UNIVERSITY OF CALIFORNIA RIVERSIDE Systematic Research on Minute Litter Bugs Dipsocoromorpha With Emphasis on Schizopteridae (Hemiptera: Heteroptera) A Dissertation submitted in partial satisfaction of the requirements for the degree of Doctor of Philosophy in Entomology by Alexander Knyshov December 2018 Dissertation Committee: Dr. Christiane Weirauch, Chairperson Dr. John Heraty Dr. Timothy Paine Dr. Mark Springer Copyright by Alexander Knyshov 2018 The Dissertation of Alexander Knyshov is approved: Committee Chairperson University of California, Riverside Acknowledgments I would to thank the following individuals and institutions who have kindly loaned or donated the material for my research: Ruth Salas and Toby Schuh (AMNH), Mick Webb (BMNH), Vasily Grebennikov (CNC), Pavel Štys (CUNI), James Boone, Rebekah Baquiran, and Margaret Thayer (FMNH), Susan Halbert (FSCA), Jim Lewis (INBio), Giar-Ann Kung (LACM), Peter Schwendinger and John Hollier (MHNG), Edward Riley (TAMU), Steve Heydon (UCD), Tom Henry (USNM), Serguei V. Triapitsyn and Doug Yanega (UCRC), Edward Riley (TAMU), Fedor Konstantinov (ZISP), Michael Ivie (MTEC), Christopher Grinter (CAS), Norm Penny and Brian Fisher (CAS), Simon van Noort (SAM), Daniela Takiya (UFRJ), and Thailand Inventory Group for Entomological Research (TIGER) Project. Robin Delapena (FMNH) and Walter Winn (FSCA) helped with sorting lots of bulk samples and recovering many of the specimens used in this study. I am grateful to Serguei Triapitsyn and Vladimir Berezovskiy (UCR) for training me and helping with permanent slide-mounting of specimens and István Mikó (Penn State) for advice on confocal microscopy. I thank the Heteropteran Systematics Lab undergraduates Bridget Gonzales, Walena Logan, Brian Vanderveer, Christy Hoong, Ishani Richardson, Joanna Mai, Michelle Ly, Kyle Whorrall for their help with specimen processing, including sorting and databasing. Members of the Weirauch lab, including Rochelle Hoey-Chamberlain, Stephanie Leon, Michael Forthman, Junxia Zhang, Eric Gordon, Paul Masonick, Madison Hernandez, Carlos Rosas, Samantha Smith, Stephanie Castillo, are acknowledged for help throughout my PhD and comments and suggestions on the manuscript drafts. Additionally, members iv of the Heraty lab are thanked for reviewing some of my manuscripts. I am very grateful to Rochelle Hoey-Chamberlain and Eric Gordon for a lot of help with specimen processing and various research ideas. I would also like to thank my dissertation committee members, Dr. John Heraty, Dr. Timothy Paine, and Dr. Mark Springer, for their guidance and help. Especially, I would like to thank Dr. Christiane Weirauch for all the help and support she provided to me during my time at UCR. I am thankful that she has given me an opportunity to extend my PhD research beyond what was originally proposed and experiment with new research ideas. The following funding sources are acknowledged: the US National Science Foundation grant “ARTS: Litter Bugs: revisionary and phylogenetic research on the least studied true bug infraorder (Insecta: Hemiptera: Dipsocoromorpha)” project (DEB-1257702), awarded to Christiane Weirauch; the UCR seed grant “Unlocking the Vault of SoCal Biota” awarded to Christiane Weirauch, Amy Litt, and John Heraty; a Dean’s Distinguished Fellowship, a Dr. Mir S. Mulla and Lelia Mulla Endowed Scholarship, and a UCR Dissertation Research Grant awarded to me. The text of this dissertation, in part or in full, is a reprint of the material as it appears in Knyshov et al. (2016) and Knyshov et al. (2018). The co-author Christiane Weirauch listed in those publications directed and supervised the research which forms the basis for this dissertation. v ABSTRACT OF THE DISSERTATION Systematic Research on Minute Litter Bugs Dipsocoromorpha With Emphasis on Schizopteridae (Hemiptera: Heteroptera) by Alexander Knyshov Doctor of Philosophy, Graduate Program in Entomology University of California, Riverside, December 2018 Dr. Christiane Weirauch, Chairperson Dipsocoromorpha, or the minute litter bugs, are a poorly studied and minuscule group of true bugs (Hemiptera: Heteroptera) with uncertain phylogenetic position, often bizarre morphology, and substantial undescribed biodiversity. A combination of taxonomic revisions, comparative morphological studies, and phylogenetic analyses based on both morphological and molecular data is employed to advance our knowledge of the group. The first chapter taxonomically revises the New World genus Chinannus and describes 26 new species. The second chapter develops and tests a cost-efficient DNA sequencing method for archival specimens. The third chapter uses and refines this method to conduct a comprehensive phylogenetic analysis of a very diverse group of dipsocoromorphans, the Corixidea genus group. The fourth chapter taxonomically revises the genus Voragocoris that belongs to the Corixidea genus group, building on the phylogeny inferred in chapter three, and describing seven new species. The fifth chapter conducts a vi comparative study of abdominal morphology in Dipsocoromorpha, and both standardizes the terminology and proposes primary homology hypotheses, that could be used for a phylogenetic reconstruction. The sixth, and last, chapter presents a comprehensive phylogenetic analysis of Dipsocoromorpha based on both morphological and molecular data. vii Table of contents Introduction ..............................................................................................................1 References ....................................................................................................9 Chapter 1: Systematics of the genus Chinannus Wygodzinsky Abstract ......................................................................................................13 Introduction ................................................................................................14 Material and Methods ................................................................................21 Results and Discussion ..............................................................................30 References ................................................................................................117 Tables and Figures ...................................................................................122 Chapter 2: Sequence capture using PCR-generated baits: a method for cost-efficient and data-rich phylogenies Abstract ....................................................................................................149 Introduction ..............................................................................................150 Material and Methods ..............................................................................155 Results and Discussion ............................................................................163 References ................................................................................................171 Tables and Figures ...................................................................................176 Supporting information ............................................................................180 Chapter 3: Phylogenetic analysis of the Corixidea Reuter genus group Abstract ....................................................................................................203 Introduction ..............................................................................................204 viii Material and Methods ..............................................................................208 Results ......................................................................................................214 Discussion ................................................................................................219 References ................................................................................................223 Tables and Figures ...................................................................................227 Supporting information ............................................................................234 Chapter 4: Taxonomic revision of the genus Voragocoris Weirauch Abstract ....................................................................................................241 Introduction ..............................................................................................241 Material and Methods ..............................................................................243 Results and Discussion ............................................................................245 References ................................................................................................262 Tables and Figures ...................................................................................264 Chapter 5: Comparative morphology of male genitalic structures across Dipsocoromorpha Abstract ....................................................................................................276 Introduction ..............................................................................................277 Material and Methods ..............................................................................280 Results ......................................................................................................284 Discussion ................................................................................................322 Conclusions ..............................................................................................330