MOLECULAR AND MORPHOLOGICAL SYSTEMATICS OF LIBELLULOIDEA (ODONATA: ANISOPTERA) AND DICTYOPTERA by JESSICA LEE WARE A Dissertation submitted to the Graduate School-New Brunswick Rutgers, The State University of New Jersey in partial fulfillment of the requirements for the degree of Doctor of Philosophy Graduate Program in Entomology written under the direction of Michael L. May and Karl Kjer and approved by ________________________ ________________________ ________________________ ________________________ New Brunswick, New Jersey [October, 2008] i ABSTRACT OF THE DISSERTATION MOLECULAR AND MORPHOLOGICAL SYSTEMATICS OF LIBELLULOIDEA (ODONATA: ANISOPTERA) AND DICTYOPTERA By JESSICA LEE WARE Dissertation Director: Michael L. May and Karl M. Kjer Libelluloidea are highly successful dragonflies with unique behavior and life histories. The systematics of Libelluloidea (Odonata: Anisoptera) has historically been in conflict, with little agreement about the number of families that are comprised in this large, heterogeneous group. For my PhD thesis, I have assembled the most comprehensive molecular and morphological libelluloid dataset to date, in an attempt to revise and simplify libelluloid taxonomy, and to answer questions about the evolutionary history of the group. I ran Bayesian and parsimony analyses to recover phylogenetic hypotheses with which I explore the success of Libelluloidea. Divergence estimation, a method by which nodes of a tree are dated, was first explored under different evolutionary models for a subset of libelluloid taxa in order to determine whether treatment of hydrogen-bonded ribosomal nucleotides affected the age of divergence estimates. Using methodology based on these ii results, I was able to estimate divergence dates and diversification rates for the first large-scale dating analysis of dragonflies. On a smaller scale, I also completed a study of Syncordulia , a vulnerable genus of endemic South African libelluloid dragonflies whose systematics was not yet confirmed. Additional studies of phylogenetic methodology were undertaken in my thesis work for another large and heterogeneous group, the Dictyoptera (Mantodea, Blattodea and Isoptera). In this study, the effect of outgroup selection was determined using a broad, comprehensive taxon set for which we had both molecular and morphological data. These results suggest that the evolution of sociality, on which much of the recent discussion in dictyopteran systematics has focused, cannot be reliably determined when different outgroups recover dramatically conflicting topologies. iii Acknowledgement and/or Dedication I would like to thank my husband Jeremy, my daughter Aeshna, my beloved dog Spider and my twin Syrus for their unfaltering support throughout my PhD work. Without them I would had difficulty overcoming the inevitable setbacks, wrong turns and confusion that seem to be a natural part of the academic process. I would also like to thank my best friend and labmate, Dana Price, for her thoughtful conversations about all things entomological and phylogenetic (or otherwise) throughout the last five years. I am deeply indebted to Leslie May for her thoughtfulness, caring, compassion, encouragement, humor, and support, which have become such a welcome aspect of my family’s life here at Rutgers. I would like to thank Karl Kjer and Mike May for their advisement and encouragement regarding both my thesis work and extracurricular affairs. I’m also very grateful to them for supporting my travel to Australia and Namibia, both of which were remarkable opportunities in terms of dragonfly collecting and personal growth. Thank you to Kim Bloodsworth and Dana Price, for their friendship and maternal support since the birth of their goddaughter, Aeshna. Thanks very much to Lashon Ware for her sisterly advice and cheerleading. To my godparents Betty and Archie McGugan, I am thankful for long-distance encouragement and all of the dragonfly books! Caren, Jessica and Leah iv Huff, the Shaws and the Gullettes have been a wonderful source of inspiration and enthusiasm, for which I am very indebted. I would like to acknowledge the Ware Irons family for their insistence, from an early age, that I go to graduate school to get my Ph. D—which I finally did! Last but not least, several fellow dragonfly lovers (enthusiasts and academics alike) were encouraging and supportive throughout my work, for which I am very grateful. I’d like to acknowledge some of them here (although I am thankful to so many more than there is room to list). I am especially grateful to Ken & Sandy Tennessen, Günther & Christine Theischinger, Stas & Lena Gorb and Jerry and Christine Louton, for welcoming me into their homes and providing specimens, advice and discussion; Dennis Paulson, for his thorough and