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Diptera: Tachinidae) Jeremy Daniel Blaschke University of Tennessee - Knoxville, [email protected]

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University of Tennessee, Knoxville Trace: Tennessee Research and Creative Exchange Doctoral Dissertations Graduate School 8-2015 Evolution and Phylogeny of the Parasitoid Subfamily Phasiinae (Diptera: Tachinidae) Jeremy Daniel Blaschke University of Tennessee - Knoxville, [email protected] Recommended Citation Blaschke, Jeremy Daniel, "Evolution and Phylogeny of the Parasitoid Subfamily Phasiinae (Diptera: Tachinidae). " PhD diss., University of Tennessee, 2015. https://trace.tennessee.edu/utk_graddiss/3399 This Dissertation is brought to you for free and open access by the Graduate School at Trace: Tennessee Research and Creative Exchange. It has been accepted for inclusion in Doctoral Dissertations by an authorized administrator of Trace: Tennessee Research and Creative Exchange. For more information, please contact [email protected]. To the Graduate Council: I am submitting herewith a dissertation written by Jeremy Daniel Blaschke entitled "Evolution and Phylogeny of the Parasitoid Subfamily Phasiinae (Diptera: Tachinidae)." I have examined the final electronic copy of this dissertation for form and content and recommend that it be accepted in partial fulfillment of the requirements for the degree of Doctor of Philosophy, with a major in Plants, Soils, and Insects. John K. Moulton, Major Professor We have read this dissertation and recommend its acceptance: Ernest Bernard, Rebecca Nichols, Brian O'Meara Accepted for the Council: Dixie L. Thompson Vice Provost and Dean of the Graduate School (Original signatures are on file with official student records.) Evolution and Phylogeny of the Parasitoid Subfamily Phasiinae (Diptera: Tachinidae) A Dissertation Presented for the Doctor of Philosophy Degree The University of Tennessee, Knoxville Jeremy Daniel Blaschke August 2015 ii Copyright © 2015 by Jeremy Daniel Blaschke All rights reserved. iii Acknowledgements First, this thesis would not have been possible without the support and direction of my major professor, Kevin Moulton. Through my time at UTK my understanding of molecular systematics and evolutionary biology has been developed and I have learned valuable techniques and skills that will aid my future research and teaching endeavors. I am very grateful to my committee members, Ernie Bernard, Becky Nichols, Brian O’Meara, and Jim O’Hara for their time, counsel, and encouragement. Reliance on fellow researchers and collaborators was essential for obtaining the rare and/or international genera for this study. John Stireman, Isaac Winkler, and the entire Stireman Lab at Wright State University, Jim O’Hara and Monty Wood from the Canadian National Collection, and Pierfilippo Cerretti from Sapienza Università di Roma provided collections from several international trips as well as their own local habitats. They have also provided valuable advice and resources on tachinid classification, identification, and history. Additional specimens or sequences were provided by Takuji Tachi, Martin Hauser, Thibault Ramage, Jaako Pojoismaki, and many others. Funding for this research was provided by the National Science Foundation Award DEB-1146290, “Collaborative Research: The Phylogeny and Evolution of World Tachinidae (Diptera)”, awarded to J.K. Moulton. Additionally, I would like to thank the Adams family for their love and support, patience, and interest (however forced) in my work. Finally, I would like to thank my wonderful wife Ally and our dear dissertation baby Jack. I am truly grateful for Ally’s love, encouragement, and the hard work and sacrifice she has made to make this degree possible for me. I look forward to many future adventures with them by my side. iv Abstract The first molecular phylogenetic analysis of the agriculturally important parasitoid subfamily Phasiinae (Diptera: Tachinidae) is presented, estimated from 128 worldwide taxa (80 genera) and approximately 7.6 kilobases of nuclear data. Special emphasis is placed on taxa with controversial taxonomic placement. The resultant phylogenetic tree is used to reconstruct ancestral character states, trace the evolution of significant adaptive traits within the Tachinidae, and test hypotheses about the classification of Phasiinae. Subfamily placements of the taxa Eutherini, Epigrimyiini, Litophasia , Strongygastrini, and Parerigonini are confidently resolved, the former three within Dexiinae and the latter two within Phasiinae. Due to sparse molecular evidence, the Imitomyiini are tentatively placed among the Phasiinae. Ancestral state reconstruction suggests a dominant and persistent trend in Phasiinae to evolve piercing structures used to insert eggs directly into host tissues. A single potential synapomorphy of Phasiinae is identified (elongated hypandrium). This phylogeny is used to update classification of worldwide phasiine genera and tribes. Many novel phylogenetic hypotheses are presented including the division of Parerigonini s. l. into three tribes: Parerigonini s. s. , Zitini, and Cylindromyiini, and the division of Phasiini s. l. into four lineages: Phasiini s. s. , Gymnosomatini, Opesiini, and Xystini. Two tribes are resurrected (Opesiini and Xystini) and one new tribe is proposed (Zitini nomen novum ). Additionally, a survey of phasiine biodiversity was conducted in Great Smoky Mountains National Park (TN, NC). Species identifications were made using morphological keys, with further evidence from 900 base pairs of the nuclear coding gene MCS. In total, 221 specimens representing 26 phasiine species were collected. Of these, 21 species are newly recorded from the park, four are new records for Tennessee, and two are new records for North Carolina. All 12 eastern Nearctic phasiine genera were represented. Updated identification keys to eastern Phasiinae are provided and DNA barcoding sequences were generated that will aid future researchers to quickly and inexpensively identify phasiine species. v Table of Contents Introduction ..................................................................................................................................... 1 Chapter I: Evolution and Phylogeny of the Parasitoid Subfamily Phasiinae (Diptera: Tachinidae) ......................................................................................................................................................... 2 Abstract ............................................................................................................................... 3 Introduction and Background ............................................................................................. 4 Tachinidae ............................................................................................................... 5 Phasiinae ................................................................................................................. 6 Taxonomy ............................................................................................................... 7 Objectives ............................................................................................................. 10 Materials and Methods ...................................................................................................... 11 Collection .............................................................................................................. 11 Genes..................................................................................................................... 15 Extraction and Amplification ................................................................................ 16 Phylogenetic Analysis ........................................................................................... 19 Ancestral State Reconstruction ............................................................................. 20 Results ............................................................................................................................... 24 Sequence data........................................................................................................ 24 Trees ...................................................................................................................... 24 Outgroups .............................................................................................................. 24 Tachininae + Exoristinae ...................................................................................... 29 Dexiinae ................................................................................................................ 29 Phasiinae ............................................................................................................... 30 Taxonomically ambiguous genera ........................................................................ 33 Clade frequencies across all analyses ................................................................... 33 Phylogenetic informativeness of genes and codon positions ................................ 34 Discussion ......................................................................................................................... 34 Classification......................................................................................................... 36 Evolution of Tachinidae ........................................................................................ 41 Conclusions ........................................................................................................... 51 List of References ............................................................................................................
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