The evolution of mating systems in black scavenger flies (Diptera: Sepsidae) Item Type text; Dissertation-Reproduction (electronic) Authors Schulz, Katja-Sabine 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 26/09/2021 22:27:05 Link to Item http://hdl.handle.net/10150/289010 INFORMATION TO USERS This manuscript has been reproduced 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. 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Contact UMI directly to order. Bell & Howell Information and Learning 300 North Zeeb Road, Ann Artxsr, Ml 48106-1346 USA 800-521-0600 THE EVOLUTION OF MATING SYSTEMS IN BLACK SCAVENGER FLIES (DIPTERA: SEPSIDAE) by Katja-Sabine Schulz 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 1999 UMI N\iinber; 9946792 UMI Microform 9946792 Copyright 1999, by UMI Company. All rights reserved. This micrororm edition is protected against unauthorized copying under Title 17, United States Code. UMI 300 North Zeeb Road Ann Arbor, MI 48103 2 THE UNIVERSITY OF ARIZONA ® GRADUATE COLLEGE As members of the Final Examination Committee, we certify that we have read the dissertation prepared by Katja-Sabine Schulz entitled The Evolution of llatinp Systems in Black Scavenger Flies (Diptera: Sepsidae). and recommend that it be accepted as fulfilling the dissertation requirement for the Degree of Doctor of Philosophy D^vid Maddison Dati j 'I- 7 Martha Hunter Date Diana V/heeler J^&ith Bronstein Date 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 chat I have read this dissertation prepared under my direction and reconmend that it be accepted as fulfilling the dissertation requirement. ^ hO Di sertation Director ^ -J JJ. Date y Davxd Maddxson 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 acknowledgement 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 judgement the proposed use of the material is in the interest of scholarship. In all other instances, however, permission must be obtained from the author. SIGNED: 4 ACKNOWLEDGEMENTS I sincerely thank my advisor David Maddison for his patience and support during the painfully slow gestation of this dissertation. Without his persistent encouragement, this work would never have been finished. I also thank the other members of my Final Examination Committee, Judi Bronstein, Molly Hunter, and Diana Wheeler for all their suggestions and consistent positive attitudes toward my work. Over the years, Dan Papaj, Martin Taylor, John Alcock, and Bob Smith served on my ever evolving dissertation committee, and each of them made significant contributions to the development of my studies. The manuscripts for this dissertation were improved by comments from my Final Committee as well as John Alcock and Wolf Blanckenhom. I am particularly grateful to Bill Eberhard for carefully reading drafts of each of my chapters and for giving much constructive criticism of my interpretations of the data. While I have not heeded all of his suggestions, his input has greatly enhanced my thinking and writing during the critical gelling phase of this dissertation. Special thanks go to Rudolf Meier and Armekati Reichardt, my collaborators on various projects. Karen Ober, Wendy Moore, and Gene Hall, my spirited companions in the Maddison lab, often made the dissertation time warp blues a little easier to bear. 5 TABLE OF CONTENTS LISTOFnGLJRES 7 LIST OF TABLES 8 ABSTRACT 9 INTRODUCTION 11 L POST-OVIPOSITION MATINGS AND PATTERNS OF SPERM PRECEDENCE IN SEPSIS PUNCTUM (FABRICIUS) 16 SUMMARY 16 INTRODUCTION 17 MATERIALS AND METHODS 22 Oviposition And Sperm Storage 22 Experimental Families For Paternity Studies 25 Paternity Analyses 28 RESULTS 30 Oviposition And Sperm Storage 30 Patterns Of Sperm Precedence 31 DISCUSSION 35 Do Males Remove Rival Sperm? 