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 9238305 Systematics of the genusMulinum pers. (Apiaceae, Hydrocotyloideae, Mulineae) Zech, James Craig, Ph.D. The Ohio State University, 1992 UMI 300 N. ZeebRd. Ann Aibor, MI 48106 SYSTEMATICS OF THE GENUS MULINUM PERS. (APIACEAE, HYDROCOTYLOIDEAE, MULINEAE) DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By James Craig Zech, A.A., B.S., M.S. ****** The Ohio State University 1992 Dissertation Committee: Approved by Daniel J. Crawford Fred Sack Tod F. Stuessy Departmen lant Biology My Parents ACKNOWLEDGEMENTS I would like to thank Dr. Tod F. Stuessy for his help and direction not only with this dissertation but also during my years at Ohio State. T thank Dr. Daniel J. Crawford for serving on my committees, for allowing me to work in his laboratory, and also for his advice and guidance. In addition, I thank my remaining committee members, Drs. Fred Sack (Advisory and Dissertation Committees) and Gary L. Floyd (Advisory Committee) for their advice and support. Without the participation of many people this work would not have been possible. I thank Dr. Lincoln Constance for his continued interest and encouragement and Dr. Thomas Duncan and the University of California, Berkeley Herbarium staff for their kindness and assistance. Because my field collecting would have been impossible without their support and direction, I thank Dr. Jorge V. Crisci at the Museo de La Plata, Argentina and Dr. Oscar O. Parra and Professor Clodomiro Marticorena at the Universidad de Concepcion Chile. I thank Maria Marta Cigliano, Claudio Contogiorgakis, Mario Gentilli, Patricia Gentilli, Liliana Katinas, Alegandra Moschetti, Alberto Ripalta, Fidel Alegandro Roig, Tod F. Stuessy, Ana Maria Zavattieri, and especially Juan Jose iii Morrone for help while in the field. Many thanks go to the directors, curators, and staff of the following herbaria: B, BM, CM, CONC, CORD, E, F, G, GH, GOET, K, L, LAU, LP, LY, MA, MO, NY, OS, OXF, P, PR, PRC, S, SGO, SI, UC, UPS, US, and W. Special thanks go to Dr. John J. Fur low (OS) for loan assistance, Maryan Benkowski (OS) for help with annotation labels, Mauricio Velayos (MA), Melica Mufioz Schick (SGO), and Esther G. de Mautino (CORD) for sending additional type material information, and Harald Riedl (W) for his efforts in type location. A very special thanks go to my colleagues, and friends, Jorge Arriagada, Maryan Benkowski, Marybeth Cosner, Patty Pacheco, Hugo Valdebenito, Loni Walker, and Jun Wen for hours of talks and constant support. I thank Sabine Zech for German translation, and Dr. R. L. Stuckey for the use of his computer. Finally, I thank my parents, A1 and Gerry Zech who have, more than anyone, supported me unconditonally during the bad times as well as the good times. iv VITA December 22, 1960 ..... Born - Grand Rapids, Michigan 1 9 8 1 ................... A.A., Grand Rapids Junior College, Grand Rapids, Michigan 1983 .................... B.S., Central Michigan University, Mt. Pleasant, Michigan 1984-1986 .............. Teaching Assistant, Central Michigan University, Mt. Pleasant, Michigan. 1986 ................... M.S., Central Michigan University, Mt. Pleasant, Michigan 1986-1991 .............. Teaching Associate, The Ohio State University, Columbus, Ohio 1988 ................... Research Assistant, Harvard University, Cambridge, Massachusetts. 1989-1991 .............. Lecturer, The Ohio State University, Columbus, Ohio 1 9 9 1 .................... Lecturer and Laboratory Instructor, Central Ohio Technical College, Newark, Ohio 1991-1992 .............. Lecturer and Laboratory Instructor, The Ohio State University - Newark Campus, Newark, Ohio PUBLICATIONS J.C. Zech. 1987. Micromorphological Characteristics of the Tegumen Layer Among Three Michigan Species of Luzula (Juncaceae). Abstract. Ohio J. Sci. 87: 5. 1988. The Juncus effusus Complex: The Latest Look. Abstract. Ohio J. Sci. 88: 12. 1989. Flavonoid Survey of Four Species Within The Tribe Mulineae (Hydrocotyloideae, Apiaceae). Abstract. Ohio J. Sci. 89: 7. 