Phylogeny of the Genus Argia (Odonata: Coenagrionidae) with Emphasis on Evolution
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Phylogeny of the Genus Argia (Odonata: Coenagrionidae) with Emphasis on Evolution of Reproductive Morphology Dissertation Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Ryan Matthew Caesar Graduate Program in Evolution, Ecology and Organismal Biology The Ohio State University 2012 Dissertation Committee: Dr. Norman Johnson, Advisor Dr. Marymegan Daly Dr. John Freudenstein Dr. Hans Klompen Copyright by Ryan Matthew Caesar 2012 Abstract The damselfly genus Argia is found throughout the New World where some species are common and abundant members of lotic freshwater and adjacent ecosystems. Argia species are not only important predators of aquatic and terrestrial invertebrates but are themselves an important prey item to a variety of other insects and vertebrates. The distribution of species is highly variable within the genus and some species are locally threatened or endangered due to range limitation and habitat loss. Damselflies and dragonflies (Odonata) may be useful indicators of aquatic ecosystem health as well as indicators of climate change. There are approximately 120 species described with at least twenty suspected undescribed species. The taxonomy of the North American species is well known, but the Central and South American species are in need of revision. The phylogeny of the genus has never been studied using modern, repeatable methods. Therefore the evolutionary history of the genus has never been thoroughly explored. The reproductive biology of Odonata is unique among insects and provides a model system for testing hypotheses related to character evolution by sexual selection and other mechanisms of evolution. Argia species have unique morphologies of male and female secondary sexual characters, the modified cerci and paraprocts of males and the ii corresponding plates of the female pro- and meso-nota that are grasped by males during copulation and oviposition. The patterns of variation in these structures, both within and among species, may reveal the extent to which sexual and natural selection help shape the current diversity of the group. This dissertation presents phylogenetic hypotheses for the genus Argia using data from external morphology and ribosomal DNA. Maximum parsimony and maximum likelihood analyses were performed on the data, resulting in topologies that are mostly congruent, well-resolved, and moderately to highly supported. The variation in male cercus morphology is examined using three dimensional morphometrics where shape is quantified from computer tomography models. The phylogenetic hypotheses are used to examine patterns of cercus variation across the genus. The same methods are applied to populations of the widespread species Argia moesta in an attempt to test whether intrasexual selection applies to these important reproductive structures. iii Dedication Dedicated to my Father, Rodney Bruce Caesar January 8, 1949 – July 22, 2009 iv Acknowledgments There are many people who directly and indirectly played key roles in my graduate training and the production of this dissertation. I cannot possibly thank these people enough within the confines of this document, but I will mention them in brief as a sincere token of my appreciation for their support, guidance, and love. I would like to foremost thank my committee members for their guidance and patience during my time as a graduate student at Ohio State: Dr. Meg Daly, Dr. Hans Klompen, Dr. John Freudenstein and my advisor Dr. Norm Johnson. Not only has their instruction been instrumental in my growth as a biologist, for which I am especially grateful. I came to Ohio State and worked for several years as a student of Dr. John Wenzel before he took a position outside of the university. John taught me more about phylogenetic systematics, natural history, and many other topics than anyone else in my life. I am indebted to him for expanding my knowledge and abilities as a biologist and educator and for his support and friendship. Without the assistance of two generous scientists my work would not have been possible. I especially thank Dr. Rosser Garrison of the California Department of Food v and Agriculture and Dr. Mark McPeek of Dartmouth College for sharing their expertise with me on odonate biology. Dr. Ajit Chaudpari of the OSU Sports Biomechanics laboratory generously lent me use of his facilities computers and software for CT image analysis. My undergraduate advisor Dr. John Abbott of the University of Texas and my master’s degree advisor Dr. Anthony Cognato of Michigan State University continue to serve as mentors and friends, and their role in shaping me as a scientist is considerable. I thank them for all they continue to do for me. As an undergraduate student, Ahalya Skandarajah assisted me with molecular laboratory work. She presented a posted of her contributions at the Annual Meeting of the Entomological Society of America in Indianapolis in 2010. Members of the Wenzel lab that overlapped with me have not only become good friends but also helped shape many of the ideas in this dissertation: Sibyl Bucheli, Hojun Song, Joe Raczkowsi, Todd Gilligan, and Glené Mynhardt. For their friendship and collegiality I am eternally grateful. Many fellow graduate students, postdocs and researchers in the Museum of Biological Diversity at Ohio State have been helpful and inspirational along the way and I thank you immensely, including: Craig Barrett, Elijah Talamas, Steve Passoa, Alejandro Valerio, Charuwat Taekul, Monica Farfan, Kaitlyn Upstrom, Annie Lindgren, Brandon Sinn, Paul Larson, Abby Reft, Luciana Gusmao. I am grateful to my immediate family members for various forms of support, including but not limited to financial, motivational, and emotional: my parents-in-law Jim vi and Rita Shave, my brother-in-law Ryan Shave and his wife Carol, my sister Jenny Cresswell and her husband Ronny, my brother Chris, and my parents Rodney and Sue. For my entire life, Mom and Dad provided gentle guidance, never imposing demands but always suggesting that I explore all options and alternatives and take the path that suited me best. I hope I have done so. Most importantly of all, I thank my loving wife Jenny. She provided vast technical assistance with the production of figures for this document as well as numerous presentations throughout my graduate career, and her skills as a production artist are immeasurable. Her support, patience, and friendship are invaluable. Along the way, she gave me the greatest gift of my life, our son August Rodney. I’m sorry I took so long, and I love you dearly! vii Vita August 16, 1975 ........................................................ Born- Fort Knox, Kentucky May 2001 .................................................................. B.A. Zoology, University of Texas May 2004 .................................................................. M.S. Entomology, Texas A&M University Publications Caesar, R. M., N. Gillette, and A.I. Cognato. 2005. Population Genetic Structure of an Edaphic Beetle (Ptiliidae) Among Late Successional Reserves in the Klamath- Siskiyou Ecoregion, California. Annals of the Entomological Society of America 98(6): 931-940. Caesar, R. M., M. Sörensson, and A. I. Cognato. 2006. Integrating DNA Data and Traditional Taxonomy to Streamline Biodiversity Assessment: An example from Edaphic Beetles in the Klamath Ecoregion, California, USA. Diversity and Distributions 12(5): 483-489. Caesar, R.M. and J.W. Wenzel. 2009. A Phylogenetic Test of Classical Species Groups in Argia (Odonata: Coenagrionidae). Entomologica Americana 115(2): 97-108. viii Fields of Study Major Field: Evolution, Ecology and Organizmal Biology Specialization: Entomology, Systematics of Odonata ix Table of Contents Abstract ............................................................................................................................... ii Dedication .......................................................................................................................... iv Acknowledgments............................................................................................................... v Vita ................................................................................................................................... viii Table of Contents ................................................................................................................ x List of Tables .................................................................................................................... xv List of Figures .................................................................................................................. xvi Chapter 1 : General Introduction ....................................................................................... 1 The Order Odonata .......................................................................................................... 1 The Genus Argia ............................................................................................................. 2 Description of Dissertation Project ................................................................................. 3 Chapter 2 : A Phylogenetic Test of Classical Species Groups in Argia (Odonata: Coenagrionidae) .................................................................................................................. 5 Introduction ....................................................................................................................