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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. Bell & Howell Information and Learning 300 North Zaeb Road. Ann Arbor, Ml 48106-1346 USA 600-521-0600 UMT EVOLUTION AND BIOGEOGRAPHY OF MALESHERBIACEAE, AN ENDEMIC FAMILY OF ARID WESTERN SOUTH AMERICA DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of the Ohio State University By Karla Marie Gengler, B.A. ***** The Ohio State University 2000 Dissertation Committee: Approved by Professor Daniel J. Crawford, Co-adviser Professor Andrea Wolfe, Co-adviser Daniel J. CrawfoE Professor John Wenzel Department of Evolution, Ecology, and Organismal Biology K , L L '^/ Andrea Wolfe, Department of Evolution, Ecology, and Organismal Biology UMI Number 9982566 Copyright 2000 by Gengler, Karla Marie Ail rights reserved. UMI* UMI Microform9982566 Copyright 2000 by Bell & Howell Information and teaming Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code. Bell & Howell Information and teaming Company 300 North Zeeb Road P.O. Box 1346 Ann Arbor, Ml 48106-1346 Copyright by Karla Marie Gengler-Nowak 2000 ABSTRACT Maiesherbiaceae, which are endemic to arid Peru, Chile, and adjacent Argentina, provide an interesting research opportunity, as the family’s small size and distribution in a virtual continental island make it ideal for phylogenetic and biogeographic studies. The family’s distribution in both Chile and Peru is unusual since the adjacent countries share very few desert taxa, so Maiesherbiaceae are an ideal group for the study of the biogeography of the entire dry region west of the Andes. To study the biogeography of the family, a phylogeny based on ITS sequence data was reconstructed. This analysis confirmed the monophyly of the family and showed it to consist of five major clades. Four of the five major clades contain morphologically very similar species; the two species comprising the Hfth share few such traits. The five major clades are herein recognized as subgenera. These subgenera correspond closely to the distributions of species in the family, suggesting that the physical history of the region strongly influenced the evolution of Maiesherbiaceae. The morphologically distinct Peruvian species form a subgenus and constitute a terminal clade in the family, indicating the family originated in Chile and subsequently migrated to Peru. By examining the geological history of the region, it was u inferred that the family split from Tumeraceae in the late Miocene. A rapid radiation giving rise to the fîve subgenera occurred in the Pliocene, with a more recent radiation event affecting much of the family in the Pleistocene. Allozyme diversities of four Peruvian inter-Andean species were examined. The data suggest that at least two of the species are the result of founder events; morphological similarities among the four closely related taxa indicate that they may be recently diverged. Founder events may be relatively common in the flora of the arid inter-Andean valleys because these valleys are quite isolated by intervening regions of high elevation. The M. humilis Poepp. species complex was also studied in detail because the four species and three varieties previously described are difficult to distinguish. A phenetic analysis of morphological data gathered from herbarium specimens. The further emphasized the similarities among groups of organisms. Five clusters are recognized as varieties of a single species, M. humilis. One former species, M. multiflora Ric., is not recognized as separate from M. humilis var. parviflora, and two others, M. gabrielae Ric. and Af. taltalina Ric., are recognized as varieties of M. humilis. Ill Dedicated to my husband, Joseph Nowak IV ACKNOWLEDGMENTS I thank my primary adviser, Dr. Daniel J. Crawford, for his thoughtful critiques and advice, which encouraged me greatly during the course of my studies. I also thank my former adviser. Dr. Tod F. Stuessy, for introducing me to Maiesherbiaceae, helping me begin my studies, and offering much-needed advice on travel in South America and morphological work. Dr. Andrea Wolfe has also been a source of encouragement and helpful critiques of my work; I further thank her for agreeing to co-ad vise me. I thank Dr. John Wenzel for use of his copies of the programs NONA, Pee-Wee, and Hennig86, his many hours of assistance with these programs, and especially for spending so much time teaching me cladistic theory. I am grateful to Dr. John Freudenstein for his help with the translation of my diagnoses into Latin and with the rules of botanical nomenclature. My phenetic and genetic diversity studies were improved greatly by the helpful comments of Dr. Theresa Culley and Lisa Wallace. My three field seasons in South America would not have been successful without the assistance, advice, and field support of the faculty, staff, and students of a number of Chilean and Peruvian institutions. I gratefully thank the botany department of the Universidad de Concepcidn for their hospitality and advice as I studied their large collection of Malesherbia. I also thank the staff of the Museo Nacional de Historia Natural In Santiago, Chile and the faculty at the Universidad de La Serena’s herbarium. My Peruvian field studies were made possible through the assistance of the faculty, staff, and students at the Museo Nacional de Historia Natural in Lima, the Universidad Nacional de San Cristobal de Huamanga in Ayacucho, Museo Contisuyo in Moquegua, the Universidad Nacional de San Agustin in Arequipa, and the Universidad Nacional del Centro in Huancayo. I extend my thanks to the Institute de Recursos Naturales of Peru for permission to collect and export specimens from that country. I thank the U.S. Department of Education for granting me a Foreign Language and Area Studies Fellowship for the study of Spanish, which was necessary for my field work. This study was funded by NSF grant DEB-9623496, the Beatley Herbarium Fund, Sigma Xi Grants-in-Aid of Research, a Tinker Foundation Grant, and the Flora of Chile Project. VI VITA November 6, 1970 Bom—Adrian, Minnesota 1993 B.A. Biology, College of St. Benedict, St. Joseph, Minnesota 1994, 1996, 1997, 1998, 1999 Graduate Teaching Associate The Ohio State University 1995, 1996 Herbarium Curatorial Assistant The Ohio State University 2000-present Instructor, Department of Biology, Denison University PUBLICATIONS I. D M. Spooner, D. De Jong, B.-Y. Sun, T.F. Stuessy, K.M. Gengler, G.L. Nesom, & P £ . Berry. 1995. Chromosome counts of Compositae from Ecuador and Venezuela. Ann. Missouri Hot. Gard. 82:596-602. 2. T.F. Stuessy & K. Gengler. 1995. Maiesherbiaceae. Pp. 297-298. In S. Takahashi et al. (eds.). The World of Plants. Asahi Shimbun, Tokyo. vii FIELDS OF STUDY Major Field: Evolution, Ecology, and Organismal Biology vui TABLE OF CONTENTS Page Dedication ..................................................................................................................... iv Acknowledgments ......................................................................................................... v Vita...............................................................................................................................vii List of Figures ............................................................................................................. xi Chapters; 1. Introduction .............................................................................................................l 1.1 Literature Cited......................................................................................9 2. Genetic diversities of four little-known species of Malesherbia (Maiesherbiaceae) endemic to the arid inter-Andean valleys of Peru .................................................................................10 2.1 Introduction ......................................................................................... 10 2.2 Materials and Methods ....................................................................... 12 2.3 Results..................................................................................................13 2.4 Discussion............................................................................................15 2.5 Literature Cited..................................................................................