The evolutionary biotogy of Conopidae (Diptera): A life history, molecular, morphological, systematic, and taxonomic approach Joel Francis Gibson B.ScHon., University of Guelph, 1999 M.Sc, Iowa State University, 2002 B.Ed., Ontario Institute for Studies in Education/University of Toronto, 2003 A thesis submitted to the Faculty of Graduate and Postdoctoral Affairs in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Biology Carleton University Ottawa, Ontario © 2011 Joel Francis Gibson Library and Archives Bibliotheque et 1*1 Canada Archives Canada Published Heritage Direction du Branch Patrimoine de Pedition 395 Wellington Street 395, rue Wellington Ottawa ON K1A 0N4 Ottawa ON K1A 0N4 Canada Canada Your Tile Votre r&ference ISBN: 978-0-494-83217-2 Our file Notre reference ISBN: 978-0-494-83217-2 NOTICE: AVIS: The author has granted a non­ L'auteur a accorde une licence non exclusive exclusive license allowing Library and permettant a la Bibliotheque et Archives Archives Canada to reproduce, Canada de reproduire, publier, archiver, publish, archive, preserve, conserve, sauvegarder, conserver, transmettre au public communicate to the public by par telecommunication ou par I'lnternet, preter, telecommunication or on the Internet, distribuer et vendre des theses partout dans le loan, distribute and sell theses monde, a des fins commerciales ou autres, sur worldwide, for commercial or non­ support microforme, papier, electronique et/ou commercial purposes, in microform, autres formats. paper, electronic and/or any other formats. The author retains copyright L'auteur conserve la propriete du droit d'auteur ownership and moral rights in this et des droits moraux qui protege cette these. Ni thesis. Neither the thesis nor la these ni des extra its substantiels de celle-ci substantial extracts from it may be ne doivent etre imprimes ou autrement printed or otherwise reproduced reproduits sans son autorisation. without the author's permission. In compliance with the Canadian Conformement a la loi canadienne sur la Privacy Act some supporting forms protection de la vie privee, quelques may have been removed from this formulaires secondaires ont ete enleves de thesis. cette these. While these forms may be included Bien que ces formulaires aient inclus dans in the document page count, their la pagination, il n'y aura aucun contenu removal does not represent any loss manquant. of content from the thesis. 1+1 Canada Little fly, Thy summer's play My thoughtless hand Has brushed away. Am not I A fly like thee? Or art not thou A man like me? For I dance And drink and sing, Till some blind hand Shall brush my wing. If thought is life And strength and breath, And the want Of thought is death, Then am I A happy fly, If I live, Or if I die. William Blake (1757-1827) Abstract Conopidae is a fascinating family commonly referred to as thick-headed flies. Over 800 species, in over 50 genera and subgenera have been described, but little phylogenetic work has been completed. Past theories on life history, conopid-host interactions, mating strategies, family placement, phylogeny, and classification have been based on very little data and analysis. Based on field observation and collections analysis, evidence of hilltopping mating behaviour is confirmed for thirteen species of Conopidae in the vicinity of Ottawa, Ontario. This represents only the second ever record of this behaviour amongst species of Conopidae. DNA sequence data from ten gene regions is used in a phylogenetic analysis often species of Conopidae and representatives of seventeen other families of Diptera. Parsimony and Bayesian analyses are used to propose a phylogenetic hypothesis that places Conopidae within Schizophora as adelphotaxon to Lauxaniidae. Also, a comparison of the informative qualities of mitochondrial versus nuclear DNA sequence data and ribosomal versus protein coding DNA sequence data is included. Further DNA sequence data from five gene regions and over fifty species of Conopidae are combined with morphological data. A cladogram recovered via parsimony analysis confirms the monophyly of five subfamilies within Conopidae. Relationships between subfamilies are also determined with morphological autapomorphies proposed for all nodes in the cladogram. A phylogenetic analysis including only morphological data for all world genera of Conopidae is completed. The recovered iii cladogram includes six monophyletic subfamilies, two monophyletic tribes within Myopinae, and eleven monophyletic tribes within Conopinae. This phylogenetic hypothesis is used as the basis of a new classification of the genera of Conopidae. Chrysidiomyia Smith, 1989 is placed as a junior synonym of Callosiconops Krober, 1940a, stat. rev. A new genus, Schedophysoconops gen. nov., and subgenus, Asiconops (Aegloconops) subgen. nov., are described. The evolution of body structures within the family is discussed. Biogeographic patterns within subfamilies are noted. The first ever key to world genera of Conopidae is