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Diptera: Mycetophilidae) SYSTEMATICS OF LEPTOMORPHUS AND PHYLOGENY OF THE SCIOPHILINI (DIPTERA: MYCETOPHILIDAE) Christopher James Borkent Department of Natural Resource Sciences McGill University, Montreal June 2012 A thesis submitted to McGill University in partial fulfillment of the requirements of the degree of Doctor of Philosophy © Christopher James Borkent, 2012 ABSTRACT A taxonomic revision and phylogenetic analysis of the world fauna of Leptomorphus Curtis (Diptera: Mycetophilidae) were undertaken. Forty-five valid species of Leptomorphus are recognized, including 12 new species and three new synonyms. Species descriptions, diagnoses and illustrations are provided for 37 species, along with a key to adults. Eight recently described Oriental and Australasian species for which material was unavailable are excluded from the key. The phylogenetic analysis used 73 adult morphological characters and supported the monophyly of Leptomorphus. Four major clades were found within the genus, but no support was found for previously recognized subgenera. A phylogenetic analysis was also conducted at the generic level for the tribe Sciophilini, to which Leptomorphus belongs. The analysis included 81 exemplar species, representing all but one of the genera currently or previously placed in the Sciophilini, scored for 96 adult morphological characters. The resulting phylogeny showed a monophyletic Sciophilini supported by four synapomorphies and containing 34 monophyletic genera: Acnemia Winnertz, Acomoptera Vockeroth, Adicroneura Vockeroth, Afrocnemia Matile, Allocotocera Mik, Anaclileia Meunier, Aneura Marshall, Austrosciophila Tonnoir, Azana Walker, Baeopterogyna Vockeroth, Cluzobra Edwards, Drepanocercus Vockeroth, Duretophragma Borkent gen. n., Eudicrana Loew, Leptomorphus Curtis, Loicia Vockeroth, Megalopelma Enderlein, Monoclona Mik, Morganiella Tonnoir & Edwards, Neoallocotocera Tonnoir, Neoaphelomera Miller, Neotrizygia Tonnoir & Edwards, Neuratelia Rondani, Paramorganiella Tonnoir, Paratinia Mik, Paratrizygia Tonnoir, Parvicellula Marshall, Phthinia Winnertz, Polylepta Winnertz, Sciophila Meigen, Stenophragma Skuse, Tasmanina Tonnoir, Taxicnemis Tonnoir & Edwards, and Trizygia Skuse. Four genera which had been placed in the Sciophilini by some previous authors (Coelophthinia Edwards, Impleta Plassmann, Speolepta Edwards, and Syntemna Winnertz) were excluded from the tribe and placed in the Gnoristini. New generic diagnoses were given for ii all genera in the tribe. The new genus Duretophragma, formerly constituting part of the paraphyletic genus Stenophragma, is described and illustrated and 10 new generic combinations resulted from recognition of the new genus. iii RÉSUMÉ Une révision taxonomique et une analyse phylogénétique de la faune mondiale du genre Leptomorphus Curtis (Diptera: Mycetophilidae) ont été effectuée. Quarante-cinq espèces valides sont reconnues, incluant 12 nouvelles espèces et trois nouveaux synonymes. Des descriptions, caractères diagnostiques et illustrations sont présentés pour 37 espèces, de même qu'une clef d'identification des adultes. Huit espèces orientales et australasiennes sont exclues de la clef parce que des spécimens représentant ces espèces n'étaient pas disponibles. L'analyse phylogénétique, basée sur 73 caractères morphologiques des adultes, supporte la monophylie de Leptomorphus et met en évidence quatre clades, mais ne soutient pas les sous-genres reconnus antérieurement. Une analyse phylogénétique de la tribu à laquelle appartient le genre Leptomorphus, Sciophilini, a également été effectuée, basée sur 96 caractères morphologiques des adultes provenant de 81 espèces exemplaires représentant tous les genres compris antérieurement ou présentement dans la tribu, sauf un. Quatre synapomorphies corroborent la monophylie de Sciophilini qui contient 34 genres également monophylétiques: Acnemia Winnertz, Acomoptera Vockeroth, Adicroneura Vockeroth, Afrocnemia Matile, Allocotocera Mik, Anaclileia Meunier, Aneura Marshall, Austrosciophila Tonnoir, Azana Walker, Baeopterogyna Vockeroth, Cluzobra Edwards, Drepanocercus Vockeroth, Duretophragma Borkent gen. n., Eudicrana Loew, Leptomorphus Curtis, Loicia Vockeroth, Megalopelma Enderlein, Monoclona Mik, Morganiella Tonnoir et Edwards, Neoallocotocera Tonnoir, Neoaphelomera Miller, Neotrizygia Tonnoir et Edwards, Neuratelia Rondani, Paramorganiella Tonnoir, Paratinia Mik, Paratrizygia Tonnoir, Parvicellula Marshall, Phthinia Winnertz, Polylepta Winnertz, Sciophila Meigen, Stenophragma Skuse, Tasmanina Tonnoir, Taxicnemis Tonnoir et Edwards, et Trizygia Skuse. Quatre genres classés par certains auteurs dans Sciophilini sont ici exclus de la tribu et placés dans Gnoristini: Coelophthinia Edwards, Impleta Plassmann, Speolepta Edwards, et Syntemna Winnertz. Des nouveaux caractères diagnostiques sont iv présentés pour tous les genres inclus dans la tribu. L'auteur propose un nouveau genre, Duretophragma, une fraction du genre paraphylétique Stenophragma, établissant ainsi 10 nouvelles combinaisons au niveau générique. Une description et illustrations du genre sont présentées. v TABLE OF CONTENTS ABSTRACT ............................................................................................................ ii RÉSUMÉ ............................................................................................................... iv LIST OF FIGURES ............................................................................................... ix LIST OF TABLES ............................................................................................... xix ACKNOWLEDGMENTS .....................................................................................xx PREFACE ........................................................................................................... xxii CONTRIBUTION OF AUTHORS.................................................................... xxiii CLAIM TO ORIGINALITY ............................................................................. xxiv CHAPTER 1. GENERAL INTRODUCTION AND LITERATURE REVIEW ............................................................................1 Systematics and diversity of Diptera .......................................................................1 Higher level phylogenetics of the Diptera ...............................................................3 Morphological studies within the Diptera................................................................4 Molecular studies within the Diptera .......................................................................5 Combined potential of molecules and morphology .................................................8 Phylogeny and classification of nematocerous Diptera .........................................10 Biology of the Mycetophilidae ..............................................................................12 Review of Mycetophilidae classification ...............................................................13 The tribe Sciophilini ..............................................................................................15 The genus Leptomorphus Curtis ............................................................................16 Objectives ..............................................................................................................18 Literature Cited ......................................................................................................19 CONNECTING STATEMENT .............................................................................35 CHAPTER 2. SYSTEMATICS AND PHYLOGENY OF LEPTOMORPHUS CURTIS (DIPTERA: MYCETOPHILIDAE) ...........................................36 Abstract ..................................................................................................................36 Table of Contents ...................................................................................................38 Introduction ............................................................................................................40 Taxonomic history .................................................................................................40 Materials and methods ...........................................................................................43 vi Distributions .......................................................................................................43 Measurements ....................................................................................................43 Terminology .......................................................................................................44 Dissections .........................................................................................................44 Phylogeny ..........................................................................................................44 Systematics of the Genus Leptomorphus Curtis ....................................................45 Diagnosis............................................................................................................45 Generic description ............................................................................................46 Biology ...............................................................................................................51 Key to adults of Leptomorphus species .................................................................53 Species descriptions ...............................................................................................66
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