A TAXONOMIC STUDY OF THE LENNOACEAE. Item Type text; Thesis-Reproduction (electronic) Authors Yatskievych, George Alfred, 1957- Publisher The University of Arizona. Rights Copyright © is held by the author. Digital access to this material is made possible by the University Libraries, University of Arizona. Further transmission, reproduction or presentation (such as public display or performance) of protected items is prohibited except with permission of the author. Download date 23/09/2021 14:29:13 Link to Item http://hdl.handle.net/10150/274684 INFORMATION TO USERS This reproduction was made from a copy of a document sent to us for microfilming. While the most advanced technology has been used to photograph and reproduce this document, the quality of the reproduction is heavily dependent upon the quality of the material submitted. The following explanation of techniques is provided to help clarify markings or notations which may appear on this reproduction. 1. 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Other University Microfilms International A TAXONOMIC STUDY OF THE LENNOACEAE by George Alfred Yatskievych A Thesis Submitted to the Faculty of the DEPARTMENT OF PLANT SCIENCES In Partial Fulfillment of the Requirements For the Degree of MASTER OF SCIENCE WITH A MAJOR IN HORTICULTURE In the Graduate College THE UNIVERSITY OF ARIZONA 19 82 STATEMENT BY AUTHOR This thesis has been submitted in partial fulfillment of requirements for an advanced degree at The University of Arizona and is deposited in the University Library to be made available to borrowers under rules of the Library. Brief quotations from this thesis are allowable without spe­ cial permission, provided that accurate acknowledgment of source is made. Requests for permission for extended quotation from or repro­ duction of this manuscript in whole or in part may be granted by the heaci of the major department or the Dean of the Graduate College when in his judgment the proposed use of the material is in the interests of scholarship. In all other instances, however, permission must be obtained from the author. SIGNED: APPROVAL BY THESIS DIRECTOR This thesis has been approved on the date shown below: CHARLES T. MASON, JR. " Date Professor of Ecology ana Evolutionary Biology ACKNOWLEDGMENTS This study was not accomplished without help. In fact, most endeavors of similar magnitude require the cooperation a great many people to insure their success. To the legion of friends, colleagues, and other researchers who made this project possible, I owe a great debt of gratitude, which is here detailed. My committee members, Charles T. Mason Jr., John R. Reeder, and Paul G. Bartels showed the utmost support during this study, and their guidance and criticisms are greatfully acknowledged. Dr. Mason, in particular, went far beyond the call of duty in helping to arrange and participating in field work, as well as providing facilities and equip ment. In addition, the counseling of Robert H. Hoshaw was very impor tant to me. Dr. Hoshaw also lent important equipment. Correspondence with Job Kuijt greatly stimulated my research, and his encouragement and suggestions played a major role in the planning of this study. Conversations with Gary Nabhan, Laurence J. Toolin, and Richard S. Felger were also very important in the development of the project. I thank the curators and staff of all the herbaria from whom I borrowed specimens for the privilege of studying their collections of Lennoaceae, and for the time and effort they spent in loaning their material to me. In particular I would like to thank Cathy Breslin and Rebecca Eiber of ARIZ for their patience in dealing with iii iv an, at times, demanding graduate student. The staff at ASU, ASC, DES, TEX, and UNLV were most courteous and helpful during the visits. I am further grateful to all the botanists with whom I have corresponded concerning this study. These researchers generously- provided opinions and data on various aspects of nomenclature, distribution, and taxonomy: Julian A. Steyermark, Dan H. Nicolson, Robert L. Dressier, Stephen D. Koch, Grady L. Webster, Jerzy Rzedowski, John Lewis, Patricia K. Holmgren, Christopher Davidson, A. Charpin, R. Doug Stone, Arthur Gibson, Neil Harriman, Reid Moran, C. Ochsenius, Arthur M. Phillips, Paloma Blanca Fdez. de Caleya, Frans Stafleu, Robert F. Thorne, and Luz Ma. Villarreal de Puga. Thomas R. VanDevender and the staff of the Arizona Natural Heritage Program were also instrumental in the acquisition of distributional data. The analysis of pollen morphology of the Lennoaceae required special equipment and techniques that I did not possess at the beginning of this project. For help with various aspects of this por­ tion I am indebted to Robert Thompson, Wayne Ferris, Michael McClure, David Bentley, and Gopi Rao. The persistence and stamina of my "field assistants" has never ceased to amaze me, and a particular thank-you is due to those energetic individuals who devoted long hours in hot, uncomfortable locales, to the strenous activity of excavating the plants essential to my study. They are: Ed Collins, Carole Jenkins, Ken Preston-Mafham, Joe Cicero, Paul Shaw, Carl Wilson, Peggy Gallagher, Sylvia Forbes, V Allen Kelley, Jane Evans, and Frank Reichenbacher. I am also grate­ ful to Alan A. Beetle for leading a field expedition to Mexico, and to Jody Lee Duek and Greg and Carol Starr, for collecting plants of Lennoaceae for me. I am grateful to Mark Jorgenson of Anza Borrego Desert State Park (California) and to Robert L. Moon of Joshua Tree National Monument (California) for permission to study plants in their parks, and for courtesies extended during my visits. The office of the Subsecretaria Forestal of the Secretaria de Agriculture y Recursos Hidraulicos, Mexico, was most helpful in issuing a permit allowing me to make collections of Lennoaceae in Mexico. Funding for portions of this study was provided by research grants from Sigma Xi, the Scientific Research Society, and from the University of Arizona Graduate Research Development Fund, whose support is gratefully acknowledged. The magnificent line drawings found in Chapter 3 were rendered by Margaret Kurzius, and the final thesis draft was typed by Joan Farmer, whose efforts are appreciated. Finally, I would like to thank my family for their continued support, and following the precedent set by Moran (1962), I would like to especially thank my parents, without whose early efforts this study would not have been possible. J TABLE OF CONTENTS Page LIST OF ILLUSTRATIONS viii ABSTRACT ix 1. INTRODUCTION 1 Taxonomic History of the Family 4 Chemistry 6 Ethnobotany 7 Cytology 10 Palynology 13 Other Morphology and Anatomy 24 Stems, Roots, and Leaves 25 Inflorescences, Flowers, and Fruits 27 Ecology and Biogeography 34 Host Plants 45 Relationships of the Family 49 2. TAXONOMIC ANALYSES 53 Evaluation at the Generic Level 54 Species Evaluations in Lennoa 56 Species Evaluations in Pholisma 58 Evolution in the Family 60 3. SYSTEMATIC TREATMENT 63 Key to the Genera of Lennoaceae 64 Key to the Formae of Lennoa madreporoides 66 Key to the Species of Pholisma 70 APPENDIX A: GRAPHIC ANALYSIS OF LENNOA MADREPOROIDES 76 APPENDIX B: SPECIMENS EXAMINED 97 LITERATURE CITED 117 vi LIST OF ILLUSTRATIONS Figure Page 1.