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Diptera: Tephritidae): New Species, Phylogeny, and Justifications

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Diptera: Tephritidae): New Species, Phylogeny, and Justifications SYSTEMATICS OF THE GENUS RHAGOLETIS (DIPTERA: TEPHRITIDAE): NEW SPECIES, PHYLOGENY, AND JUSTIFICATIONS By Daniel Lloyd Hulbert A DISSERTATION Submitted to Michigan State University in partial fulfillment of the requirements for the degree of Entomology—Doctor of Philosophy 2018 ABSTRACT SYSTEMATICS OF THE GENUS RHAGOLETIS (DIPTERA: TEPHRITIDAE): NEW SPECIES, PHYLOGENY, AND JUSTIFICATIONS By Daniel Lloyd Hulbert Flies of Rhagoletis (Diptera: Tephritidae) are economically important fruit pests (infesting specialty fruit crops including apples, blueberries and cherries), which also serve as models for studying modes of speciation and coevolutionary relationships with their hymenopteran parasitoids. There are new species within the genus which have not been previously formally described. One of these species is within the tabellaria species group. I describe the morphology of Rhagoletis bushi Hulbert & Smith, its geographic distribution, host association, phylogenetic relationships, and identify an associated species of parasitoid wasp. The new species infests the fruit of buffaloberry (Shepherdia argentea) in the Northern Great Plains of North America. There is a suite of morphological characters, and a unique host plant association, that are diagnostic of R. bushi. Further evidence for the validity of R. bushi and its placement within the tabellaria species group comes from DNA sequence data from multiple genetic loci. The phylogenetic relationships among Rhagoletis species groups remain unresolved despite analyses based on morphology, allozymes, and mitochondrial DNA. Most Nearctic Rhagoletis belong to one of five species groups (pomonella, tabellaria, cingulata, suavis, and ribicola groups), with two unplaced species (R. fausta and R. juniperina), all of which appear to be part of a larger monophyletic group that also includes some Palearctic taxa. Regarding the overall phylogeny of the genus, my goals were to 1) resolve phylogenetic relationships using mitochondrial (COI) and nuclear (28S, CAD, period, AATS) DNA sequences, and 2) to identify the monophyletic group containing these Nearctic species. Using Bayesian analysis of a combined dataset with 4399 aligned nucleotides, I inferred a well-supported monophyletic group containing the five Nearctic Rhagoletis species groups, plus R. fausta, R. juniperina, and two Palearctic species: R. batava and R. flavigenualis. Within this larger monophyletic assemblage, the five Nearctic species groups together are monophyletic as are four of the five individual species groups (not ribicola). Palearctic and Neotropical Rhagoletis were resolved into well- supported clades of taxa often sharing closely related host plants. A well-resolved phylogeny of Rhagoletis is a valuable tool for future work addressing questions pertaining to how history, geography and ecology have shaped the phylogenetic patterns we observe in the genus. It is often claimed that systematic biology is fundamental to all other areas of biology. I critically evaluate the acceptance of this claim by entomologists critically as it relates to the field of entomology. I also critically describe the justification and valuations for systematic biology using the framework of Boltanski and Thévenot’s realms of worth and the philosophical framework for justification using virtues, desserts and outcomes. In order to accomplish these purposes, I critically analyze and review relevant entomological literature and interview practitioners of entomology and insect systematic biology. I find justification for systematic biology overwhelming takes the form of appeals to utilitarianism (both internally and externally focused) and are most relevant in the Industrial World. Additionally, some justifications given also pertain to the Civic World and to virtue. Evaluation of justification in systematic biology is important, especially as our globe becomes increasingly ecologically and politically unstable. Dedicated to Jean Hulbert. iv ACKNOWLEDGEMENTS I would like to acknowledge the C. S. Mott Predoctoral Fellowship in Sustainable Agriculture for funding a portion of my program. Thank you to Lyman Briggs College for providing consistent teaching assistant opportunities to help pay my stipend. I would also like to acknowledge the Michigan State University Dissertation Completion Fellowship for funding my final semester. I would like to thank our colleagues who assisted in insect collections including Amanda E. Nelson, Bernhard Merz, Dorie Bush, Daniel Frias, Guy Bush, Jorge Graziano, John Jenkins, J. P. Luz, James Smith, Martín Aluja, Meredith Doellman, Megan Frayer, Parita Shah, Rufus Isaacs, R. Pavia, Stewart Berlocher, Thomas