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TERMITE SOCIAL EVOLUTION by Timothy George Myles a Dissertation Submitted to the Faculty of the DEPARTMENT of ENTOMOLOGY in Part

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TERMITE SOCIAL EVOLUTION by Timothy George Myles a Dissertation Submitted to the Faculty of the DEPARTMENT of ENTOMOLOGY in Part Termite social evolution Item Type text; Dissertation-Reproduction (electronic) Authors Myles, Timothy George 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 06/10/2021 05:46:53 Link to Item http://hdl.handle.net/10150/558092 TERMITE SOCIAL EVOLUTION by Timothy George Myles A Dissertation Submitted to the Faculty of the DEPARTMENT OF ENTOMOLOGY In Partial Fulfillment of the Requirements For the Degree of DOCTOR OF PHILOSOPHY In the Graduate College THE UNIVERSITY OF ARIZONA 19 8 8 2 THE UNIVERSITY OF ARIZONA GRADUATE COLLEGE As members of the Final Examination Committee, we certify that we have read the dissertation prepared by Timothy George Myles_____________________ entitled TERMITE SOCIAL EVOLUTION and recommend that it be accepted as fulfilling the dissertation requirement Doctor of Philosophy the Degree of Robert L. Date / Diana E. Wheele, Date William B. Heed 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 .y/ ' Dissertation Director William L. Nutting D a t e / 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 major department or the Dean of the Graduate College when in his or her 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. 4 ACKNOWLEDGEMENTS The most vital support is financial and I have many debts of gratitude to express to those who helped me financially over the last seven years. I thank Dr. Nutting for offering me the Dow Fellowship which financed my first two years at Arizona and for obtaining some additional support from Merck Chemical Co. I thank the Department of Entomology Faculty for providing me with a teaching assistantship for the next two years. I thank my parents for generous financial support at critical times. I thank Dr. Bob Smith for rescuing me twice from destitution with employment over the last two and a half years. I thank my uncle, Vale Myles, for a gift that kept me afloat during early 1988 while I was writing chapter four. I also thank the Entomology Faculty for assistance to attend several meetings and for approving money for page charges on several publications. For help with page charges I especially thank Dr. Roger Huber. I am grateful for travel awards from Sigma Xi and the Graduate Student Program Development Fund. I thank my wife, Alicesara, for working full time through two pregnancies and then returning immediately to work, for maintaining all the finances on the home front, and for being a wonderful mother and loving wife. I thank my step daughters, Darunee and Andrea, for being an enormous help to Alicesara at work and for the loving care they have given to Tisha and Tyler—to Da and Andrea I dedicate chapter five. I thank my parents, George and Florence Lee Myles for their continuous faith, encouragement, and love. Through it all Bob Smith has been a counselor, ally, and friend. He encouraged my thinking with enthusiasm and lavish moral support. Without him I may not have made it. Bob’s friendship has made the time so much more enjoyable and meaningful. I also wish to thank Dr. Ross Zimmerman and Patrick Hahn for intelligent and entertaining discussions about termite social behavior that enriched my understanding. I thank Drs. Steve Buchman, David Byrne, Justin Schmidt, and Floyd Werner for reviews of early drafts of chapter three. I also thank Dr. Scott Sakaluk, Dr. David Zeh, and Patrick Hahn for thoughtful reviews of chapter five which helped to improve it. I thank my committee members, Drs. Robert Smith, Diana Wheeler, William Heed, Astrid Kodric-Brown, and in particular my advisor William Nutting for showing great forbearance over the long course of my studies. They have all been subjected to my long and "turgid" writing style in drafts of several manuscripts, preliminary exams, and the present weighty dissertation. My thinking and writing have benefitted substantially from their input. I thank Dr. Con Slobodchikoff for inviting me to participate in the conference at NAU on The Ecology of Social Behavior, out of which grew chapter six. Lastly, I thank Dan Hogue for showing me how to use Lotus and the H/P plotter, Dave Bentley for help on the SEM, and Justine Collins for assistance in preparation of tables for the final draft. 5 TABLE OF CONTENTS Page LIST OF ILLUSTRATIONS................................................................................................... 9 LIST OF P L A T E S.................................................................................................................... 12 LIST OF TABLES . ............................................. 14 ABSTRACT .............................................................................................................................. 16 CHAPTER ONE: HISTORICAL AND CONCEPTUAL BACKGROUND ON THE THE QUESTION OF TERMITE SOCIAL EVOLUTION Introduction......................... 1........................... .......................................................... 18 Historical Background................................................................................................ 20 Conceptual Background.............................................................................................. 28 CHAPTER TWO: REVIEW OF SECONDARY REPRODUCTION IN THE ISOPTERA Introduction......................................................................... 33 Methods ............ 35 Results ............................................................................................................ 36 Discussion ......................................................................................! ............................ 85 , Definitions of Primary and Secondary Reproductives............................ 85 Major Consequences of Secondary Reproduction.................................... 85 Definitions of Replacement and Supplementary R eproductives.......... 85 Developmental Origins and Types of Secondaries ................................. 86 Pseudergate Reproductive P otential........................................................... 86 Phylogenetic Patterns of Secondary R eproduction................................. 87 Neotenic Differentiation............................................................................... 90 Extent of Female Physogastry in Relation to Type of Secondaries ... 91 Factors Prompting Secondary Differentiation ......................................... 92 Secondary Reproduction in Relation to Nesting-Feeding Strategies . 93 1) Neotenic Production in Relation to Dry Endoxylophagy . 93 2) Secondary Reproduction in Relation to Damp Endoxylophagy 94 3) Secondary Reproduction in Relation to Xylophagous Foraging 95 4) Secondary Reproduction in Relation to Arboreal Xylophagy 96 5) Secondary Reproduction in Relation to Epigeous Nesting 96 6) Secondary Reproduction in Relation to Mound Building . 98 7) Secondary Reproduction in Relation to Humivory ........... 100 8) Secondary Reproduction in Relation to Grass and Litter H arvesting......................... 101 9) Secondary Reproduction in Relation to Nest Inquilinism . 102 Summary of Secondary Reproduction in to Nesting-Feeding Strategy........................................... 102 On the Origin of Neotenic Reproduction ................................................ 103 Hypotheses Concerning How Secondary Reproduction Was Important in Subsequent Social E volu tion .......................... 104 Bartz’s Inbreeding Induced Relatedness Asymmetry Hypothesis ................................................... 104 Resource Inheritance Interpretations.......... ................................. 105 Comparisons of Secondary Reproduction in Lower and Higher Term ites.................... 106 TABLE OF CONTENTS (continued) 6 Page The Primitive Nature of the Neotenic Caste ........................................... 107 CHAPTER THREE: ON THE ORIGIN OF THE SOLDIER CASTE IN TERMITES Introduction..................................................................................... 108 Alternative Interpretations of Reproductive Soldiers The Selfish Soldier Hypothesis .................................................................. I ll The Defensive Soldier Hypothesis ............................................................ 112 Occurrence of Reproductive Soldiers in Termopsidae ......................................... 112 Significance of Reproductive Soldiers..................................................................... 113 Primitive Ecology Favored Replacement Reproduction and Sibling
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