REDEFINITION AND REVISION OF THE GENUS TAENIAPTERA MACQUART, 1835 (DIPTERA: MICROPEZIDAE) A Thesis Presented to The Faculty of Graduate Studies of The University of Guelph by MORGAN D. JACKSON In partial fulfillment of requirements for the degree of Master of Science April, 2011 © Morgan D. 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I+l Canada ABSTRACT REDEFINITION AND REVISION OF THE GENUS TAENIAPTERA MACQUART, 1835 (D1PTERA: MICROPEZIDAE) Morgan D. Jackson Advisors: University of Guelph, 2011 Professor S.A. Marshall Special Graduate Faculty J.H. Skevington The genus Taeniaptera Macquart is redefined and revised following a phylogenetic analysis of select Taeniapterinae. The phylogenetic analysis utilized DNA characters (12S rRNA, cytochrome-c-oxidase subunit I, wingless, and a portion of the c- p-s region of CAD) as well as morphological characters (external structure, male & female genitalia). Data was analyzed primarily using maximum parsimony with Bayesian Inference used for total molecular evidence. The existing Taeniapterine tribal classifications were recovered as paraphyletic. Taeniaptera, Hemichaeta Hennig, ParagraUomyia Hendel and GraUomyia Rondani were all elevated to the generic level. Taeniaptera is redefined to include only species with a closed wing cell T4+5 {Taeniaptera trivittata Macquart, T. lasciva (Fabricius), T. nigriceps Hennig, T. vulpes Cresson, T. knutsonii n. sp., T. longapilis n. sp., and T. promiscua n. sp.), and belong to a larger clade characterized by an axe-shaped maxillary palpus. Taeniaptera is revised with an illustrated key to species and detailed species (re)descriptions. Acknowledgements I'd like to begin by thanking my supervisors, Drs. Stephen A. Marshall and Jeffrey H. Skevington. Steve first introduced me to the fascinating world of systematic entomology during my undergraduate studies, and has provided me with multiple opportunities to explore the never ending diversity of flies through class work and independent study, which has culminated in this study. Steve's encyclopedic knowledge of the lnsecta, endless patience critiquing and improving my writing, and prowess with a camera have all been invaluable to my education and career development. Jeff provided me a glimpse into the world of molecular phylogenetics and the potential gains and pitfalls associated with this field of study, and an alternative yet complimentary approach to systematics and taxonomy. Steve and Jeff also supported my research financially, with NSERC Discovery Grants (SAM, JHS), University of Guelph funding (SAM), AAFC funding (JHS), and NSF funds through the Fly Tree of Life project (JHS). I'd also like to thank my third committee member, Dr. Robert Hanner for yet another viewpoint on the science of taxonomy. I'd like to thank my friends and colleagues in the University of Guelph Insect collection (Matthew Bergeron, Adam Brunke, Dr. Matthias Buck, Phil Careless, Dave Cheung, Gil Goncalves Miranda, Adam Jewiss-Gaines, Joel Kits, John Klymko, Nichelle Lomas, Stephen Luk, Dr. Owen Lonsdale, Steve Paiero, Rob Pivar, and Andrew Young) for their continual support, academic opinions and debate, and for making the insect systematics lab a hell of a lot of fun, even on the most stressful of days. A special thank you to Scott Kelso, who taught me the art of the PCR, the way of the pipette, and showed me how a molecular lab should be run. I owe a large portion of my data to him, and he made my transition to the "dark side" (mostly) painless. Likewise, Joel Gibson was always around to discuss troubles with techniques or new theories. The CNC lunch crew (Drs. Jim O'Hara, Brad Sinclair, Scott Brooks, Jim Cumming, Pat Bouchard and Gary Gibson) provided me with more opinions on entomology and academia than I can remember while also providing a source of daily entertainment as I listened to their "debates". I'd like to thank the University of Guelph and the North American Dipterists Society (NADS) for providing me travel grants to attend the International Congress of Dipterology in Costa Rica and explore the