PHYLOGENETIC SYSTEMATICS OF THE CERYLONID SERIES OF CUCUJOIDEA (COLEOPTERA) by JAMES A. ROBERTSON (Under the Direction of Joseph V. McHugh) ABSTRACT We conduct a large-scale phylogenetic investigation of the Cerylonid Series (C.S.) of Cucujoidea, a diverse group of cucujoid beetles comprising 9,600 species classified in eight families, using morphological data (76 taxa ! 147 adult and larval characters), molecular data (341 taxa ! 9 genes: 18S, 28S, H3, 12S, 16S, COI, COII, CAD and ArgK) and a combination of the two datasets. In total, our analyses suggest the following: the C.S. is a monophyletic group based on both morphological and molecular evidence; the superfamily Cucujoidea is paraphyletic with respect to the remaining superfamilies in the series Cucujiformia; the C.S. represents a unique clade within Cucujiformia and should be recognized as its own supferfamily, Coccinelloidea, within the series; Byturidae and Biphyllidae should be transferred to Cleroidea; the C.S. families Corylophidae, Coccinellidae, Latridiidae, and Discolomatidae, are monophyletic; Cerylonidae, Endomychidae, and Bothrideridae are paraphyletic. Bothrideridae is split into two distinct families comprising the former Bothriderinae (as Bothrideridae) and the other including the remaining subfamilies (as Teredidae); the cerylonid subfamily Euxestinae is included within Teredidae; the new concept of Cerylonidae includes the following subfamilies: Ceryloninae, Ostomopsinae, Murmidiinae, Discolomatinae and Loeblioryloninae (inserte sedis); the status of the putative new C.S. family, Akalyptoischiidae, is uncertain; the endomychid subfamily Anamorphinae is elevated to familial status, as Anamorphidae. INDEX WORDS: Cucujoidea, Coleoptera, Alexiidae, Bothrideridae, Cerylonidae, Corylophidae, Coccinellidae, Latridiidae, Endomychidae PHYLOGENETIC SYSTEMATICS OF THE CERYLONID SERIES OF CUCUJOIDEA (COLEOPTERA) by JAMES A. ROBERTSON B.S., Brigham Young University, 2002 A Dissertation Submitted to the Graduate Faculty of The University of Georgia in Partial Fulfillment of the Requirements for the Degree DOCTOR OF PHILOSOPHY ATHENS, GEORGIA 2010 © 2010 James A. Robertson All Rights Reserved PHYLOGENETIC SYSTEMATICS OF THE CERYLONID SERIES OF CUCUJOIDEA (COLEOPTERA) by JAMES A. ROBERTSON Major Professor: Joseph V. McHugh Committee: Mark A. Farmer Kenneth G. Ross Electronic Version Approved: Maureen Grasso Dean of the Graduate School The University of Georgia July 2010 DEDICATION This work is dedicated to my dear wife, Emily Robertson, who has believed in me from day one. Without her continued support, love, sacrifice, and taking care of things in the home, I simply could not have accomplished this work. Indeed, the completion of this dissertation is just as much of an accomplishment for her as it is for me. iv ACKNOWLEDGEMENTS I am truly grateful for the many opportunities that I have received throughout my graduate tenure. I have been fortunate to attend and present my research at numerous professional meetings; I was able to participate in a number highly specialized workshops that greatly added to my professional development and research tool belt; I had the opportunity to do fieldwork in remote areas on the planet in search of wonderful beetles; I was able to perform research in a number of excellent research laboratories; I had the opportunity to instruct several Entomology courses and thus gain teaching experience as well as having the experience of working with students. These and other opportunities were made available to me largely due to the thoughtfulness and generosity of others on my behalf. I thank the members of my committee, Joseph McHugh, Mark Farmer, and Kenneth Ross, for their support, guidance, and hand in my professional development. I thank them for their time spent on my behalf. Thanks also goes to Mark Brown, the graduate coordinator, for his encouragement, support, and fair departmental assignments. Likewise the staff in the Entomology department has been very helpful and kind to me during my employment here. I wish to thank the curators and other individuals responsible for the loan of specimens used throughout both the morphological and molecular components of my dissertation research. This research simply could not have occurred with their help. Michael A. Ivie, John F. Lawrence, Adam !lipi"ski, and Joseph McHugh provided assistance and direction with various morphological techniques. v I also wish to thank the fellow graduate students who overlapped with me during my time in the McHugh lab: Floyd Shockley, Juanita Forrester, Adriano Giorgi, Nathan Lord, and Chris Hartley. I thank the above individuals for their kindness, respect, guidance, help with various phases of my research, but most of all for their camaraderie and friendship. Above all, three individuals made this research possible: Adam !lipi"ski who generously provided extremely rare specimens for my use during the morphological component of my research. Likewise, he provided many critical, very difficult to obtain, molecular-grade specimens for the molecular study. I thank him for this generosity, loan and donation of specimens, training and mentoring, and gracious hosting during my stay at CSIRO. This research simply would not have been possible without his immense help. Michael Whiting was extremely generous and gracious with hosting several visits to his lab to generate the molecular data presented in this dissertation. Once again, this research would not have been possible without his great help. Joseph McHugh is truly an amazing person. His kindness, generosity, and mentoring have had a profound effect on me. I am constantly amazed by his kindness to others, professionalism, and productivity. His scientific rigor and standard for excellence in research are outstanding. I thank him for his friendship and advising, supporting my research, providing direction, and his constant patience and understanding. Special thanks also goes to Joe’s family for the sacrifices they have inevitably made by extension of Joe’s efforts in my behalf. I consider Joe a great mentor and very close friend. This research was funded through NSF PEET grant DEB-0329115 (to J.V. McHugh, M.F. Whiting, and K.B. Miller) and NSF AToL grant EF-0531665 to M.F. Whiting (subcontract to J.V. McHugh). Additionally generous support was provided through the H. H. Ross vi Endowment Fund and the Department of Entomology at the University of Georgia for travel to present this research in various stages at professional meetings, to work with Adam !lipi"ski at CSIRO Entomology, Canberra, Australia, and to help support fieldwork. vii TABLE OF CONTENTS Page ACKNOWLEDGEMENTS.........................................................................................................v CHAPTER 1 INTRODUCTION.....................................................................................................1 2 REVIEW OF THE CERYLONID SERIES OF CUCUJOIDEA (COLEOPTERA) ....4 References ...........................................................................................................8 3 SEARCHING FOR NATURAL LINEAGES WITHIN THE CERYLONID SERIES (COLEOPTERA: CUCUJOIDEA) .........................................................................12 Abstract .............................................................................................................13 Introduction .......................................................................................................14 Materials and Methods.......................................................................................16 Results...............................................................................................................19 Discussion..........................................................................................................21 Conclusions .......................................................................................................32 Acknowledgements............................................................................................33 References .........................................................................................................35 Figure Captions..................................................................................................41 4 A MORPHOLOGICAL STUDY AND CLADISTIC ANALYSIS OF THE CERYLONID SERIES (COLEOPTERA: CUCUJOIDEA) .....................................49 Abstract .............................................................................................................50 viii Introduction .......................................................................................................51 Materials and Methods.......................................................................................53 Results...............................................................................................................87 Discussion..........................................................................................................88 Acknowledgements..........................................................................................103 References .......................................................................................................104 Figure Captions................................................................................................108 5 HOST PREFERENCE AND PHYLOGENY OF CUCUJOIDEA (COLEOPTERA)124 Abstract ...........................................................................................................125 Introduction