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NEODIPRION LECONTEI</Em> University of Kentucky UKnowledge Theses and Dissertations--Biology Biology 2017 EXAMINING THE ROLE OF HOST USE ON DIVERGENCE IN THE REDHEADED PINE SAWFLY, NEODIPRION LECONTEI, ACROSS MULTIPLE SPATIAL SCALES Robin Kimberly Bagley University of Kentucky, [email protected] Author ORCID Identifier: https://orcid.org/0000-0003-2209-0521 Digital Object Identifier: https://doi.org/10.13023/ETD.2017.285 Right click to open a feedback form in a new tab to let us know how this document benefits ou.y Recommended Citation Bagley, Robin Kimberly, "EXAMINING THE ROLE OF HOST USE ON DIVERGENCE IN THE REDHEADED PINE SAWFLY, NEODIPRION LECONTEI, ACROSS MULTIPLE SPATIAL SCALES" (2017). Theses and Dissertations--Biology. 46. https://uknowledge.uky.edu/biology_etds/46 This Doctoral Dissertation is brought to you for free and open access by the Biology at UKnowledge. It has been accepted for inclusion in Theses and Dissertations--Biology by an authorized administrator of UKnowledge. For more information, please contact [email protected]. STUDENT AGREEMENT: I represent that my thesis or dissertation and abstract are my original work. Proper attribution has been given to all outside sources. I understand that I am solely responsible for obtaining any needed copyright permissions. I have obtained needed written permission statement(s) from the owner(s) of each third-party copyrighted matter to be included in my work, allowing electronic distribution (if such use is not permitted by the fair use doctrine) which will be submitted to UKnowledge as Additional File. I hereby grant to The University of Kentucky and its agents the irrevocable, non-exclusive, and royalty-free license to archive and make accessible my work in whole or in part in all forms of media, now or hereafter known. I agree that the document mentioned above may be made available immediately for worldwide access unless an embargo applies. I retain all other ownership rights to the copyright of my work. I also retain the right to use in future works (such as articles or books) all or part of my work. I understand that I am free to register the copyright to my work. REVIEW, APPROVAL AND ACCEPTANCE The document mentioned above has been reviewed and accepted by the student’s advisor, on behalf of the advisory committee, and by the Director of Graduate Studies (DGS), on behalf of the program; we verify that this is the final, approved version of the student’s thesis including all changes required by the advisory committee. The undersigned agree to abide by the statements above. Robin Kimberly Bagley, Student Dr. Catherine R. Linnen, Major Professor Dr. David F. Westneat, Director of Graduate Studies EXAMINING THE ROLE OF HOST USE ON DIVERGENCE IN THE REDHEADED PINE SAWFLY, NEODIPRION LECONTEI, ACROSS MULTIPLE SPATIAL SCALES _____________________________________ DISSERTATION _____________________________________ A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in the College of Arts and Sciences at the University of Kentucky By Robin Kimberly Bagley Lexington, Kentucky Director: Dr. Catherine R. Linnen, Assistant Professor of Biology Lexington, Kentucky 2017 Copyright © Robin Kimberly Bagley 2017 ABSTRACT OF DISSERTATION EXAMINING THE ROLE OF HOST USE ON DIVERGENCE IN THE REDHEADED PINE SAWFLY, NEODIPRION LECONTEI, ACROSS MULTIPLE SPATIAL SCALES Phytophagous insects make up over one quarter of described species on Earth, and this incredible diversity seems directly linked to feeding on plants. Comparative studies of sister groups have shown shifts to herbivory are consistently associated with increased species diversity in insects, but the reasons for this diversification remain unclear. While other explanations, such as decreased extinction rates or influences on population structure, exist, one prominent hypothesis suggests shifts and subsequent adaptation to novel host plants can lead to the evolution of reproductive barriers. Given their extreme specialization on host plants in the genus Pinus and intimate, life-long association with their host plants, divergent host use has been suspected to drive speciation in the conifer sawfly genus Neodiprion. Previous work showed host shifts coincide with speciation events in the genus; but could not determine if these host shifts initiated speciation or if they occurred after other reproductive barriers arose. Determining the contribution and timing of host shifts relative to speciation will require examination of populations at the earliest stages of divergence, before post-speciation changes amass. If host shifts frequently drive speciation in the genus, there will likely be evidence of host-driven divergence within species occurring on a wide range of host plants. The goal of this