Comparative Analysis of Courtship in <I>Agelenopsis</I> Funnel-Web Spiders
Total Page:16
File Type:pdf, Size:1020Kb
University of Tennessee, Knoxville TRACE: Tennessee Research and Creative Exchange Doctoral Dissertations Graduate School 5-2012 Comparative analysis of courtship in Agelenopsis funnel-web spiders (Araneae, Agelenidae) with an emphasis on potential isolating mechanisms Audra Blair Galasso [email protected] Follow this and additional works at: https://trace.tennessee.edu/utk_graddiss Part of the Behavior and Ethology Commons Recommended Citation Galasso, Audra Blair, "Comparative analysis of courtship in Agelenopsis funnel-web spiders (Araneae, Agelenidae) with an emphasis on potential isolating mechanisms. " PhD diss., University of Tennessee, 2012. https://trace.tennessee.edu/utk_graddiss/1377 This Dissertation is brought to you for free and open access by the Graduate School at TRACE: Tennessee Research and Creative Exchange. It has been accepted for inclusion in Doctoral Dissertations by an authorized administrator of TRACE: Tennessee Research and Creative Exchange. For more information, please contact [email protected]. To the Graduate Council: I am submitting herewith a dissertation written by Audra Blair Galasso entitled "Comparative analysis of courtship in Agelenopsis funnel-web spiders (Araneae, Agelenidae) with an emphasis on potential isolating mechanisms." I have examined the final electronic copy of this dissertation for form and content and recommend that it be accepted in partial fulfillment of the requirements for the degree of Doctor of Philosophy, with a major in Ecology and Evolutionary Biology. Susan E. Riechert, Major Professor We have read this dissertation and recommend its acceptance: Gordon M. Burghardt, James A. Fordyce, Todd M. Freeberg Accepted for the Council: Carolyn R. Hodges Vice Provost and Dean of the Graduate School (Original signatures are on file with official studentecor r ds.) Comparative analysis of courtship in Agelenopsis funnel-web spiders (Araneae, Agelenidae) with an emphasis on potential isolating mechanisms A Dissertation Presented for the Doctor of Philosophy Degree The University of Tennessee, Knoxville Audra Blair Galasso May 2012 Copyright © 2012 by Audra Blair Galasso All rights reserved. ii DEDICATION I dedicate this dissertation to my parents, Joyce and Everett Payne, who always encouraged in me a love of books, knowledge and the natural world. iii ACKNOWLEDGEMENTS I wish to thank my advisor, Susan Riechert, for providing me with intellectual freedom and steadfast support throughout my time in graduate school. She introduced me to the fascinating world of spider behavior, and her lab was always an interesting place to be. I would also like to thank my committee members, Gordon Burghardt, Jim Fordyce and Todd Freeberg. Our interactions in and out of the classroom broadened my understanding of animal behavior and evolutionary biology. I would also like to express my appreciation to the Department of Ecology & Evolutionary Biology, which provided me with a teaching assistantship and an intellectually stimulating environment. I would like to acknowledge those individuals who helped me collect spiders for this project: Steve Galasso, Kim Kennard, Julie Morgan, Jonathan Pruitt and Casey Willingham. I am grateful to George Uetz for allowing me to use his vibration recording equipment at the University of Cincinnati and to his students, Shira Gordon and Brian Moskalik for assistance with that equipment. My fellow students offered stimulating discussion and support, particularly Mackenzie Taylor and my fellow Riechert Lab members – Jonathan Pruitt, Sarah Duncan, Jamie Troupe, Sara Bengston and Austin Dillon. I would also like to acknowledge funding for this project, which came from NSF grant 0315901 from the Evolutionary Processes Program to Susan Riechert, an EEB Summer Research Grant and the Vincent Roth Fund for Systematic Research from the American Arachnological Society. iv Finally, I wish to express deep appreciation to my arachnophobic husband, Steve Galasso, who moved to Knoxville to support my graduate education and even helped me collect and feed spiders. v ABSTRACT The genus Agelenopsis (Araneae, Agelenidae) is a group of morphologically similar funnel-web spiders with overlapping habitat requirements and geographic ranges. Yet, molecular evidence suggests this group has undergone recent speciation with no evidence of hybridization. In this dissertation, I explored courtship divergence as a possible explanation for the formation and coexistence of these species. Courtship behavior patterns, sequences, and vibratory signals were compared across 12 Agelenopsis species and three related outgroup species. Courtship in Agelenopsis was found to be comparatively long but not completely species-specific. To investigate mechanisms of reproductive isolation within the genus, interspecific crosses were staged in the lab between species for two closely related species pairs, one pair being sympatric in distribution and the other peripatric. Vibratory courtship did not function as a reproductive isolating mechanism for either pair. In the sympatric species pair, males displayed significantly less courtship toward heterospecific females than to conspecific females and no interspecific copulations were attempted. Females did not reject males based on their vibratory courtship, rather males rejected females, apparently based on chemical cues. In contrast, there was interspecific courtship and copulation in the peripatric species pair. However, males appeared to encounter mechanical difficulties copulating with heterospecific females and no hybrid offspring were produced. Thus, the sympatric pair was reproductively isolated by barriers acting early in the mating vi process, while the geographically isolated pair appeared to be isolated by barriers acting very late in the mating process. Though these results did not support a role for vibratory courtship in reproductive isolation, vibratory courtship appeared to be quite important in Agelenopsis. Males continued to court females even after females accepted them, and even behavior occurring late in the courtship sequence varied among species. In comparison to the two more distantly related outgroup species, vibratory courtship in Agelenopsis seems to be under relatively strong sexual selection for long, complex displays. vii TABLE OF CONTENTS INTRODUCTION .................................................................................................. 1 Courtship Function and Evolution ..................................................................... 2 The genus Agelenopsis: Natural History and Speciation ................................... 3 Goals of the Dissertation Research ................................................................... 5 Literature Cited .................................................................................................. 7 CHAPTER I COMPARATIVE ANALYSIS OF COURTSHIP IN FUNNEL-WEB SPIDERS .............................................................................................................. 9 Abstract ........................................................................................................... 10 Introduction ..................................................................................................... 11 Methods .......................................................................................................... 17 Collections and rearing conditions ............................................................... 17 Courtship trials ............................................................................................. 19 Quantitative description of courtship ............................................................ 20 Relative durations of two courtship phases and incidence of mixed phase behavior ....................................................................................................... 23 Tests to determine if species can be discriminated by courtship behavior .. 23 Phylogenetic analysis of courtship ............................................................... 26 Results ............................................................................................................ 28 General courtship features ........................................................................... 28 viii Behavior patterns exhibited ......................................................................... 30 Test for differences in rearing history ........................................................... 31 Species differences in courtship variables ................................................... 32 Assessing prevalence of courtship after female acceptance ....................... 33 Species-specificity of courtship .................................................................... 34 Phylogenetic analyses ................................................................................. 36 Discussion ....................................................................................................... 39 General features of funnel-web spider courtship ......................................... 39 Possible mechanisms of courtship divergence in Agelenopsis: Drift and phylogeny .................................................................................................... 43 Possible mechanisms of courtship divergence in Agelenopsis: Reinforcement ............................................................................................. 44 Possible mechanisms of courtship divergence in Agelenopsis: Sexual selection ......................................................................................................