wilderness of Costa Rica in search of more flies. I'd like to thank the institutions that allowed me access to their collections or provided loans of Taeniaptera specimens to Dr. Stephen Marshall: CMNH - Section of Insects and Spiders, Carnegie Museum of Natural History, Pittsburgh, PA; CNCI - Canadian National Collection of Insects, Agriculture and Agri-Food Canada, Ottawa, ON; INPA - Colecao Sistematica da Entomologia, Instituto Nacional de Pesquisas da Amazonia, Manaus; MTEC - Montana State University Entomology Collection, Entomology Research Laboratory, Montana State University, Bozeman, MT; UNCB Museo de Historia Natural, Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Sante Fe' de Bogota', Colombia USNM - United States National Museum, Smithsonian Institute, Washington DC. Last but certainly not least, 1 want to thank my wife, Renee, who has always supported and stood by me. She has put up with my collecting on vacations, long hours working in the lab or at home, and has always been waiting with a smile whenever I return from a collecting trip or conference. m Table of Contents Acknowledgements i Table of Contents iv List of Figures vii List of Tables xiv 1. Introduction 1 1.1 General Biology 1 1.2 Classification of the Taeniapterinae 3 1.4 The Genus Taeniaptera Macquart 1835 4 2. Phylogeny of the Taeniapterinae 7 2.1 Introduction 7 2.2 Methodology - Molecular Phylogeny 8 2.2.1 Taxon Sampling and Storage 8 2.2.2 DNA Extraction 9 2.2.3 DNA Amplification 9 2.2.4 DNA Sequencing and editing 10 2.2.5 Sequence Alignment 11 2.2.6 Concatenated Molecular Dataset Parsimony Analysis 11 2.2.7 Bayesian Analysis 12 2.2.8 Sequence Congruency Test 13 2.2.9 Taxon Sampling - Morphological Parsimony Analysis 14 2.2.10 Matrix Coding and Specimen Preparation 14 2.2.11 Morphological Dataset & Total Evidence Parsimony Analyses 15 iv 2.3 Results 15 2.3.1 Sequencing Success and phylogenetic utility-white, EF-la and 28S 15 2.3.2 12S 16 2.3.3 COI 16 2.3.4 wingless 17 2.3.5 CAD 18 2.3.6 Concatenated Molecular Dataset Parsimony Analysis 18 2.3.7 Molecular Dataset Bayesian Analyses 19 2.3.8 Molecular Data Congruency 19 2.3.9 Morphological Dataset Parsimony Analyses 20 2.3.10 Total Evidence Parsimony Analysis 20 2.4 Discussion of Relationships 21 2.4.1 Total Evidence Parsimony Analysis 21 2.4.2 Bayesian Analysis of Concatenated Molecular Dataset 27 2.4.3 Gene sampling and utility 28 2.5 Taxonomic Recommendations Based on Total Evidence Analysis 30 3. Redefinition of the genus Taeniaptera Macquart 1835 and revision of the Taeniaptera s.s 35 4. Conclusions & Future Work 66 5. Figures & Tables 68 6. References 145 7. Appendices 157 7.1 List of Abbreviations 157 v 7.2 Illustration of Characters used in Morphological Analysis 159 VI List of Figures Figure 1: Graphical representation of Hennig's generic relationships of the Taeniapterinae (Hennig, 1934; 1935a; 1935b; 1936) 68 Figure 2: Phylogeny of the Taeniapterinae from Albuquerque 1986 69 Figure 3: The subgenera of Taeniaptera and included species according to Albuquerque (1986). ** - type species for the subgenus 70 Figure 4: Preferred tree for 12S parsimony analysis (tree length = 798, CI = 0.503, RI = 0.672). Numbers below nodes represent; bootstrap support/Total Bremer support. Numbers after species names are unique specimen identifiers. Coloured boxes represent new generic combinations: Red - Taeniaptera, Green - Grallomyia, Blue - Paragrallomyia, Grey - Poecilotylus 71 Figure 5: Preferred tree for COI HEB parsimony analysis (tree length = 1492, CI = 0.286, RI = 0.478). Numbers above or below nodes represent; bootstrap support/jackknife support/Total Bremer support. Numbers after species names are unique specimen identifiers. Coloured boxes represent new generic combinations: Red - Taeniaptera, Green - Grallomyia, Yellow - Hemichaeta, Blue - Paragrallomyia, Grey - Poecilotylus. 72 Figure 6: Single most parsimonious tree for COI PJ parsimony analysis (tree length = 1737, CI = 0.341, RI = 0.518). Numbers above or below nodes represent; bootstrap support/jackknife support/Total Bremer support.