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Phylogenetics, Evolution and Systematics of Holodonata with Special Focus on Wing Structure Evolution: Morphological, Molecular and Fossil Evidence

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Phylogenetics, Evolution and Systematics of Holodonata with Special Focus on Wing Structure Evolution: Morphological, Molecular and Fossil Evidence PHYLOGENETICS, EVOLUTION AND SYSTEMATICS OF HOLODONATA WITH SPECIAL FOCUS ON WING STRUCTURE EVOLUTION: MORPHOLOGICAL, MOLECULAR AND FOSSIL EVIDENCE By SETH M. BYBEE A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2008 1 © 2008 Seth M. Bybee 2 To my wife for her endless supply of support, encouragement and attention and my children for their never ending curiosity. To my parents for all their love and guidance and my siblings for their examples. To all those that gave me a chance. 3 ACKNOWLEDGMENTS I am indebted to the long hours of dedicated help and support that I received from all those who served on my graduate committee. This project would not have been possible without their endless generosity, advice, and council. I am sincerely grateful to Dr. Alexandr Rasnitsyn and the paleoentomological staff at the Paleoentomological Institute, Dr. Andrew Ross at The Natural History Museum, and Dr. André Nel at le Muséum National d'Histoire Naturelle, who helped to make this research possible and who were superb hosts. Many thanks to Dr. David Grimaldi for his support, encouragement and insight into this research. Thanks to Dr. David Wahl for his help with optimizing the photographic equipment. This research was supported by grants from the Society of Systematic Biologists, the Explorers Club, the Systematic Association, the Entomological Society of America, and the Florida Entomological Society. The DNA component of this research was funded by NSF grants DEB 0120718, NSF DEB 0206505, NSF Research Experience for Undergraduates (REU) program DBI-0139501, Brigham Young University's Office of Original Research and Creative Activities, and the Entomological Foundation. This research represents a large collaborative effort from the odonate community and I am grateful for their generosity with specimen donations and identifications. I am particularly grateful to Jeff Skevington, Dennis Paulson, Akihiko Sasamoto, Karen Gaines, Rasmus Hovmöller, Jurg DeMarmels, L.-J. Wang, Adrian Trapero, Robert Larsen, Ken Tennessen, Bill Mauffray and the International Odonate Research Institute, Gainesville, FL, USA. I give special thanks to Andrew Rehn for providing specimens, specimen identification, discussions and character coding of his character systems, and his thoughtful critique of my research. I am also grateful to the personnel both current and past from the Branham and Whiting 4 laboratories for their collection of specimens, useful comments on this manuscript, and encouragement. 5 TABLE OF CONTENTS page ACKNOWLEDGMENTS ...............................................................................................................4 LIST OF TABLES.........................................................................................................................10 LIST OF FIGURES .......................................................................................................................11 ABSTRACT...................................................................................................................................12 CHAPTER 1 PHYLOGENETICS................................................................................................................14 Introduction.............................................................................................................................14 What Types of Data Are Used in Insect Phylogenetics?........................................................15 Morphological Data.........................................................................................................16 Molecular Data ................................................................................................................17 Combined Analysis or Total Evidence............................................................................18 What Are the Major Modern Methodologies Used in Phylogenetics?...................................18 Maximum Parsimony ......................................................................................................19 Statistical Approaches .....................................................................................................20 Maximum likelihood (ML) ......................................................................................20 Bayesian ...................................................................................................................20 What Is Going on in Insect Phylogenetic Studies?.................................................................21 2 SCANNING ROCKS FOR DATA: FOSSIL IMAGING AND DATABASING..................25 Introduction.............................................................................................................................25 Materials and Methods ...........................................................................................................26 Collections Studied..........................................................................................................26 Scanning ..........................................................................................................................26 Macrophotography ..........................................................................................................29 Results.....................................................................................................................................29 Discussion...............................................................................................................................30 Conclusions.............................................................................................................................32 3 ODONATA PHYLOGENY ...................................................................................................36 Introduction.............................................................................................................................36 Materials and Methods ...........................................................................................................39 Taxon sampling ...............................................................................................................39 Laboratory Methods ........................................................................................................40 Data Analysis...................................................................................................................41 Results.....................................................................................................................................43 DO Analysis ....................................................................................................................43 6 Bayesian Analysis ...........................................................................................................44 Parsimony Analyses ........................................................................................................45 Additional Analyses (Fossils Excluded) .........................................................................45 Data Completeness, Support, Congruence and Gene Contribution ................................46 Data completeness:...................................................................................................46 Support: ....................................................................................................................47 Congruence: .............................................................................................................47 Overall Gene Contribution.......................................................................................48 Discussion...............................................................................................................................49 Odonata............................................................................................................................49 Epiprocta..........................................................................................................................50 Anisoptera........................................................................................................................50 Superfamily and Family Monophyly...............................................................................52 Zygoptera.........................................................................................................................54 Superfamily and Family Monophyly...............................................................................55 Contribution of Fossil Taxa.............................................................................................59 Recommendations for Classification Based on the Phylogenies ....................................60 Conclusions.............................................................................................................................60 4 ODONATA WING EVOLUTION.........................................................................................70 Introduction.............................................................................................................................70 Materials and Methods ...........................................................................................................71 Character Optimization (Smart) ......................................................................................71
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