1989. A Preliminary Flavonoid Survey of the Genus Mulinum Pers. As Well As Other Taxa of the Tribe Mulineae (Hydrocotyloideae, Apiaceae). Abstract. Amer. J. Bot. 76: 285. 1990. Phenetic Analysis of Fruit Characters of the Genus Mulinum Pers. (Mulineae, Hydrocotyloideae, Apiaceae). Abstract. Ohio J. Sci. 90: 6. 1990. Phenetic Analysis of Fruit Characters of the Genus Mulinum Pers. As Well As Other Genera Within the Mulineae (Hydrocotyloideae, Apiaceae). Abstract. Amer. J. Bot. 77: 437. 1991. The Cushion Habit and the Genus Mulinum Pers. (Apiaceae). Abstract. Ohio J. Sci. 91: 27. 1991. Mulinum Pers. (Apiaceae) and the Cushion Habit: The Environmental Factors Which Cause Cushion Formation and Relationships to the Biogeography and Phylogeny of the Genus. Abstract. Amer. J. Bot. 78: 231. 1992. Classical Taxonomic Problems Encountered During the Revision of the Genus Mulinum Pers. Abstract. Ohio J. Sci. 92: 17. Zech, J. C. and D. E. Wujek. 1986. The Micromorphological Characteristics of the Tegumen Layer Between and Among Species of Juncus and Luzula including Varieties of the Juncus effusus Complex. Abstract. Pap. Michigan Acad. Sci. 1990. Scanning Electron Microscopy of Seeds in the Taxonomy of Michigan Juncus. Michigan Bot. 29: 3-18. FIELDS OF STUDY Major Field: Plant Biology Studies in Plant Systematics with Professor Tod F. Stuessy vi TABLE OF CONTENTS DEDICATION ........................................... ii ACKNOWLEDGEMENTS ..................................... iii VITA ................................................ v LIST OF TABLES ....................................... ix LIST OF FIGURES ........................ xi CHAPTER PAGE I. REVISION OF THE GENUS MULINUM PERS. (APIACEAE, HYDROCOTYLOIDEAE, MULINEAE) ...... 1 Introduction ............................. 2 Taxonomic History ........................ 3 Generic Relationships ................... 8 Habitat and Distribution ................ 11 Reproductive Morphology ................. 12 Chromosome Number ........................ 15 Phylogenetic Analysis of Mulinum ........ 16 Biogeography ............................. 24 Taxonomic Categories .................... 28 Morphology and Taxonomic Criteria ....... 28 Taxonomy .............................. 33 Key to Species of M ulinum........... 36 Excluded and Doubtful Names ............. 118 Acknowledgements ......................... 125 Literature Cited ........... 126 Numerical List of Taxa .................. 142 Index to Numbered Collections Examined ... 143 vii II. EVOLUTION OF CUSHION HABIT IN MULINUM (APIACEAE) .................................. 155 Introduction ........................... 156 General Factors Affecting Cushion Formation............................. 160 Factors Affecting Mulinum Cushion Formation ............................ 166 Evolution of the Cushion Habit in Mulinum ............................... 172 Literature Cited... ..................... 178 III. FLAVONOIDS OF MULINUM (APIACEAE, HYDROCOTYLOIDEAE, MULINEAE) ................ 187 Introduction ........................... 187 Materials and Methods.................... 191 Acquisition of Plant Material ......... 191 Extraction, Isolation, and Identification of Flavonoids ........ 191 Results ................................. 193 Discussion ............................. 196 Subfamilial and Tribal Flavonoid Variation .......................... 196 Generic and Infrageneric Flavonoid Variation inMulinum ................ 199 Evolution of the Flavonoid System in Mulinum ............................. 201 Literature Cited ..................... 2 05 LIST OF REFERENCES .................................. 210 viii LIST OF TABLES TABLE PAGE 1. Chromosome numbers for species of Mulinum .... 17 2. Characters and character states used in the cladistic analysis of Mulinum. 0 = primitive; 1,2 = derived ............................... 18 3. Character state matrix for phylogenetic reconstruction of Mulinum. 0 = primitive; 1, 2, = derived.