Hoffmeister, Valery Korneyev, Wee Yee, Yue Ming, Peter Rausse and Scott Beck. I would like to thank Sydney Barosko, Morgan Potter, and Mitch Katkic for their help in the laboratory. Thank you to Severyn Korneyev for his taxonomic and morphological expertise. Thank you to my advisor, James J. Smith, for giving me the initial opportunity, and for his assistance and mentorship over the years of my program. I would like to thank the other members of my committee, Anthony Cognato, Ronald Debry, and Craig Harris for their guidance. Finally, I would like to thank all of the friends and family members who have supported me throughout the years of my program. Especially my parents, Nancy Scheer and Thomas Hulbert who, in so many ways, have supported me. v PREFACE My dissertation is likely somewhat unconventional when compared to others in entomology PhD programs and I would like to give an explanation before the main document. This dissertation has two chapters that would be considered typical entomology research where I detail my research on the systematics of the Rhagoletis (Diptera: Tephritidae). However, the third chapter addresses sociology of entomology and systematic biology. In that third chapter, I investigate and discuss the social dimensions of systematic biology. Why is that sociology chapter present in an otherwise conventional entomological dissertation of insect systematics? Early in my PhD program, I applied for and received the C. S. Mott Predoctoral Fellowship in Sustainable Agriculture. Part of the requirements of the fellowship included my participation in the Graduate Specialization in Ecological Food and Farming Systems (EFFS). The EFFS specialization “fosters understanding of interdisciplinary concepts critical to sustainable agriculture and food systems.” Functionally, the EFFS specialization is designed to give bio/geo/chemical scientists a stronger and formal background in the social sciences and vice versa. In fulfillment of my participation in the EFFS specialization, I have worked on a research project investigating the social justifications for systematic biologists given by entomologists. It may initially seem that the three major chapters of my dissertation make strange neighbors however, as I elaborate in chapter three, I believe considerations of justification and social context are very important things for natural scientists to include in their work. Additionally, the chapter of this dissertation titled “Description of a new species…” has already been published in the journal Insect Systematics and Diversity (Hulbert et al. 2018). As such, the chapter is not technically the nomenclatural act for the new species, although it does vi contain a description of it. To address this, the “Nomenclature” section of the chapter has been altered relative to the published work to reflect the fact that the present dissertation does not contain any official nomenclatural acts. Also, I have removed all instances of the text “n. sp.” from the chapter. I believe these actions I have taken are sufficient to avoid any taxonomic confusion and blunders, but my purpose of including these details in the dissertation overview is to make my actions and intensions explicit. vii TABLE OF CONTENTS LIST OF TABLES ........................................................................................................................ x LIST OF FIGURES .................................................................................................................... xii INTRODUCTION......................................................................................................................... 1 Why study systematics? ............................................................................................................ 1 Systematics defined ................................................................................................................. 1 Justification ............................................................................................................................. 2 Introduction to Rhagoletis ........................................................................................................ 4 Diversity of small organisms .................................................................................................. 4 Overview of the genus ............................................................................................................. 5 Previous phylogenetic analyses .............................................................................................. 7 Dissertation overview................................................................................................................ 9 CHAPTER 1: DESCRIPTION OF A NEW RHAGOLETIS (DIPTERA: TEPHRITIDAE) SPECIES IN THE TABELLARIA SPECIES GROUP ...........................................................
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