included. IV Acknowledgements Funding was supplied by an NSERC Postgraduate Scholarship, an Ontario Graduate Scholarship, an Entomological Society of Canada Graduate Research- Travel Scholarship, a Biological Survey of Canada Scholarship, a Dipterology Fund Student Research and Travel Grant, a Willi Hennig Society Marie Stopes Travel Award, Carleton University Faculty of Graduate Studies and Research Travel Bursaries, and a Carleton University Biology Department Graduate Scholarship. Funding and equipment were also supplied by J.H. Skevington from his Agriculture and Agri-Food Canada Research budget, an NSERC Discovery Grant, and an NSF grant to B. Wiegmann (FLYTREE AToL, EF-0334948). Funding and travel costs were also supplied by S.B. Peck from his NSERC Discovery Grant. My thanks go to Dr. Skevington and Prof. Peck for not only funding my research, but also for providing insight and guidance. Thanks as well to Prof. F. Chapleau, the other member of my advisory committee, who likewise provided insight at important stages of my research. I was extremely fortunate to be able to meet and consult with Dr. Sidney Camras of the Field Museum of Natural History in Chicago, IL. I am grateful for the chance to work with and gain direct insight from one of the true fathers of conopidology. I also had the opportunity to collaborate with M. Mei and F.C. Thompson, two other eminent scientists investigating Conopidae. Thanks to M. Mei, M. Irwin, S. Winterton, S. Gillespie, N. Cordes, M. Hauser, K. Bayless, M. Locke, J. Cumming, S. Cumming, H. Cumming, S. Brooks, J. Corrigan, M. v Foldvari, M. Jackson, C. Kehlmaier, 0. Lonsdale, J. O'Hara, S. Marshall, M. Pollet, T. Wheeler, S. Gaimari, J. Savage, and B. Sinclair for providing specimens and/or identifications. Thanks, as well, to my contacts at all insect collections that facilitated visits and/or loans of materials. C. Lewis provided assistance with programming MrBayes. M. Sorenson provided assistance with determining branch supports for trees. Portions of this thesis, as submitted for publication, were improved based on editorial input from G. Capretta, J. O'Hara, A. Vogler, S. Gaimari, and S. Cameron, as well as numerous anonymous reviewers. Special thanks go to my molecular lab colleagues. M. Jackson, J. Kits, G.F.G. Miranda, and W. Knee. Their words of caution, innovation, and sympathy were much appreciated during long hours of pipetting, aligning, and analyzing. The utmost appreciation is extended to S. Kelso, the guru and mentor of the molecular lab, without whom no sequence is possible. Special thanks as well are extended to colleagues in the Canadian National Collection of Insects, Arachnids, and Nematodes. M. Locke, S. Brooks, B. Sinclair, 0. Lonsdale, J. O'Hara, P. Bouchard, J. Cumming, and G. Gibson all provided valuable insight into all matters taxonomic, morphological, editorial, phylogenetic, and budgetary. Finally, and most importantly, my thanks are extended to my family. My parents, Frank and Marina, and my brothers, Ben and Will, offered constant encouragement and support. My daughters, Anabel and Elsa, provided the most essential part of this project: unfiltered joy and love. My wife, Gina, supplied unending support, patience, insight, inspiration, and love. No mere words could ever express my eternal gratitude. vi Table of Contents List of tables ix List of figures x Preface xiv 1. Introduction to Conopidae 1.1. The use of Conopidae in biological research 1 1.2. Previous systematic research on Conopidae 4 1.3. Molecular and morphological methodology employed 6 1.4. Organization of thesis 10 2. Hilltopping in species of Conopidae in the Ottawa area 2.1. Introduction 12 2.2. Materials and methods 13 2.3. Results 14 2.4. Discussion 16 3. Placement of Conopidae (Diptera) within Schizophora based on mtDNA and nrDNA gene regions 3.1. Introduction 19 3.2. Materials and methods 23 3.3. Results 27 3.4. Discussion 33 vii 4. Phylogenetic analysis of relationships between genera of Conopidae based on molecular and morphological data 4.1. Introduction 43 4.2. Materials and methods 45 4.3. Results 73 4.4. Discussion 79 5. Revision of the genera of Conopidae based on morphological data 5.1. Introduction 99 5.2. Materials and methods 103 5.3. Results 128 5.4. Discussion 137 5.5. Key to world genera of Conopidae 178 6. General conclusion 197 General bibliography 203 Tables 246 Figures 283 Vlll List of tables Table 2.1. Specimens of Conopidae (Diptera) from the CNC collected in the Ottawa area, noting those collected on hilltops 246 Table 3.1. List of taxa included in Chapter 3 analysis with GenBank accession numbers 247 Table 3.2. Primer oligonucleotides used for PCR amplification of selected gene segments in Chapter 3 249 Table 3.3. Summary of results for individual and concatenated gene partitions 250 Table 4.1. List of taxa included in Chapter 4 analysis with GenBank accession numbers 252 Table 4.2. Primer oligonucleotides used for PCR amplification of selected gene segments in Chapter 4 255 Table 4.3.