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Taxonomy and Phylogeny of Piophilidae (Diptera)

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Taxonomy and Phylogeny of Piophilidae (Diptera) Taxonomy and Phylogeny of Piophilidae (Diptera) Sabrina Rochefort Department of Natural Resource Sciences McGill University, Montreal August 2015 A thesis submitted to McGill University in partial fulfillment of the requirements of the degree of Master of Science © Sabrina Rochefort, 2015 ABSTRACT The worldwide generic classification of Piophilidae (Diptera) is tested using a morphological and molecular phylogenetic analysis, and the Nearctic species of the family are revised. The taxonomic revision includes geographic distributions, capture notes, species descriptions and an identification key to the 43 Nearctic species. Based on the phylogenetic analysis, 20 genera are recognized in the family. Five genera are synonymized: Neopiophila McAlpine, Boreopiophila Frey and Parapiophila McAlpine with Arctopiophila Duda; Neottiophilum Frauenfeld with Mycetaulus Loew; and Stearibia Lioy with Prochyliza Walker. One new Holarctic genus, Borealicola, is described, and a second new genus, not described in this thesis, is recognized for the Australian species Protopiophila vitrea McAlpine. Four new species are described: Arctopiophila mcalpinei, A. variefrontis, Borealicola madaros, and B. skevingtoni. Eighteen new combinations are proposed: Arctopiophila atrifrons (Melander & Spuler), A. baechlii (Merz), A. dudai (Frey), A. flavipes (Zetterstedt), A. kugluktuk (Rochefort & Wheeler), A. lonchaeoides (Zetterstedt), A. nigritellus (Melander), A. nitidissima (Melander & Spuler), A. pectiniventris (Duda), A. penicillata (Steyskal), A. setaluna (McAlpine), A. tomentosa (Frey), A. vulgaris (Fallén), A. xanthostoma (Melander & Spuler), Borealicola fulviceps (Holmgren), B. pseudovulgaris (Ozerov), Mycetaulus praeustum (Meigen) and Prochyliza nigriceps (Meigen). ii RÉSUMÉ La classification mondiale des genres appartenant à la famille des Piophilidae (Diptère) est examinée à l’aide d’une analyse phylogénique incluant des caractères morphologiques et moléculaires, et les espèces Néarctique de la famille sont révisées. Cette dernière inclus des distributions géographiques, des notes sur les méthodes de captures, des descriptions d’espèces et d’une clé d’identification des 43 espèces Néarctique. Suite à l’analyse phylogénique mondiale, 20 genres sont reconnus dans la famille. Cinq genre sont devenus des synonymes: Neopiophila McAlpine, Boreopiophila Frey et Parapiophila McAlpine avec Arctopiophila Duda; Neottiophilum Frauenfeld avec Mycetaulus Loew; et Stearibia Lioy avec Prochyliza Walker. Un nouveau genre Holarctique, Borealicola, est décrit, et un second nouveau genre, non décrit dans cette thèse, est reconnu pour l’espèce Australienne Protopiophila vitrea McAlpine. Quatre nouvelles espèces sont décrites: Arctopiophila mcalpinei, A. variefrontis, Borealicola madaros, et B. skevingtoni. Il y a, de plus, dix-huit nouvelles combinaisons: Arctopiophila atrifrons (Melander & Spuler), A. baechlii (Merz), A. dudai (Frey), A. flavipes (Zetterstedt), A. kugluktuk (Rochefort & Wheeler), A. lonchaeoides (Zetterstedt), A. nigritellus (Melander), A. nitidissima (Melander & Spuler), A. pectiniventris (Duda), A. penicillata (Steyskal), A. setaluna (McAlpine), A. tomentosa (Frey), A. vulgaris (Fallén), A. xanthostoma (Melander & Spuler), Borealicola fulviceps (Holmgren), B. pseudovulgaris (Ozerov), Mycetaulus praeustum (Meigen) et Prochyliza nigriceps (Meigen). iii TABLE OF CONTENTS Abstract............................................................................................................................................ii Résumé...........................................................................................................................................iii List of figures..................................................................................................................................vi List of appendices.........................................................................................................................viii Acknowledgements.........................................................................................................................ix Preface..............................................................................................................................................x Contributions of authors.................................................................................................................xi CHAPTER 1. Introduction and literature review......................................................................1 Introduction......................................................................................................................................1 Piophilidae as study organisms........................................................................................................2 Evolutionary relationships of Piophilidae........................................................................................3 Piophilidae classification: McAlpine’s (1977) and Ozerov’s (2004) view…….........................…4 The need for a new classification and taxonomic revision..............................................................5 Objectives........................................................................................................................................6 References........................................................................................................................................8 Connecting statement.....................................................................................................................11 CHAPTER 2. Phylogeny of the family Piophilidae and revision of the Nearctic species.....12 Introduction....................................................................................................................................12 Materials and methods...................................................................................................................14 Specimen examination and description..............................................................................14 Selection of outgroups and exemplar species for phylogenetic analysis...........................15 Character states for morphological phylogenetic analysis.................................................19 Morphological phylogeny..................................................................................................21 Molecular phylogeny analysis...........................................................................................22 Results and discussion...................................................................................................................22 Phylogeny of Piophilidae: Morphological characters........................................................22 Phylogeny of Piophilidae: Molecular characters...............................................................27 Taxonomic revision of Nearctic Piophilidae species.........................................................28 Species descriptions...............................................................................................28 iv Identification key to Nearctic Piophilidae............................................................131 Revised classification of Piophilidae...................................................................136 References....................................................................................................................................165 CHAPTER 3. General conclusion............................................................................................175 References....................................................................................................................................176 v LIST OF FIGURES Figure 1. One of the 30 most parsimonious trees based on morphology. Species names follow literature prior to revised classification in this thesis. Dark squares = uniquely derived character states; white squares = homoplasious character states. Bremer values are shown below branches........................................................................................................................143 Figure 2. One of the 30 most parsimonious trees based on morphology. Species names follow literature prior to revised classification in this thesis. Dark squares = uniquely derived character states; white squares = homoplasious character states. Bremer values are shown below branches........................................................................................................................144 Figure 3. Majority consensus tree (70%) of 30 most parsimonious trees. Percentage frequency values are shown below branches...........................................................................................145 Figure 4. Maximum likelihood tree of Piophilidae exemplar species using 483 base pairs of Cytochrome Oxidase 1 (CO1). Bootstrap values above branches..........................................146 Figure 5. Maximum likelihood tree of Piophilidae exemplar species using 658 base pairs of Cytochrome Oxidase 1 (CO1). Bootstrap values above branches..........................................147 Figure 6. Nearctic distribution of Actenoptera avalona..............................................................148 Figure 7. Nearctic distribution of Actenoptera hilarella.............................................................148 Figure 8. Nearctic distribution of Allopiophila luteata (circles) and A. testacea (triangles).......149 Figure 9. Nearctic distribution
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