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U·M·I University Microfilms International a 8Ell & Howell Information Company 300 North Zeeb Road Patterns of homoplasy in North American Astragalus L. (Fabaceae). Item Type text; Dissertation-Reproduction (electronic) Authors Sanderson, Michael John. 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 10/10/2021 18:39:52 Link to Item http://hdl.handle.net/10150/184764 INFORMATION TO USERS The most advanced technology has been used to photo­ graph and reproduce this manuscript 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 UIVn 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 re­ produced 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. These are also available as one exposure on a standard 35mm slide or as a 17" x 23" black and white photographic print for an additional charge. 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. U·M·I University Microfilms International A 8ell & Howell Information Company 300 North Zeeb Road. Ann Arbor. M148106-1346 USA 313/761-4700 800/521-0600 Order Number 9000144 Patterns of homoplasy in North American Astragalus L. (Fabaceae) SandersOl~, Michael John, Ph.D. The University of Arizona, 1989 U·M·I 300 N. Zeeb Rd. Ann Arbor, MI 48106 PATTERNS OF HOMOPLASY IN NORTH AMERICAN ASTRAGALUS L. (FABACEAE) by MICHAEL JOHN SANDERSON A Dissertation Submitted to the Faculty of the DEPARTMENT OF ECOLOGY AND EVOLUTIONARY BIOLOGY In Partial Fulfillment of the Requirements For the Degree of DOCTOR OF PHILOSOPHY In the Graduate College THE UNIVERSITY OF ARIZONA 1 9 8 9 THE UNIVERSITY OF ARIZONA GRADUATE COLLEGE As members of the Final Examination Committee, we certify that we have read the dissertation prepared by __M_i_c_h_a __ e_l __ J_. __ s__ a_n_d_e_r_s_o __ n__________________ ___ entitled Patterns of homoplasy in North American Astragalus and recommend that it be accepted as fulfilling the dissertation requirement for the Degree of Doctor of Philosophy M. Donoghue Date ' ~~~~ R. Hoshaw ~~f{-.~~~ R. Robichaux Date Final approval and acceptance of this dissertation is contingent upon the candidate's submission of the final copy of the dissertation to the Graduate College. I hereby certify that I have read this dissertation prepared under my direction and recommend that it be accepted as fulfilling the dissertation requirement. lJ\\~t~:r, .b~, \\-JL- Dissertation Director Date ) 3 STATEMENT BY AUTHOR This dissertation 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 dissertation are allowable without special permission, provided that accurate acknowledgment of source is made. Requests for permission for extended quotation from or reproduction of this manuscript in whole or in part may be granted by the head of the maj or department or the Dean of the Graduate College when in his or her judgement 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: 4 ACKNOWLEDGMENTS Michael Donoghue initially proposed the idea of working on the "hopeless" genus Astragalus, although he should not be held accountable for my desire to pursue a proj ect of the present scope. His enthusiasm for studying evolutionary problems via cladistic methods was infectious and greatly influenced the course of my work. As always, his tenacious reading of early drafts of the dissertation improved it immeasurably. Thanks also to the other members of my committee, Dr. William Heed, Dr. Robert Robichaux, Dr. Robert Hoshaw, .and Dr. Richard Strauss, all of whom provided much needed criticism of the present work. Rupert Barneby gave badly needed encouragement to my early thoughts on working in the genus. I thank him for taking the time to respond at such length to my letters and for hosting my visit to the New York Botanical Garden in 1986. The present work has benefited greatly from the intellectual climate provided in Dr. Donoghue's lab by Geeta Bharathan, Mark Porter, and Jim Malusa. Larry Hufford advised me on many developmental issues and the ideas about character resolution in Chapter Three grew from conversations with him. Ned