35 Why Do Males Pursue Post-oviposition Matings? 37 Is The Last Male Advantage Associated With All Matings? 42 Co-segregation Of RAPD Markers In Paternity Analyses 44 n. THE TIMING OF MATINGS IN BLACK SCAVENGER FLIES 57 SUMMARY 57 INTRODUCTION 58 MATERIALS AND METHODS 60 RESULTS 65 DISCUSSION 69 in. MALE-FEMALE ASSOCIATIONS DURING OVIPOSITION IN BLACK SCAVENGER FLIES 86 SUMMARY 86 INTRODUCTION 87 MATERIALS AND METHODS 89 RESULTS 93 DISCUSSION 95 Adaptive Significance Of Escorting Behaviour 95 Evolutionary History Of Escorting Behaviour 98 6 TABLE OF CONTENTS - Continued IV. GEOGRAPHIC VARIATION IN THE BEHAVIOUR AND MORPHOLOGY OF BLACK SCAVENGER FLIES 112 SUMMARY 112 INTRODUCTION 113 MATERIALS AND METHODS 116 Patterns Of Sexual Size Dimorphism 116 Sex Ratios At Oviposition Sites In The Field 119 RESULTS 120 Patterns Of Sexual Size Dimorphism 120 Sex Ratios At Oviposition Sites In The Field 122 DISCUSSION 123 V. THE GENETIC BASIS OF PRE-COPULATORY ESCORTING BEHAVIOUR IN SEPSIS PUNCTUM (FABRICIUS) 143 SUMMARY 143 INTRODUCTION 144 MATERIALS AND METHODS 146 Parental Stocks And Substrate 146 Individual Male Assays For The Expression Of Escorting Behaviour 147 Pooled Assays For The Expression Of Escorting Behaviour 149 RESULTS 154 Individual Male Assays For The Expression Of Escorting Behaviour 154 Pooled Assays For The Expression Of Escorting Behaviour 156 DISCUSSION 158 CONCLUSIONS 178 APPENDIX A: Nucleotide Sequences Of Diagnostic RAPD Primers 186 APPENDIX B: Populations, Habitats, And Substrates Used For Behavioural Observations And For The Collection Of Founders For Laboratory Populations 187 REFERENCES 188 7 LIST OF HGURES FIGURE 1.1, The mating pattern of S. punctiim 47 FIGURE 1.2, Outline of the mating regime used to produce families for the analysis of patterns of sperm precedence 49 FIGURE 2.1, Representative stages of ovarian development in S. punctiim 74 FIGURE 2.2, Results of ovarian dissections of sepsid females collected in copula in the field 76 FIGURE 2.3, Results of ovarian dissections of Themira piitris females caught solitary or in copula in the field 78 FIGURE 2.4, Phylogenetic relationships of sepsid genera with information on the mating behaviour of gravid females 80 FIGURE 3.1, Ovipositing S. punctum female accompanied by an escorting male 104 FIGURE 3.2, Phylogenetic relationships of sepsid genera with information on the occurrence of escorting behaviour 106 FIGURE 3.3, Foreleg of a male Xenosepsis fukuharai 108 FIGURE 4.1, Mean developmental times and tibia length measurements for S. punctum males and females from escorting and non-escorting lines 131 FIGURE 4.2, Frequency distributions of sex ratios at oviposition sites in the field 133 FIGURE 5.1, Breeding designs for comparisons of the expression of escorting behaviour 164 FIGURE 5.2, Distribution of the proportion of escorted oviposition bouts for male offspring of F2 generation crosses 166 FIGURE 5.3, Mean escorting scores of S. punctum males from different crosses 168 8 LIST OF TABLES TABLE 1.1, Oviposition patterns of S. punctum females 51 TABLE 1.2, Offspring produced by S. punctum females that had mated only once 53 TABLE 1.3, Distribution of RAPD markers that were diagnostic for either putative father in offspring sample 1 and 2 of experimental families 55 TABLE 2.1, The timing of matings in sepsid females 82 TABLE 2.2, Results of focal observations of sepsid females at oviposition sites in the field (timing of matings) 84 TABLE 3.1, Results of focal observations of sepsid females (escorting behaviour) 110 TABLE 4.1, Number of replicate offspring clutches assayed