The ecology and evolution of tachinid-host associations Item Type text; Dissertation-Reproduction (electronic) Authors Stireman, John Oscar 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 03/10/2021 23:55:09 Link to Item http://hdl.handle.net/10150/289745 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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ProQuest Information and Leaming 300 North Zeeb Road, Ann Arbor, Ml 48106-1346 USA 800-521-0000 NOTE TO USERS This reproduction is the best copy available. UMI' THE ECOLOGY AND EVOLUTION OF TACHINID-HOST ASSOCIATIONS by John Oscar Stireman III 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 200 1 UMI Number: 3010196 UMI ® UMI Microform 3010196 Copyright 2001 by Bell & Howell Information and Learning Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code. Bell & Howell Information and Learning Company 300 North Zeeb Road P.O. Box 1346 Ann Arbor, Ml 48106-1346 2 THE UNIVERSITY OF ARIZONA GRADUATE COLLEGE As members of the Final Examination Conmiittee, we certify that we have read the dissertation prepared by John Oscar Stireman III entitled Thp Frnlngy and F vn 1 r 111 nn nf Tarhirn'rl-Mngt Associations and recommend that it be accepted as fulfilling the dissertation requirement for the Degree of Doctor of Philosophy ^ Date BAnviMpjAn. 4-lg-Q/ Date Date , , _^l(S/d ( \ 7 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 that I have read this dissertation prepared under my direction and recommend that it be accepted as fulfilling the dissertation requirement. Dissertation Difector Datfe 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 the 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. SIGNED 4 ACKNOWLEDGEMENTS I am grateful to the many people that contributed to my dissertation work in a wide variety of ways. I thank those who served on my committee over the course of my graduate study: Nancy Moran, Dan Papaj, Judie Bronstein, David Maddison, Molly Hunter, and Bob Smith. All have generously provided expertise in their respective areas that vastly improved the quality of the studies contained in this dissertation. I would like to make special note of the contribution of Dan Papaj to the behavioral studies I conducted and David Maddison to the phylogenetic study. Without their expertise, these projects would not have been possible. I would especially like to thank my advisor Dr. Nancy Moran for serving as my advisor despite our rather divergent interests, generously providing laboratory space and facilities, and critically evaluating my dissertation work. I am also indebted to Dr. Elizabeth Bemays for her encouragement and counseling regarding many of the studies contained herein. Mike Singer has profoundly influenced all the studies in this dissertation due to our extensive discussions of the interactions between parasitoids and their hosts, his collaboration on several of the projects described in this dissertation, and his gracious donations of advice and assistance. He also provided an excellent field companion given his vast knowledge of natural history. Matt Kaplan was also invaluable in generously providing time and expertise in order to teach me how to collect and analyze molecular sequence data. He, along with the other "Varones" (Harold Greeney, Caleb Gordon) made my graduate career one of the most interesting and exhilarating times of my life. I would like to thank many other students and post-docs who contributed to my dissertation and graduate study including: The Moran lab (Patrick Abbot, Dan Funk, Helen Dunbar, Jay Withgott, Alex Mira, Apama Telang, Barry Sullender), EEB graduate students/post-docs (Marshall Hedin, Jen Weeks, Gita Bodner, Eileen Hebets), and Entomology graduate students (Kevin Moulton, Dan Hahn, Matt Johnston, John Hoekstra, Jun Isoe, Barret Klein, Karen Ober). I would also like to thank several professors and University staff that have provided their time and/or expertise including; Yves Carriere, Carl Olson, Reg Chapman, John Jaenicke, and Hayward Spangler. Several students, amateur naturalists, and biologists have helped with field work and/or provided data, information, and identifications concerning Lepidoptera and/or tachinids including: Jim O'Hara, Monty Wood, Bruce Walsh, Ray Nagle, Jim Tuttle, Jim Brock, Nate Hubert, Patrick Phoebus, and Sucheta Katakkar. Finally, I would like to acknowledge my family for their enormous support. My wife. Shannon and my son, Aaron, have graciously put up with me and supported me through this long process, and my mother, Dianne, has always encouraged me to pursue my interests. I am greatly thankful to my father, John Stireman jr., for encouraging my interest in ecology, evolution, and entomology, and for helping me develop the ability to think critically. 5 TABLE OF CONTENTS ABSTRACT 7 1 INTRODUCTION 9 2 PRESENT STUDY 21 REFERENCES 27 A MULTI-SCALE SPATIAL AND TEMPORAL VARIATION IN THE PARASITOID COMMUNITY OF AN EXOPHYTIC POLYPHAGOUS CATERPILLAR 32 B HOST SELECTION CUES IN A GENERALIST TACHINID PARASITOID; THE IMPORTANCE OF HOST MOVEMENT 79 C LEARNING IN A GENERALIST TACHINID PARASITOID 119 D PHYLOGENETIC RELATIONSHIPS OF TACHINID FLIES IN THE SUBFAMILY EXORISTINAE (TACHINIDAE: DIPTERA) BASED ON 28S RDNA AND EFla, AND ANALYSES OF HOST RELATED CHARACTER EVOLUTION 162 TABLE OF CONTENTS - Continued ECOLOGICAL AND EVOLUTIONARY DETERMINANTS OF TACHINID SPECIES RICHNESS AND PARASITISM RATES IN A COMMUNITY OF MACROLEPIDOPTERA 253 DETERMINANTS OF HOST USE BY A TACHINID PARASITOID COMMUNITY 315 7 ABSTRACT The Tachinidae is a taxonomically and ecologically diverse clade of parasitoids for which evolutionary and ecological relationships with hosts are largely unknown. Here, I employed a multidisciplinary approach to evaluate the determinants of patterns of host use in the Tachinidae. First, I examined spatio-temporal variation in the tachinid-dominated parasitoid assemblage of one lepidopteran species Grammia geneura. The parasitoid assemblage and parasitism rates varied dramatically among and within sampling sites, seasons, and years. I show that this variability may be a function of habitat-specific parasitism and indirect interactions between this host and other Macrolepidoptera through shared tachinid parasitoids. I then experimentally examined the host selection process in the tachinid Exorista mella. Host movement was an important elicitor of attack behavior. Flies also responded to odors associated with food plants of their host. Experienced flies attacked hosts more readily than did inexperienced flies. Based on these results, I proposed a host selection scenario for this tachinid species. E. mella also learned to associate colors with hosts and avoided deterrent models that they had experienced. However, I failed to find evidence for odor learning. Learning of host-associated cues by E. mella may allow this parasitoid to take advantage of abundant host populations and maintain host-searching efficiency in an unpredictable environment. To examine how host-associated characteristics evolved in the Tachinidae, I 8 reconstructed the evolutionary