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 Aroor Ml -18106-1346 USA 313 761-4700 800 521 0600 Order Number 9120T36 Systematics ofAralxa (Araliaceae) Wen, Jun, Ph.D. The Ohio State University, 1991 UMI 300 N. Zeeb Rd. Ann Arbor, MI 48106 NOTE TO USERS THE ORIGINAL DOCUMENT RECEIVED BY U.M.I. CONTAINED PAGES WITH SLANTED PRINT. PAGES WERE FILMED AS RECEIVED. THIS REPRODUCTION IS THE BEST AVAILABLE COPY. SYSTEMATICS OF ARALIA (ARALIACEAE) DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of the Ohio State University By Jun Wen, B.S. ***** The Ohio State University 1991 Dissertation Committee: Approved by: Daniel J. Crawford Michael Evans Tod F. Stuessy Advisor Department of Plant Biology Dedicated to Hanmin ii ACKNOWLEDGMENTS This work would never have been completed without generous help from numerous people. Professor Tod F. Stuessy, my major advisor, is gratefully acknowledged for his generous advice, constant encouragement and continuous support throughout all phases of my graduate study. I also greatly appreciate my committee members, Professors Daniel J. Crawford and Michael Evans for their advice and critical review of the manuscript. Discussions with other persons of the systematics program at The Ohio State University have also contributed to this study. Jorge E. Arriagada, Melanie DeVore, John J. Furlow, Thomas G. Lammers, Paul 0. Lewis, David M. Spooner, Ronald L. Stuckey, and James C. Zech deserve special mention. I would also like to express my appreciation to Dr. David Frodin, well known systematist of Araliaceae, for stimulating discussions. This study was supported by many foundations and institutions. Financial support for field work was provided by the National Science Foundation (BSR-8815325), Central China Agricultural University, the Explorers Club, Ohio State University-Hubei Province Exchange Program, Sigma Xi, and the Tinker Foundation. Travels to European herbaria were supported by an Alumni Research Award of The Ohio State University and a John Ayers Herbarium Travel Award from the American Society of Plant Taxonomists. Appreciation is expressed to the American Association of University Women for an International Fellowship (1988-89) and to The Ohio State University for a Presidential Fellowship (1990-91). Numerous friends and colleagues provided help and assistance during my field and herbarium travel trips to China, Mexico, eastern U.S., California and Great Britain. Special thanks go to Diane Bridson, Martin Cheek, Zhiyuan Chen, Fernando Chiang, Nancy Coile, Sally Dawson, Barbara Ertter, Kuomei Feng, James W. Hardin, De-yuan Hong, Chiming Hu, Samuel B. Jones, Jr., Thomas G. Lammers, Na Li, Zhengwei Lu, Xiaozhong Luo, Ze-hui Pan, John F. Pruski, Edward E. Schilling, David M. Spooner, Guangfu Tao, Jose L. Villasenor R., Wencai Zhang, Peizhong Zheng and Souqing Zou. I am indebted to numerous local people from Guangdong, Hubei, Jiangxi and Yunnan of China for providing vehicles, food, lodging, and field assistances. The curators of A, B, BKF, BM, CM, E, G, GH, HIB, IBSC, K, K-W, KUN, L, LE, LU, MO, MU, NAS, NF, NY, OS, P, PE, PH, TAI, TAIF, U, UC, UNCC, US, W, and WU are gratefully acknowledged for permitting the examination of specimens, either through loans or during visits. Finally, I express my sincere and special thanks to my husband, Hanmin Dong, for all of his support, consideration, unfailing encouragement and many helpful suggestions. iv VITA December 27, 1963............ Born Hubei, China, P. R. 1980.......................... Jing Chou College-preparatory School, Hubeir China, P. R. 1984-8 5 ................ Teaching Assistant, Department of Forestry and Horticulture, Central China Agricultural University 1985-199 1.............. Graduate Associate, Department of Plant Biology, The Ohio State University PUBLICATIONS Wen, J. 1987. The epidermal characters and systematics of Aralia (Araliaceae). Ohio J. Sci. 87: 5. and T. F. Stuessy. 