INFORMATION TO USERS This manuscript has been reproduced from the microfilm master. UMI filTTiR the text directfy from the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter face, while others may be firom any type of computer printer. H ie 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 inq)roper alignment can adversefy afiect 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 comer and continuing fiom 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. Photogr^hs 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 aiy photographs or illustrations appearing in this copy for an additional charge. Contact UMI directly to order. UMI A Bell & Howell information Company 300 North Zeeb Road. Ann Arbor. Ml 48106-1346 USA 3l3.'761-4700 800/521-0600 PHYLOGENY AND BIOGEOGRAPHY OF PAEONIA (PAEONIACEAE) DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Tao Sang, B.S., M.S. ***** The Ohio State University 1995 Dissertation Committee: Approved b; Daniel J. Crawford Keith R. Davis Tod F. Stuessy Advlso: Department of Plant Biology DMI Number: 9544679 UMI Microform 9544679 Copyright 1995, by DMI Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code. UMI 300 North Zeeb Road Ann Arbor, MI 48103 To My Father and Wanxiao 11 ACKNOWLEDGEMENTS The completion of this project would not have been possible without the generous assistance of numerous people. I wish to express my sincere appreciation to my major professor, Tod F. Stuessy, for his invaluable advice, constant encouragement, and continuous support throughout all phases of my graduate study. Sincere appreciation is also expressed to Dr. Daniel J. Crawford for his generous help and invaluable guidance in all aspects of my research and study at OSU. I am deeply indebted to Dr. G. Ledyard Stebbins for his support and encouragement of the project as well as helpful discussions. I thank Professor De-yuan Hong for his continuous support, stimulating suggestions, and generous help in many aspects of this project. I thank Dr. Keith Davis for serving in my advisory, general exam, and dissertation committee, and for his valuable comments on my dissertation draft. Numerous friends and colleagues provided help and assistance during my field trips to California, Bulgaria, China, Greece, and Spain. Special thanks go to Abdeslam E. Aallali, David J. Keil, Xue-yi Li, Joaguin M. Mesa, Kaiyu Pan, Yianlong Pei, D. Petkova, Tim S. Ross, Dagui Tang, Dimitris Tzandoudakis, and Dessi Usunov. I thank Bruce iii Bartholomew, G. L. Osti, Loren H. Rieseberg, and Nigel Rowland for providing plant materials; and the Royal Botanic Gardens, Kew, for allowing collection of leaf materials. Thanks also go to my colleagues and friends, Jorge Arriagada, Erica Armstrong, Marybeth Cosner, Melanie Devore, Betsy Esselman, Karla Gengler, Seung-Chul Kim, Hongqi Li, Ming Yang, Xuanli Yao, James Zech, and Liming Zhao for their help and useful discussions. I am grateful to Mr. Roy Klehm (Klehm Nursery, South Barrington, Illinois) for his generous financial support of the field work. This research was also supported by a National Science Foundation Doctoral Dissertation Improvement Grant DEB-9321616 to T.F.S. and T.S., a Sigma Xi Grant-in-Aid of Research, a Graduate Student Research Grant from American Society of Plant Taxonomists, two Janice Beatley Herbarium (OS) Awards, and a Graduate Student Alumni Research Award from the Ohio State University. The curators of the following herbaria are acknowledged for allowing study of specimens, either through loans or during visits: ATM, GH, GRA, K, KUN, NY, PE, SO, SOM, SZ. UC, UFA, US, and WUK. Finally, I give my most special thanks to my wife, Wanxiao, for her love, support, understanding, and help during four years of my graduate study. IV VITA January 19, 1966 ........ Born - Sichuan, P. R. China 1986 .................... B.S., Botany, Fudan University, Shanghai, P. R. China 1989 .................... M.S., Botany, Fudan University, Shanghai, P. R. China 1990-91 ................. Teaching Assistant, Department of Botany, Ohio University, Athens, Ohio 1991-94 ................. Graduate Teaching/Research Associate, Department of Plant Biology, The Ohio State University, Columbus, Ohio 1994-95 ................. Presidential Fellow, The Ohio State University, Columbus, Ohio PUBLICATIONS Yu, C. -Y., M. Zhang, P. -S. Hsu, and T. Sang. 1990. Cladistic analysis of Populus. Bulletin of Botanical Research 10; 69-76. Abu-Asab, M, S., P. D. Cantino, J. W. Nowiche, and T. Sang. 1993. Systematic implications of pollen morphology in Caryopteris (Labiatae). Systematic Botany 18: 502-515. Sang, T., D. J. Crawford, S. -C. Kim, and T. F. Stuessy. 