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INFORMATION TO USERS The most advanced technology has been used to photo graph and reproduce this manuscript from the microfilm master. UMI films the original text directly from the copy submitted. Thus, some dissertation copies are in typewriter face, while others may be from a computer printer. 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 copyrighted material had to be removed, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, charts) are re produced by sectioning the original, beginning at the upper left-hand comer and continuing from left to right in equal sections with small overlaps. Each oversize page is available as one exposure on a standard 35 mm slide or as a 17" x 23" black and white photographic print for an additional charge. Photographs included in the original manuscript have been reproduced xerographically in this copy. 35 mm slides or 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. • , % Accessing the WorldsUMI Information s 'ice 1938 300 North Zeeb Road Aru-Artie V 3 B '336 USA 8726732 THE SYSTEMATICS OF SIMSIA (COMPOSITAE HELIANTHEAE) Spooner, David Michael, Ph. The Ohio State University, 1987 UMI 300 N. Zeeb Rd. Ann Arbor. MI 48106 PLEASE NOTE. In ail 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 mark V 1. Glossy photographs or pages _______ 2. Colored illustrations, paper or print _ _ _ _ _ 3. Photographs with dark background S 4. Illustrations are poor co p y ________ 5. Pages with black marks, not original copy S 6. 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Other Best Copy Available, P ho toed As Received University Microfilms International THE SYSTEMATICS CF SIMSIA (OOMPOSITAE: HELIANTOEAE) DISSERTATION Presented in Partial Fulfillment of the Requirements for th e Degree of Doctor of Philosophy in the Graduate School of The Ohio State University By David Michael Spooner, B.A., M.S, * * * * * 1987 Dissertation Ccnmittee: Approved By: Dr. Daniel J. Crawford Dr. Gary Floyd Dr. Tbd F. Stuessy Dr. Thomas N. T aylor Depar o f Botany Copyright by David Hichael Spooner 1987 Dedicated to My fa n ily ACKNOWLEDGMENTS I gratefully acknowledge nunerous people for their help throughout this work. Tod F. Stuessy, my major professor, suggested the problem and has consistently provided help and encouragement throughout all phases of this work and my graduate study. Appreciation is also extended to ray cornuttee members, Daniel J. Crawford, Gary L. Floyd, and Thomas N. Taylor for their advice and critical reading of the manuscript. Nunerous friends and colleagues provided help in various ways. David M. Brandenburg, Eldward E. Schilling, and Charles M. Triplehorn all provided funds and one month each of their time in the field. Armando Burgos, Oscar R. Dorado, Jose Guevara B., and Jose L. Villasenor R. also accompanied and assisted me in many ways in Mexico or Colombia Wayne J. Elisens, Katherine L. Gross, Paul 0. Lewis, Donald H. Les, John J. Fur low, Ho-Yih Liu and Richard Whitkus provided statistical advice. Harold Robinson and John J. Strother provided valuable discussions on species lim its and generic problems in Simsia. Wayne J. Elisens and Stephen J. Wolf provided photographic assistance. Elton F. Paddock helped in the interpretation of meiotic configurations. Dale Johnson helped locate obscure literature references. Michael O. Dillon provided cytological material of Simsia dombeyana Peru. Others added in nunerous other ways, including valuable discussions, the loan of specimens, help with obtaining collecting permits, or welcoming me into their homes while I was in foreign countries. They include Mireyra D. Correa A., Arthur Cronguist, Santiago Diaz P., Luis D. Gomez P., Robert K. Jansen, David J. Keil, Stephen D. Koch, Thomas G. Lammers, Ho-Yih Liu, John F. Pruski, T. P. Ramanoorthy, Mario Sousa, B illie M. Turner, Lowell E. Urbatsch, and Thomas L. Wendt. David Dennis of the College of Biological Sciences drew the line drawings. The Ohio State University Instructional and Research Computer Center provided computer time. Grant support was provided by the National Science Foundation (BSR-8313268), The Society of Sigma Xi, The Ohio State University Graduate School (Alunni Research Award), The Ohio State University Department of Botany, and the American Society of Plant Taxonomists (Herbariun Travel Award). Appreciation is expressed to The Ohio State University for the award of a