1988. A phylogenetic analysis of Nyssaceae. Ohio J. Sci. 88: 9. Tseng, C. C., K. L. Hedges,Y. C. Chen and J. Wen. 1988. Evaluation of extraction methods for two-dimensional electrophoresis of leaf proteins. Amer. J. Bot. 75 (6), pt.2: 133. Wen, J. and D. J. Crawford. 1989. Flavonoid chemistry of Aralia chinensis and A . spinosa (Araliaceae), an eastern Asian and North American disjunct species pair. Ohio J. Sci. 89: 18. Wen, J. and T. F. Stuessy. 1989. A phylogenetic analysis of Nyssaceae. Amer. J. Bot. 76 (6), suppl.: 282. Wen, J. 1990. Generic delimitations of Aralia (Araliaceae). Ohio J. Sci. 90: 20. Tao, G., J. Wen, T. F. Stuessy, and J. J. Furlow. 1990. A phenetic analysis of Aralia elata complex from eastern Asia. Ohio J. Sci. 90: 20; Amer. J. Boc. 7 7 ^6), suppl.: Tseng, C. C., K. L. Hedges, Y. C. Chen and J. Wen. 1990. Protein profile as a taxonomic character in Aralia (Araliaceae). Amer. J. Bot. 77 (6), suppl.: FIELDS OF STUDY Major Field: Plant Biology Studies in Plant Systematics and Biogeography with Professor Tod F. Stuessy vi TABLE OF CONTENTS ACKNOWLEDGMENTS ...................................... iii VITA ................................................... v LIST OF T A B L E S ...................................... xiii LIST OF F I G U R E S ........................... ............ X Chapter I. GENERIC DELIMITATION OF ARALIA (ARALIACEAE) 1 chapter II. INFRAGENERIC CLASSIFICATION OF ARALIA (ARALIACEAE) ...................................... 27 Chapter III. SYSTEMATICS OF ARALIA SECT. DIMORPHANTHUS MIQUEL (ARALIACEAE)................................ 57 Introduction ...................................... 57 Taxonomic History ................................ 59 Synopsis of Classification ....................... 63 Concepts of Species and Varieties ............... 66 Morphology and Taxonomic Criteria ................ 70 Reproductive Biology.............................. 81 Chromosome numbers.......................................84 Phylogenetic Analyses .............................. 87 Modes of Speciation................................ 114 Biogeography ...................................... 12 0 T a x o n o m y ........................................ 134 BIBLIOGRAPHY ........................................ 312 vii LIST OF FIGURES FIGURE PAGE 2.1. Strict consensus cladogram of taxa of Aralia ................................ 40 3.1. Generalized leaf architecture of species of sect. Dimorphanthus, showing the accessory leaflets (AL) and pinnae (AP) at each division of the rachises......................... 73 3.2. Cuticular patterns on leaflet abaxial surface of species of sect. Dimorphanthus: a. coronulate (A. spinosa); b. striated (A. thomsonii) . (X 600).......................... 77 3.3. Strict consensus cladogram of taxa of sect. Dimorphanthus.................................... 93 3.4. Strict consensus cladogram of taxa of series Chinensis................................. 99 3.5. Strict consensus cladogram of taxa of series Dimorphanthus............................ 105 3.6. Strict consensus cladogram of taxa of series Foliolosav............................... Ill 3.7. Generalized paleogeography of North Atlantic area in Early Eocene, showing the possible routes of connection between Europe and North America (from Tiffney 1985b).................................. 125 3.8. Generalized paleogeography of the Tethys Seaway, Early to Middle Eocene, also showing the Turgai Straits separating Asia from Europe (from Tiffney 1985b)............127 3.9. Distributions of Aralia foliolosa and A. merrillii .............................. 146 3.10. Distributions of Aralia ferox and A. montana......................................156 3.11. Distributions of Aralia armata and A. malabarica ................................. 164 3.12. Distributions of Aralia finlaysoniana var. finlaysoniana, A. finlaysoniana var. setacea and A.spinifolia .................. 172 viii 3.13. Distributions of A. thomsonii, A. searelliana, A. vietnamensis, A. chinensis var. chinensis, and A. chinensis var. longibractea .............................. 189 3.14. Distributions of Aralia dasyphylla and A. urticifolia ...........................