1994. Radiation of the endemic genus Dendroseris (Asteraceae) on the Juan Fernandez Island: evidence from sequence of the ITS regions of nuclear ribosomal DNA. American Journal of Botany 81: 1494-1501. Sang, T. 1995. New measurements of distribution of homoplasy and reliability of parsimonious cladograms. Taxon 44: 77-82. v Sang, T., D. J. Crawford, and T. F. Stuessy. 1995. Documentation of reticulate evolution in peonies (Paeonia) using internal transcribed spacers sequences of nuclear ribosomal DNA: implications for biogeography and concerted evolution. Proceeding of National Academy of Sciences, USA 92: 6813-6817. Sang, T., D. J. Crawford, T. F. Stuessy, and M. S. Silva O. 1995. ITS sequences and the origin and evolution of the genus Robinsonia (Asteraceae) on the Juan Fernandez Island. Systematic Botany 20: 55-64. Abstracts Presented To Meetings Hsu, P. -S. and T. Sang. 1988. What is the exact meaning of species? In "Abstract of International Symposium on Botanical Gardens", pp.10. Nanjing: Nanjing Botanical Garden. Sang, T. 1992. Cytology, cytogenetics, and phylogeny of Paeonia (Paeoniaceae): A preliminary report. The Ohio Journal of Science 92: 13. Abstract. Sang, T. 1993. Homoplasy distribution ratio: A new measurement of reliability of parsimonious cladograms based on levels and distribution of homoplasy. American Journal of Botany 80: 174-175. Abstract. Stuessy, T. F. and T. Sang. 1993. Phylogeny of Barnadesioideae and early evolution of Compositae. The Ohio Journal of Science 93: 41. Abstract. Crawford, D. J., T. Sang, S. -C. Kim, and T. F. Stuessy. 1994. Evolution and origin of the endemic genus Dendroseris (Asteraceae) on the Juan Fernandez Islands: evidence from nuclear and chloroplast DNA. International Compositae Conference, Royal Botanic Gardens, Kew: 18. Abstract. Lee, N. S., T. Sang, and D. J. Crawford. 1994. ITS sequences divergence between disjunct taxa in eastern Asia and North America. American Journal of Botany 81: 167. Abstract. Sang, T. 1994. A field study of peony species in western China. The Ohio Journal of Science 94: 21. Abstract. Sang, T., D. J. Crawford, S. -C. Kim, and T. F. Stuessy. 1994. Radiation of the endemic genus Dendroseris (Asteraceae) on the Juan Fernandez Islands: evidence from sequence of the internal transcribed spacer (ITS) region of the nuclear ribosomal DNA (nrDNA). American Journal of Botany 81: 184. Abstract. vi Sang, T., D. J. Crawford, and T. F. Stuessy. 1994. Phylogeny of Paeonia (Paeoniaceae) inferred from ITS sequences and biogeographic implication. American Journal of Botany 81: 183. Abstract. Sang, T., D. J. Crawford, T. F. Stuessy, and M. S. Silva O. 1994. ITS sequences and the origin and evolution of the genus Robinsonia (Asteraceae) on the Juan Fernandez Island. American Journal of Botany 81: 183-184. Abstract. Stuessy, T. F., D. J. Crawford, T. Sang, and R. Rodriguez. 1994. Morphological and molecular divergence between the two species of Rhaphithamnus (Verbenaceae). American Journal of Botany 81: 190. Abstract. J Stuessy, T. F., T. Sang, and M. DeVore. 1994. Phylogeny and Biogeography of Barnadesioideae (Compositae). American Journal of Botany 81: 190-191. Abstract. Stuessy, T. F., T. Sang, and M. DeVore. 1994. The Phylogeny and Biogeography of Barnadesioideae (Compositae). International Compositae Conference, Royal Botanic Gardens, Kew: 25. Abstract. Sang, T. 1995. Evolution of peonie (Paeonia) in the Mediterranean region. The Ohio Journal of Science 95: 38. Abstract. Sang, T., D. J. Crawford, and T. F. Stuessy. 1995. Documentation of reticulate evolution in Paeonia (Paeoniaceae): implications for concerted evolution and biogeography. American Journal of Botany 82: 159- 160. Abstract. Sang, T., D. J. Crawford, and T. F. Stuessy. 1995. Comparison of chloroplast matK gene and nrDNA ITS region provides new insights into the phylogeny of Paeonia section Paeonia and molecular evolution. American Journal of Botany 82: 159. FIELDS OF STUDY Major field: Plant Biology Studies in Plant Evolution and Systematics