SYSTEMATICS OF THE ARCHIBORBORINAE (DIPTERA: SPHAEROCERIDAE) A Thesis Presented to The Faculty of Graduate Studies of The University of Guelph by JOEL H. KITS In partial fulfilment of requirements for the degree of Doctor of Philosophy August, 2011 © Joel Kits, 2011 ABSTRACT SYSTEMATICS OF THE ARCHIBORBORINAE (DIPTERA: SPHAEROCERIDAE) Joel H. Kits Advisor: University of Guelph, 2011 Professor Stephen A. Marshall The Archiborborinae comprise a diverse clade of flies in the family Sphaeroceridae. This thesis presents the first phylogenetic analysis and a thorough taxonomic revision of the subfamily. The phylogenetic revision includes morphological data from all species, and molecular data from a subset of 21 ingroup species. Although the group here treated as the Archiborborinae has been traditionally treated as a tribe within the subfamily Copromyzinae, analysis of morphological, molecular, and combined datasets supports the monophyly of the Archiborborinae and shows that the Archiborborinae and Copromyzinae are not sister taxa. The Copromyzinae are more closely related to the Sphaerocerinae and possibly to the enigmatic genus Pycnopota than they are to the Archiborborinae. The elevation of the clade to subfamily rank is supported on the basis of this evidence. Basal relationships within the Archiborborinae are difficult to resolve, but the phylogenetic evidence generally supports a division of the subfamily into the following 8 genera: Antrops Enderlein 1909, Penola Richards 1941, Frutillaria Richards 1961, Boreantrops gen. nov. , Coloantrops gen. nov. , Maculantrops gen. nov. , Photantrops gen. nov. , and Poecilantrops gen. nov. The genus Archiborborus , until recently a paraphyletic assemblage including most of the described species in the subfamily, is treated as a junior synonym of Antrops (syn. nov. ) All genera are described and a generic key is provided. A total of 122 species, including 25 previously described and 89 new, are fully described and illustrated; another 8 new species are diagnosed but not formally named. iii ACKNOWLEDGEMENTS I am grateful to the museum curators who provided loans of material: Jeffrey Cumming and James O’Hara (CNCI), Christophe Daugeron (MNHN), David Grimaldi (AMNH), Carlos Lamas (MZSP), Frank Menzel (DEI), Allen Norrbom (USNM), Mark O’Brien (UMMZ), Gary Parsons (MSUC), James Pitts (EMUS), Adrian Plant (NMWC), Gary Steck (FSCA), Margaret Thayer and James Boone (FMNH), Nigel Wyatt (BMNH), Joachim Ziegler (ZMHB), and Greg Zolnerowich (KSUC). Allen Norrbom was particularly helpful in facilitating transfer loans of material he obtained from a number of collections. René Covarrubias, Gábor Lengyel, Roberto Gonzalez, and Nigel Wyatt provided information on type material. Christian González and Jeffrey Skevington assisted with permits and logistics for collecting trips. All of the collectors who obtained the material used in this revision are appreciated; I especially thank Keith Bayliss and Marc Pollett who provided very useful new material during the course of the project. Joel Gibson and Jeffrey Skevington provided valuable advice during my genetic work. Scott Kelso provided great training and assistance while I was learning molecular techniques. Robert Hanner provided COI barcode sequences from a number of specimens. Matthias Buck provided advice on laboratory techniques. Steve Paiero provided invaluable assistance with curatorial tasks. Adam Jewiss- Gaines databased most of specimens examined for this study. All of the members of the Marshall lab are thanked for support and valuable discussions, particularly David Cheung for his help with technical matters and Gil Goncalves Miranda and Morgan Jackson for advice on phylogenetic analyses. My committee members Robert Hanner, Gard Otis, and Paul Sibley provided valuable advice and feedback. I would particularly like to thank my advisor Stephen Marshall for all of his support during the course of my research. His enthusiasm and passion helped develop my own interest in entomology and inspired me to begin the degree that led to this thesis. He also provided the excellent habitus photos of living specimens included in this thesis. Funding was provided by NSERC discovery grants to Stephen Marshall, OGS and NSERC postgraduate scholarships to JHK, and a grant from the Dipterology Fund to JHK. Sequencing was supported in part by Agriculture and Agri-Food Canada funding. iv Contents Acknowledgements ...................................................................................................................................... iii 1. Introduction ............................................................................................................................................... 1 Research objectives ................................................................................................................................... 4 2. Phylogenetics of the Archborborinae ........................................................................................................ 6 Introduction ............................................................................................................................................... 6 Material and methods ................................................................................................................................ 6 Taxon sampling: .................................................................................................................................... 6 Morphological characters:..................................................................................................................... 6 Molecular characters: ............................................................................................................................ 8 Extraction: ............................................................................................................................................. 8 Amplification and purification: ............................................................................................................. 9 Sequencing: ........................................................................................................................................... 9 Alignment: .......................................................................................................................................... 10 Data analysis: ...................................................................................................................................... 10 Results ..................................................................................................................................................... 12 Discussion ............................................................................................................................................... 20 3. A revision of Frutillaria Richards and Penola Richards (Diptera: Sphaeroceridae: Archiborborinae) . 30 Introduction: ............................................................................................................................................ 30 Material and Methods: ............................................................................................................................ 31 Phylogeny: .............................................................................................................................................. 32 Habitat and biology: ................................................................................................................................ 33 Keys to the known species of Penola Richards and Frutillaria Richards .............................................. 34 4. Revision of the Archiborborinae (other than Frutillaria and Penola ) .................................................... 81 Introduction ............................................................................................................................................. 81 Material and Methods ............................................................................................................................. 81 Key to the genera of the Archiborborinae ............................................................................................... 82 5. Antrops Enderlein 1909 .......................................................................................................................... 84 Key to the species of Antrops ................................................................................................................. 86 Antrops guaramacalensis group ........................................................................................................... 139 Antrops orbitalis group ......................................................................................................................... 154 Antrops quadrilobus group ................................................................................................................... 179 6. Boreantrops gen. nov. ........................................................................................................................... 199 Key to the species of Boreantrops ........................................................................................................ 201 v Boreantrops mexicanus group .............................................................................................................. 204 Boreantrops emarginatus group: .........................................................................................................