Presidential Fellowship which provided roe with an inencumbered final year to complete this work. I thank the curators of the following herbaria for the loan of specim ens: ARIZ, ASU, B, BAB, BM, BR, C, CAS, CGE, CHAPA, CM, COL, CR, DBS, DS, DUKE, DK£, F, F I, G, GH, QOET, U , IND, JE, K, L, LE, LIL, LL, LPB, M, MA, MASS, MEXU, MICH, MPU, MY, MC, CKF, P, PH, FMA, PCM, Q, QCA, RSA, S, S I, SMU, SR9C, TENN, TEX, U, UC, UPS, US, VEN, and W. Finally, I thank ray wife, Ann K. Spooner, without whose patience, support, and care of our fanily this work never would have taken p la c e . VITA November 1, 1949 ................................. Born - Downy, C a lifo rn ia 1971 ............................................................ B.S., Miami University, Oxford, Chio 1971-197 4 ................................................. U.S. Army Medical Field Service School, Ft. Sam Houston, Texas, Honor G raduate; Honorable Discharge, Army, 1974 1972-197 3 ................................................ University of Maryland, College Park, Maryland, 1972-1973 1974-1976 ..................................... ...M .S., Ohio University, Athens, Ohio 1977 .............................................................Instructor, Ohio University Continuing Education 1978-1980 ..............................................F ield Botanist, Ohio Department of Natural Resources, Division of Natural Areas and Preserves 1982-1987 ................................................. Graduate Teaching Associate and Graduate Research Associate, Botany Department, Hie Ohio State University, Columbus PUBLICATIONS Journal Articles Spooner, D. M. 1982. Wetlands in Teays-age valleys in extreme southeastern Ohio: Formation and flora. 89-99. IN: B. McDonald, ed. Proceedings of symposium on wetlands of the unglaciated Appalachian region, Morgantown, West V irginia, May 26-28, 1982. ________, and J.S. Shelly. 1983. Die national historical distribution of Platanthera peramoena (A. Gray) A. Gray (Orchidaceae) and its status in Ohio. Rhodora 84: 55-64. v _______ , A.W. C usick, B. A ndreas, and D.M. A nderson. 1983. N otes on Ohio vascular plants considered for listing as federally endangered or threatened species. Castanea 48t 247-255. _______ . 1983. Hie northern range of Phnradendron serotinum (Viscaceae), and its status in Ohio. Bull. Torrey Bot. Club. 110: 489-493. _______ . 1984. Reproductive features of Dentaria laciniata and E. diphylla (Cruciferae), and the implications in the taxonany of the eastern North American Dentaria complex. Amer. J. Bot. 71: 999-1005. _______ . 1984. Infraspecific variation in Gratiola viscidula Pennell (Scrophulariaceae). Rhodora 86: 79-87. _______ . 1984. Abstracts written for 50 Chio endangered or threatened species. IN: R.M. McCance and J.F. Bums (eds.). Chio Endangered and Threatened Vascular Plants. Ohio Department of Natural Resources, Colvmbcs. ________, A.W. Q jsic k , G.W. H a ll, and J.M . B askin. 1985. O b serv atio n s on the distribution and ecology of Sida hermaphrodite (L.) Rusby (Malvaceae). Sida 11: 215-225. Stuessy, T.F., D.M. Spooner, and K.A. Evans. 1986. Adaptive significance of neuter ray corollas in Helianthus grosseserratj ir (Ccrapositae). Amer. Midi. N aturalist 115: 191-197. ________, and D.M. Spooner. In P re s s . A daptive and p h y lo g e n e tic significance of paleae in the Conpositae. Taxon Spooner, D.M., T.F. Stuessy, D. J. Crawford, and M. Silva 0. In Press. Chromosome numbers fran the flora of the Juan Fernandez Islands II. Rhodora. Spooner, D.M. In Press. Simeia in “Flora de Nicaragua". Fieldiana, B ot. A b s tra c ts Spooner, D.M. 1976. Sexual and apcroictic reproduction in Dentaria laciniata (Cruciferae). Amer. J. Bot. 62 (Abetr.); 62. ________. 1981. D is tr ib u tio n a l s ta tu s o f some p re v io u s ly presumed rare Ohio plants. Ohio J. Sci. (Abetr.) 81: 20. ________, and J .S . S h e lly . 1982. The n a tio n a l h i s t o r i c a l distribution of Plan tan thera peramrwna (A. Gray) A. Gray vi {Orchidaceae) and its status in Ohio. Ohio J. Sci. (Abstr.) 82: I I . ________. 1982. Rare p la n ts and th e n a tu r a l a re a q u a lity o f w etlands along Teays-age valleys in southeastern Ohio. Chio J- Sci. (Abstr.) 82: 11. ________. 1983. Wetlands in Teays-age valleys in extrenx southeastern Ohio; formation and flora. Chio Biological Survey Informative Circular 11: 4. ________, and G.F. Hall. 1983. Sida hsrmaphrndifra- a cannon rarity. Chio J.