go to Dr. Nicholas Hind, from the Royal Botanic Gardens, Kew, for his important comments and suggestions on my research, to Jose L. Panero, from MSC, for sharing with me important information concerning Andean Heliantheae, and to Javier Francisco- Ortega, for his valuable comments on nomenclatural problems. Discussions with other persons of the plant systematics program at The Ohio State University have also contributed to this study, including Karla Gengler, Paul 0. Lewis, Tao Sang, Jun Wen, and James Zech. Special thanks go to Dr. John Furlow (OS) for loan assistance. I also thank Dr. H. W. Lack for important information regarding types at B, and Hugo Valdebenito and Fausto Sarmiento for obtaining valuable information about types at QPSL. I also thank the curators of the following herbaria from which specimens were borrowed for this study, and who provided, in some cases, space and valuable information during my visits: A, B, BM, CAS, COL, CR, DS, ECON, F, FSU, G, GH, K, LL, LP, MEDEL, MO, NY, P, UC, US, WIS. Finally, I express sincere and special thanks to my wife, Violeta, for supporting me unconditionally during all the years spent as a graduate student. iv VITA November 26, 1949........ Born Santiago, Chile. 1977 ...................... Licenciado en Biologxa, Universidad de Concepcion, Concepcion,Chile. 1977-1988................ Instructor, Dept.of Botany, Universidad de Concepcion, Chile. 1987 ...................... M.S., Universidad de Concepcion, Concepcion, Chile. 198 8 ...................... Research Assistant, The Ohio State University Herbarium. 1989-1991.................Teaching Associate, The Ohio State University, Columbus, Ohio. 1991-199 2 .................Lecturer, The Ohio State University, Columbus, Ohio. 1992-199 4 ................. Lecturer and Laboratory Instructor, The Ohio State University - Newark Campus. v PUBLICATIONS Stuessy, T.F. and J. Arriagada. 1989. Systematics of Desmanthodium (Compositae, Heliantheae). Amer. J. Bot. 76(6): 273. [Abstract]. Stuessy, T.F. and J. Arriagada. 1989. Systematics of Desmanthodium (Compositae, Heliantheae) . Ohio Acad.. Sci. 89(2): 7. [Abstract]. Arriagada, J.E. 1990. Chromosome studies in Clibadium (Compositae, Heliantheae). Ohio Acad. Sci. 90(2): 5. [Abstract]. Arriagada, J.E. Preliminary studies and infrageneric classification in the genus Clibadium (Compositae, Heliantheae). Ohio Acad. Sci. 90(2): 5. [Abstract]. Arriagada, J. and T.F. Stuessy. 1990. A new species and subgenus of Desmanthodium (Compositae, Heliantheae) from southern Mexico. Brittonia 42: 283-285. Dxaz-Piedrahita, S. and J. Arriagada. 1992. Una nueva especie de Clibadium (Compositae, Heliantheae) para Colombia. Rev. Acad. Colomb. Cienc. 18: 301-304. Stuessy, T.F. and J. Arriagada. 1993. Chromosome counts in Clibadium (Compositae, Heliantheae) from Latin America. Brittonia 45: 17 2-176. Arriagada, J. 1993. Notas sobre las relaciones genericas de Clibadium (Compositae, Heliantheae). Rev. Acad. Colomb. Cienc. 18: 465-468. vi Arriagada, J. 1994. Revision of the genus Clibadium (Compositae, Heliantheae). Ohio Acad. Sci. 94(2):2l. [Abstract] . Arriagada, J. 1994. Systematics of Clibadium (Compositae, Heliantheae). Amer. J. Bot. 81: 140. [Abstract]. FIELD OF STUDY Major field: Plant Biology Studies in Plant Systematics with Professor Tod F. Stuessy TABLE OF CONTENTS DEDICATION.............................................. ii ACKNOWLEDGEMENTS....................................... iii VITA.................................................... v LIST OF TABLES......................................... X LIST OF FIGURES....................................... xi INTRODUCTION............................................ 1 TAXONOMIC HISTORY...................................... 5 GENERIC AFFINITIES .................................... 12 CONCEPTS OF SPECIES AND VARIETIES..................... 28 CHROMOSOME NUMBERS..................................... 31 HYBRIDIZATION......... 34 HABITAT AND DISTRIBUTION ...................... 36 PLANT-INSECT INTERACTIONS.............................. 38 ETHNOBOTANY........... 42 PHYLOGENETIC ANALYSIS.................................. 48 MODES OF SPECIATION.................................... 58 MORPHOLOGY AND TAXONOMIC CRITERIA..................... 64 TAXONOMY................................................ 7 6 EXCLUDED AND DOUBTFUL NAMES........................... 235 APPENDICES A. Numerical list of species................... 237 B. Index to numbered collections examined...... 239 viii C. Notas sobre las relaciones genericas de Clibadium L. (Compositae,Heliantheae) ...... 260 D. Chromosome counts in Clibadium (Compositae, Heliantheae) from Latin America............. 265 E. Scatter diagram of representative collections of Clibadium grande. C. grandifolium, and C. pacificum................................ 271 LIST OF REFERENCES..................................... 273 ix LIST OF TABLES TABLE PAGE 1 . Characters and character states used in cladistic analysis of Clibadium and related genera. Characters 3, 4, 8, 12, 14, 15, 18, and 22 are ordered, but the rest of the multistate characters are unordered.................................... 19 2 . Character state matrix for phylogenetic reconstruction of Clibadium and related genera. See Table 1 for description of characters and states....................... 22 3. Chromosome numbers of species of Clibadium.. 32 All u = 16. 4. Characters and character states used in cladistic analysis of species of Clibadium. Characters 4, 5, 7, 13, and 15 are ordered; the rest of multistate characters are unordered.................... 50 Character state matrix for phylogenetic reconstruction of species of Clibadium. See Table 4 for description of characters and states.................... 51 x LIST OF FIGURES FIGURE PAGE 1. Distribution of Clibadium and related genera.
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