dissertation is to examine populations of the red-headed pine sawfly, Neodiprion lecontei, an abundant, well-studied pest species that occurs on multiple hosts throughout its range, for evidence of host-driven divergence. Using a combination of reduced representation genomic sequencing, population genomics, and ecological assays, I specifically look for evidence of 1) genetic differentiation between populations utilizing different host plants, 2) ecological divergence in female oviposition preference, larval performance, and ovipositor morphology between populations on different hosts, and 3) ecologically-driven reproductive isolation between genetically and ecologically divergent populations. Each chapter of this dissertation examines the role of host use in driving ecological, genetic, and/or reproductive divergence within N. lecontei at a different spatial scale. First, I surveyed range-wide patterns of diversity. I identified three genetic clusters, dated the divergence of these clusters to the late Pleistocene, and found evidence that both dispersal limitation (geography) and host use contribute to genetic differentiation within N. lecontei. Next, I looked within one of these genetic clusters for additional evidence of the role of host in driving divergence. Sawflies in this cluster primarily utilize two hosts which differ significantly in needle architecture. Although I found no evidence of neutral genetic differentiation between hosts exists, I did detect spatial and temporal differences in host use, and host-specific differences in ovipositor morphology, a performance-related trait. Finally, I examine a single site where N. lecontei utilizes three structurally divergent species of pine. Although there was little genetic structure, no sexual isolation, and no distinct host preferences, the host types were partially temporally isolated and varied in ovipositor morphology and larval performance across on the three hosts. Overall, although divergent host use consistently resulted in divergent ovipositor morphology, a reduction in gene flow via temporal or geographic isolation may be required before additional forms of ecological and genetic differentiation can develop. Together these results suggest host shifts alone may not be enough to drive population divergence and speciation in Neodiprion. KEYWORDS: Host-associated divergence, Neodiprion sawflies, isolation-by-distance, isolation-by-environment, ecological speciation Robin K. Bagley Student signature July 24, 2017 Date EXAMINING THE ROLE OF HOST USE ON DIVERGENCE IN THE REDHEADED PINE SAWFLY, NEODIPRION LECONTEI, ACROSS MULTIPLE SPATIAL SCALES By Robin Kimberly Bagley Catherine R. Linnen Director of Dissertation David F. Westneat Director of Graduate Studies July 24, 2017 Date For my Poppy. Master crabber, vacation planner, weather watcher, and the best grandfather ever. Thank you for always watching over me. “Around here, however, we don’t look backwards for very long. We keep moving forward, opening up new doors and doing new things, because we’re curious… and curiosity keeps leading us down new paths.” -Walt Disney Acknowledgements During my time at UK, I have been lucky to have the opportunity to interact with many people who are not only talented scientists, but also wonderful and inspiring people. First, I would like to thank my advisor, Catherine Linnen. It will always be a bit of a mystery to me, but I will always be thankful that you took a chance on a random, sawfly-loving weirdo from a tiny school in New Jersey. From the very first emails we exchanged in 2009 (!), I have always been struck by your kindness, refreshingly down-to- earth nature, and willingness to share your ideas and knowledge. No matter what else was going on, your door was always open to discuss experimental design, the latest results, manuscript revisions, grant proposals, talks, posters, critter care… anything and everything I needed advice on. Thank you for not only celebrating all the good times, but also for commiserating with me in the not so good times. Working with you has made me a better scientist in every way I can think of; and I look forward to collaborating with you in the future as we expand the sawfly empire. To my committee – thank you for providing your wisdom, practical advice, and encouragement at each step along this journey. Your feedback and support have both been influential not only on my dissertation, but also on my development as a scientist. There are many steps in my journey that I would have missed were it not for the support of the incredibly kind and patient staff of the Biology Department. I would especially like to thank Bev Taulbee and Jacquie Burke, both of whom have not only gone out of their way to make sure I file all my
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