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ZEEB ROAD, ANN ARBOR, Ml 48106 18 BEDFORD ROW, LONDON WC1 R 4EJ, ENGLAND 7922558 SHIELDS, WILLIAM MICHAEL PHILDPATRY, INBREEDING, AND THE ADAPTIVE ADVANTAGES 11F SEX. THE OHIO STATE UNIVERSITY, PH.D., 1979 COPR. 1979 SHIELDS, WILLIAM MICHAEL University Microfilms International j o o n . z e e b r o a d , a n n a r b o r , m i a s i o g <0/ Copyright by William Michael Shields 1979 PHILDPATRY, INBREEDING, AND THE ADAPTIVE ADVANTAGES DF SEX. DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy in the Graduate School of The Dhio State University By William lYiichael Shields, A.B., IY1.5. ****** The Ohio State University 1979 Reading Committee: Approved By Thomas C. Grubb, Jr. Paul A. Colinvaux George C. Williams Adviser (/ Department of Zoology ACKNOWLEDGMENTS All that follows is a collective effort. My personal endeavor has been shaped by the written and spoken words of colleagues too numerous to mention individually. Yet, I would feel remiss if I failed to publicly acknowledge those whose extraordinary patience and perserverance were most directly responsible for my task's completion. It is to them that much of the clarity in what follows owes its existence. I hope that it is obvious that any turbidity is my own, and my gratitude implies no responsibility for the errors which follow. My primary intellectual debt is to the work of George C. Williams, which helped my understanding of adaptation, and much more, which I hope will be obvious to the reader. I owe an open mind on the question of inbreeding to science fiction writer Theodore Sturgeon, though I hope our vocations are not confused. On a more personal level, I initiated this project while a student at Livingston College, Rutgers University. I thank the many faculty members and students who listened to my initial ramblings with tolerance. I am especially grateful to J. Burger, J. Chase, C. Leek, and T. Perper for encouraging me to pursue my ideas. I am also grateful to ii all my colleagues at the Ohio 5tate University For generously tutoring me in many necessary disciplines. Those who gave inordinately in continuous challenging discussion included K. Beal, K. Bildstein, C. Cook, J. Downhower, A. Gaunt, R. lYlitchell, 5. Peters, W. Peters, L. Putnam, and G. Pierce. My treatments of the topics to follow have benefitted from dicussion with T. lAIood, R. Milkman, B. Schaal, and especially C. Doner. I am particularly grateful to P. Greenwood and P. Harvey who unselfishly shared their thoughts and data Dn inbreeding, dispersal, and the structure of science. C. Birky, P. Colinvaux, T. Grubb, Jr., B. Murray, A. Hopkins, G. Williams, and IY1. Cunningham will have waded through the entire manuscript saving me considerable embarassment. I am both grateful and in awe. D. Dennis prepared most of the figures and S. McDowell helped with the typing. I acknowledge the following for their kind permission to make extensive use of previously published and copyrighted materials: J.L. Harper (Figs. 4-7); R. Selander and Academic Press, New York (Figs. 27 and 35); A. Beattie and 5pringer-Verlag (Fig. 8); J. Johnston and IAI. Heed and University of Chicago Press (Fig. 11); G. Rheinewald and Ardea (Fig. 12); J. Crow and M. Kimura and Harper and Row (Fig. 19); K. Mather and Chapman and Hall (Fig. 21); 5. Wright and University of Chicago Press (Fig. 23); M. Price and I\!. Ulaser and MacMillen Press (Fig. 29); iii J. Maynard 5mith and Cambridge University Press (Fig. 30); R. Highton and Plenum Press (Fig. 34); B. Wallace and IA1. IAI. Morton (Table 9); and G. Stebbins and Columbia University Press (Table 12) and Cold Spring Harbor Biological Laboratory (Table 13). Finally, I am indebted to the very special people whose intellectual and emotional support not only influenced this work profoundly, but also made life more bearable, stimulating, and joyous. I thank Barbara Brownsmith, Michael Cunningham, Bertram Murray, Thomas Grubb, and most especially Lea Shields and Janice Crook. It would certainly have been impossible without them. VITA 23 November 1947.........Born - Rockville Center, New York 1974.......................A.B., Livingston College of Rutgers University, New Brunswick, New Jersey 1974 - 1979............ ..Teaching Associate, Department of Zoology, The Ohio State University, Columbus, Ohio 1976.......................M.S., The Ohio State University, Columbus, Ohio 1976....................... Instructor in Biology, lAlilmington College, lAlilmington, Dhio 197 8 ....................... Lecturer in Biology, The Dhio State University, Lima, Ohio 197 9....................... Assistant Professor of Biology, State University of New York College Df Environmental Science and Forestry, Syracuse, New York PUBLICATIONS SHIELDS, IA1. 1Y1. and T. C. BRUBB, JR. 1974. Winter bird densities on north and south slopes. Wilson Bull. 86: 125-130. SHIELDS, W. IY1. , T. C. GRUBB, JR., and A. TEL IS. 1974. The use of native plants by monk parakeets in New Jersey. Wilson Bull. £5:172-173. SHIELDS, W. IY1, 1977. The effect of time of day on avian census results. Auk 94:380-383. 5HIELD5, W. 1Y1. 1577. The social significance of winter plumage variability: a comment. Evolution 31:905-907. v VITA (cant.) SHIELDS, IAJ. IYI. 1979. Avian census techniques: an analy­ tical review. In, Insectivorous birds and forest eco­ systems. (J.G. Dickson, R .N. Conner, R.R. Fleet, J.A. Jackson, and J.C. Kroll, eds.), Academic Press, (Mew York, In Press. SHIELDS, IAJ. IYI. 197S. Ground squirrel alarm calls: nepo­ tism or parental care? Amer. JMatur., In Press. SHIELDS, IAJ. IYI. and K. L. BILD5TEIIM. 1979. Birds versus bats: behavioral interactions at a localized food source. Ecology, In Press. GRUBB, T. C., JR. and IAJ. IYI. SHIELD5. 1977. Bald eagle interferes with an active osprey nest. Auk 94:140. FIELD OF STUDY lYlajor Field: Behavioral Ecology and Sociobiology Studies in Behavioral Ecology and Sociobiology. Professor Thomas C. Grubb, Jr. vi TABLE OF C0NTEI\1T5 Page ACKNOWLEDGMENTS ...................................... ii VITA .............................................. v LIST OF TABLES .......................................... ix LIST OF FIGURES ........................................ xi Chapter 1. INTRODUCTION ................................... 1 Function Versus Effect ................... 3 Ultimate Versus Proximate ......... 9 Philopatry, Inbreeding, and 5ex: An Overview ................................... IS 2. PATTERNS OF DISPERSAL ........................ 23 Passive Dispersal: Plants ............... 27 Passive Dispersal: Marine Inverte­ brates ......... 43 Active Dispersal: Insects and Ver­ tebrates SO 3. WHY PHILOPATRY: DISPER5AL AND INBREEDING .. 85 4. THE RELATIVITY OF RELATEDNESS AND INBREED­ ING ........................................... 107 Concepts of Relatedness; Definitions of Inbreeding ............................. 107 Genotypic Consequences of Inbreeding ... 125 Genomic Consequences of Inbreeding ..... 138 Phenotypic Consequences of Inbreeding .. 140 5. THE DISADVANTAGES OF INBREEDING ............. 144 Genotypic Disadvantages .......... 146 Genotypic Disadvantages: An Alternative View ........................................ 153 vii Chapter Page 6. AN ADVANTAGE FDR INBREEDING; COADAPTATIDN AND HYBRIDIZATION ............................. 169 Genetic Differentiation and Outbreeding Depression. ................ 174 Origin of Outbreeding Depression ........ 190 Origin and Maintenance of Philopatry .... 215 7. INBREEDING AND 5EX: PARADOX LD5T? ........... 235 The Disadvantages of Sex .................. 239 The Advantages of Sex ............ 244 Inbreeding and Sex:

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