INFORMATION TO USERS This manuscript has been reproduced from themicrofilm 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 qnali^ of the copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandardmargins, and improperalignment can adversely afreet 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 from left to right in equal sections withsmall overlaps. Each original is also photographed in one exposure and is included in reduced form at the back of the book. Photographs included in theoriginal 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. UMI University Microfilms International A Bell & Howell Information Company 3 0 0 North Z eeb Road. Ann Arbor. Ml 48106-1346 USA 313.'761-4700 800/521-0600 Order Number 9505185 Systematic studies of Calyceraceae DeVore, Melanie Lynn, Ph.D. The Ohio State University, 1994 Copyright ©1994 by DeVore, Melanie Lynn. All rights reserved. UMI 300 N. Zeeb Rd. Ann Arbor, MI 48106 SYSTEMATIC STUDIES OF CALYCERACEAE DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of the Ohio State University By Melanie Lynn DeVore, B.S. ***** The Ohio State University 1994 Dissertation Committee: Approved by: Daniel J. Crawford Michael L. Evans Tod F. Stuessy Adyisor-V Department of'Rlant Biology Copyright by Melanie L. DeVore 1994 To Neil A. Harriman, who along with serendipity, hooked me on vascular plant systematics. 11 ACKNOWLEDGMENTS I gratefully acknowledge a number of individuals and institutions for their help throughout this work. Tod F. Stuessy, my dissertation advisor provided help, as well as stimulating conversation and ideas concerning the early evolution of the Compositae and related families. I also extend appreciation to Daniel J. Crawford and Michael L. Evans for their advice and critical reading of the manuscript. I would also like to acknowledge Thomas and Edith Taylor who supported my first year of graduate studies at OSU and influenced some of my views regarding plant evolution. This dissertation would not have been possible without the support of faculty, graduate students, and staff, from the Departamento de Botanica, Universidad de Concepcion, Concepcion, Chile; Divisiones Entomologie, Plantas Vasculares, and Laboratorio de Sistematica y Biologie Evolutive (LASBE) of Museo de La Plata; and Institute de Botanica Darwinion. All of these institutions contributed to this study by supplying field vehicles and support, access to herbarium collections and library research. Ill Appreciation is expressed to; The American Society of Plant Taxonomists (Field Work Initiation Grant), The Ohio State University (Beatley Herbarium Award from OS, Latin American Studies Travel Award, and Graduate Student Alumni Research Award), and Sigma Xi for financial support. The following herbaria provided loans or access to herbarium collections: CONC, F, GH, LL, LP, MICH, MO, NY, OS, S, SI, TEX, US. Conversations with the following individuals concerning Asteraceae, Calyceraceae, or Goodeniaceae were extremely helpful: K. Bremer, A.L. Cabrera, A. Cronquist, V. Funk, E. Harris, T. hammers, C. Marticorena, R. Olmstead, and H. Robinson. Appreciation is also expressed to Jorge Arriagada for photographing type material and Monica Taylor for editing segments of the manuscript. Last but not least, I would like to thank my parents Don and Nancy DeVore for all their love, encouragement, and patience. Without their help, this work would not have been possible. IV VITA August 17, 1963............ Born - Warren, Ohio 1985....................... B. S. Geology, The University of Wisconsin-Oshkosh 1985-1987.................. Teaching Assistant, Depatment of Biology and Microbiology, The University of Wisconsin-Oshkosh 1987-1993.................. Graduate Associate, Department of Plant Biology, The Ohio State University 199 0....................... American Society of Plant Taxonomists, Field Work Initiation Grant 199 1....................... Sigma Xi, Grant-in-Aid of Research 1991....................... The Ohio State University Herbarium, Beatley Award 1991....................... The Smithsonian Institution, 10-week Graduate Student Fellowship 1991 The Ohio State University, Latin American Studies Program Travel Award 1991....................... The Ohio State University, Graduate Student Alumni Research Award 1993-1994 Lecturer, Department of Biological Sciences, Sam Houston State University V 1994 Botanist, Texas Regional Institute of Environmental Studies, Sam Houston State University 1994- Assistant Professor, Department of Biological Sciences, Sam Houston State University PUBLICATIONS DeVore, M. L. and N. A. Harriman. 