Friedman also helped me on my first attempts at serial sectioning. Renee Rusler deserves much of the credit for all my field work, since I probably would not have gone without her, and she continually prodded me into looking over one more hill for those damn plants. Geeta, George Ferguson, Marty W., and most especially George Davenport also accompanied me on field trips and helped to make all of them memorable. I thank the curators of several herbaria in Ylhich I worked at various times, including NY, RSA, NM, and the especially the staff at UC (Barbara Errter and Elizabeth Neese). Dr. C.T. Mason and Becky van Devender are especially deserving of my thanks for providing a "home" in the U of A herbarium. I would like to acknowledge a debt to four present and former faculty members in the Department of Ecology and Evolutionary Biology, each of whom at early stages in my graduate studies helped awaken in me an interest in "real" biology. They are Dr. James H. Brown, Dr. Charles H. Lowe, Dr. Robert Hoshaw, and Dr. Charles T. Mason. A special word of thanks is also due Rick Michod, my first major professor, who in very many ways is responsible for my involvement in evolutionary biology. Finally, I thank my parents for providing more than occasional financial support throughout my lengthy stay in graduate school. This research was supported by an Herbarium Travel Grant from the American Society of Plant Taxonomists, and by a McGinnies Scholarship from the Office of Arid Lands Studies at the University of Arizona. ---------.--~--- 5 TABLE OF CONTENTS LIST OF TABLES 7 LIST OF FIGURES . 8 ABSTRACT 11 1. INTRODUCTION: HOMOPLASY AND EVOLUTION . 13 Homoplasy: Definitions and History . • . 13 Homoplasy and Homology . 16 Classical View (Similarity Alone) ..• 17 Evolutionary View (Similarity plus Common Ancestry) ........•... 20 Transformational View ("Transformation" regardless of Similarity) . • . • . 25 Taxic View (Homology equals Synapomorphy) . 26 Homology as Information continuity . 30 Homoplasy and Homology Defined . • . 31 Homoplasy in Relation to Evolution and Phylogeny •.•.... 33 Homoplasy and orthogenesis · · · · · · 33 Homoplasy and Tendencies · · · · · · 36 General Patterns of Homoplasy · · · · · · 41 2. PHYLOGENETIC ANALYSIS OF NORTH AMERICAN ASTRAGALUS 46 Introduction . 46 Overview of the Genus . 46 Monophyly of the Genus and its Relationships . • . 50 Methods .•••.••... • • • 55 Scope of Present Study • • • • 55 Characters and Taxa Used • • • 65 Polymorphism • • . • . • • • • • • • 76 Correlation of Characters · • • 77 Quantitative Characters .•.. • • • 80 Rooting ••.•.. • . • • • 84 Algorithms Employed .... 87 Results .••.•........ • • • • 88 Overview of Trees Found • • • 88 Notes on Selected Major Clades · . 91 Discussion . • . • . • • • • 108 Description of the Phylogeny • •• . 108 Relation of Results to Classical taxonomies . • • . 109 -_.- -.---------- 6 The Nature of Astragalus species: Cladistic Implications . • . · 121 Exclusion of South American Species · 123 Evolution of Selected Characters · 125 3. STATISTICAL TESTS FOR HOMOPLASTIC TENDENCIES 148 Introduction . • . 148 Methods . .. • . 152 Reconstructing the Distribution of Homoplasy. 152 Measures vf spatial Pattern . 156 Null Models for the Expected Distribution . 162 Tests of Significance . • . • . 169 The Computer Program . • . 170 Studies Employed and Characters Analyzed . 172 Results ...•..•......... 183 Behavior of the Null Models . •. 183 Clustering in Three Focal Characters . 186 Clustering in 14 Additional Characters • 191 Clustering in the Iguanid Lizard Study . 192 Discussion • . • • • . 192 APPENDIX 1: PATTERNS OF VARIATION IN LEVELS OF HOMOPLASY . 202 APPENDIX 2: DESCRIPTIONS OF CHARACTERS USED IN ANALYSIS OF CHAPTER TWO . 256 APPENDIX 3: CLADISTIC ANALYSIS OF ASTRAGALUS SECTION ARGOPHYLLI GRAY . 280 APPENDIX 4: LISTING OF PROGRAM FOR ANALYSIS OF HOMOPLASTIC TENDENCIES . 287 APPENDIX 5: MORPHOMETRIC VARIATION IN FRUITS OF ASTRAGALUS .. 305 APPENDIX 6: NEW TAXA OF ASTRAGALUS
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