thoughtful commentary about my phylogenies and flight behavior project; Seth Bybee and Erik Pilgrim for advice, discussion, support, encouragement, and for being such wonderful colleagues. Thefollowingchapters were previouslypublishedinpeer-reviewedjournals: Chapter1: Ware,J.L.,M.L.May,K.M.Kjer,2007.Phylogenyofthehigher Libelluloidea(Anisoptera:Odonata):anexplorationofthemostspeciosesuperfamilyof dragonflies. Molecular Phylogenetics and Evolution , 45(1): 289310 Chapter2: Ware,J.L.,Ho,S.Y.W.,Kjer,K.M.2008.Divergencedatesoflibelluloid dragonflies (Odonata:Anisoptera) estimatedfromrRNAusingpaired-sitesubstitution models. Molecular Phylogenetics and Evolution , 47: 426432 Chapter5: Ware,J.L.,Litman,J.Klass,K-D,Spearman,L.2008.Resolvinginternal relationshipswithinthe Dictyoptera posesnewquestions:difficultyinthe placementof Polyphaga. Systematic Entomology, 33 :429450 v This thesis is dedicated to Jeremy Huff, Spider Ware Huff and Aeshna Ware Huff, for all of their patience while I worked on my computer aligning data instead of playing with them outside. vi Table of Contents Abstract………………………………………………………….pages ii-iii Acknowledgements and/or Dedication…….……………………pages iv-vi List of Tables……………………………………………………page ix List of Illustrations………………………………………………pages x-xi Chapter 1: Phylogeny of the higher Libelluloidea (Anisoptera: Odonata): an exploration of the most speciose superfamily of dragonflies. Previously published in Molecular Phylogenetics and Evolution… …………..pages 1-53 Chapter 2: Divergence dates of libelluloid dragonflies (Odonata: Anisoptera) estimated from rRNA using paired-site substitution models. Previously published in Molecular Phylogenetics and Evolution .………….pages 54-78 Chapter 3: Biogeography and divergence estimation of the relic Cape dragonfly genus Syncordulia: global significance and implications for conservation. Submitted to the Journal of Biogeography… …..pages 79-107 Chapter 4: Understanding phylogenetic relationships, homoplasy and divergence estimates within Libelluloidea (Anisoptera: Odonata): an exploration of the usefulness of molecular versus adult and larval morphological characters…………………………………………. .pages 108-200 vii Chapter 5: Resolving internal relationships within the Dictyoptera poses new questions: difficulty in the placement of Polyphaga . Previously published in Systematic Entomology …………………………pages 200-257 Curriculum Vitae……………………………………………...pages 259-260 viii List of tables Table 1, Chapter 1………………….Page 42 Table 2, Chapter 1………………….Page 44 Table 3, Chapter 1………………….Page 53 Table 1, Chapter 2………………….Page 73 Table 2, Chapter 2………………….Page 74 Table S1, Chapter 2………………...Page 75 Table S2, Chapter 2………………...Page 77 Table 1, Chapter 3………………….Page 97 Table 2, Chapter 3………………….Page 101 Table 3, Chapter 3………………….Page 102 Table 1, Chapter 4………………….Page 158 Table 2, Chapter 4………………….Page 185 Table 3, Chapter 4………………….Page 187 Table 4, Chapter 4………………….Page 189 Table 5, Chapter 4………………….Page 189 Table 6, Chapter 4………………….Page 190 Table 7, Chapter 4 …………………Pages 191-198 Table 8, Chapter 4 …………………Pages 198-200 Table 1, Chapter 5………………….Page 254 Table 2, Chapter 5………………….Page 256 Table 3, Chapter 5………………….Page 258 ix List of illustrations Figure 1, Chapter 1………………….Page 34 Figure 2, Chapter 1………………….Page 35 Figure 3, Chapter 1………………….Page 36 Figure 4a, Chapter 1…………………Page 37 Figure 4b, Chapter 1………………...Page 38 Figure 5, Chapter 1………………….Page 39 Figure 6, Chapter 1………………….Page 40 Figure 7, Chapter 1………………….Page 41 Figure 1, Chapter 2………………….Page 71 Figure 2, Chapter 2………………….Page 72 Figure 1, Chapter 3………………….Page 103 Figure 2, Chapter 3………………….Page 104 Figure 3, Chapter 3………………….Page 105 Figure 4, Chapter 3………………….Page 106 Figure 5, Chapter 3………………….Page 107 Figure 1a, Chapter 4…………………Page 151 Figure 1b, Chapter 4…………………Page 152 Figure 2, Chapter 4…………………..Page 153 Figure 3, Chapter 4………………….Page 154 x Figure 4, Chapter 4………………….Page 155 Figure 5, Chapter 4………………….Page 156 Figure 6, Chapter 4………………….Page 157 Figure 1, Chapter 5………………….Page 249 Figure 2, Chapter 5………………….Page 250 Figure 3, Chapter 5………………….Page 251 Figure 4, Chapter 5………………….Page 252 Figure 5, Chapter 5………………….Page 253 xi 1 Chapter 1: Phylogeny of the higher Libelluloidea (Anisoptera: Odonata): an exploration of the most speciose superfamily of dragonflies JessicaWare 1,Michael May 1,andKarlKjer 2 1 Department of Entomology, Rutgers University, 93 Lipman Drive, New Brunswick, NJ, 08901, USA; 2 Department of Ecology, Evolution and Natural Resources, Rutgers University, 14 College Farm Road, New Brunswick, NJ, 08901, USA Abstract Although libelluloiddragonfliesarediverse,numerous,andcommonlyobservedand studied,their phylogenetichistoryis uncertain.Over150yearsoftaxonomicstudyof LibelluloideaRambur1842,beginningwithHagen(1840),Rambur(1842),andSelys