1987. Juncus exil is Osterhout (Juncaceae), an overlooked synonym of J. confusus Coville. Sida 12: 243. DeVore, M. L. and T. N. Taylor. 1988. Mesozoic seed plants: a pollen organ from the Triassic of Antarctica [abstract]. Amer. J. Bot. 75: 105. DeVore, M. L. 1990. The occurrence of Acicarpha tribuloides Juss. in Eastern North America [abstract]. Ohio J. Sci. 85: 5. DeVore, M. L. 1990. The ballast distribution of Acicarpha tribuloides Juss. in eastern North America [abstract]. Amer. J. Bot. 77: 206. DeVore, M. L. 1991. Acicarpha tribuloides (Calyceraceae) in eastern North America. Rhodora 93: 26-35. DeVore, M. L. and Tod F. Stuessy 1991. Time and place of origin of Asteraceae, Calyceraceae, and Goodeniaceae [abstract]. Amer. J. Bot. 78: 178. DeVore, M. L. 1992. A new species of Calycera from northern Argentina [abstract]. Ohio J. Sci. 92: 17. DeVore, M. L. 1993. The systematics and biogeography of Acicarpha (Calyceraceae) [abstract]. Ohio J. Sci. 93: 41. DeVore, M. L. 1993. The systematics of Acicarpha (Calyceraceae) [abstract]. Amer. J. Bot. 80: 144 vi DeVore, M. L. 1994. Chromosome numbers in Calyceraceae [abstract]. Published abstracts. The Texas Academy of Science Meeting, Houston, TX, March 1994. DeVore, M. L. In press. Systematics of Argentinean Calycera (Calyceraceae) species [abstract]. Amer. J. Bot. 81: DeVore, M. L. In press. Calyceraceae and its relationships with Compositae [abstract]. Abstracts for The International Compositae Congress. Stuessy, T. F., T. Sang, and M. L. DeVore. In press. The Phylogeny and biogeography of Barnadesioideae [abstract]. Abstracts for The International Compositae Congress. Pesecreta, T. C., V. I. Sullivan, and M. L. DeVore. In press. The connective base of Acicarpha tribuloides (Calyceraceae). Amer. J. Bot. 81: DeVore, M. L. and T. F. Stuessy. In press. Time and place of origin of Compositae, Calyceraceae, and Goodeniaceae. In C. Jeffrey (ed.). Advances in Compositae. Academic Press, London. FIELD OF STUDY Major Field: Plant Biology Studies in Plant Systematics with Tod F. Stuessy V l l TABLE OF CONTENTS ACKNOWLEDGMENTS....................................... iii VITA.................................................. V LIST OF TABLES......................................... X LIST OF FIGURES........................................ xi INTRODUCTION........................................... 1 CHAPTER PAGE I. PLACE AND TIME OF ORIGIN OF ASTERACEAE, WITH ADDITIONAL COMMENTS ON CALYCERACEAE AND GOODENIACEAE ........................... 5 Introduction ........................... 5 Review of Morphology of Asteraceae, Calyceraceae, and Goodeniaceae......... 8 Present Distributions of Asteraceae and Goodeniaceae........................... 20 Trends in Fossil Records of Asteraceae and Goodeniaceae....................... 23 Tectonic Events in the Southern Hemisphere............................. 27 Paleoclimatic D a t a ..................... 30 References............................. 36 II. CYTOLOGY OF CALYCERACEAE................. 55 Introduction ........................... 55 Material and M e t h o d s ................... 56 Results................................. 62 Cytology in relation to morphology and distribution of Acicarpha ............. 62 Cytology and Relationships Within Calycera............................... 63 viii Nastanthus and its Relationship with Calycera......................... 65 Estimate of Ancestral Base Number of Calyceraceae....................... 68 Cytology and Biogeography............... 70 Literature cited......................... 71 III. SYSTEMATICS OF ACICARPHA .................. 76 Introduction ........................... 76 Taxonomic History ....................... 77 Generic Relationships ................... 78 Phenetic studies of the Acicarpha tribuloides Complex ................... 79 Cladistic Analysis ..................... 91 T a x o n o m y .................................102 Refere n c e s ............................... 132 IV. SYSTEMATICS OF CALYCERA...................... 140 Introduction ........................... 140 Taxonomic History......................... 141