A REVIEW OF THE IRONCLAD BEETLES OF THE WORLD (COLEOPTERA ZOPHERIDAE: PHELLOPSINI AND ZOPHERINI) by Ian Andrew Foley A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science in Entomology Montana State University Bozeman, Montana May 2006 COPYRIGHT by © Ian Andrew Foley 2006 All Rights Reserved ii APPROVAL of a thesis submitted by Ian Andrew Foley This thesis has been read by each member of the thesis committee and has been found to be satisfactory regarding content, English usage, format, citations, bibliographic style, and consistency, and is ready for submission to the Division of Graduate Education. Michael A. Ivie, Ph. D. Chair Approved for the Department of Plant Sciences and Plant Pathology John E. Sherwood, Ph. D. Approved for the Division of Graduate Education Joseph J. Fedock, Ph. D. iii STATEMENT OF PERMISSION TO USE In presenting this thesis in partial fulfillment of the requirements for a master’s degree at Montana State University, I agree that the Library shall make it available to borrowers under rules of the Library. If I have indicated my intention to copyright this thesis by including a copyright notice page, copying is allowable only for scholarly purposes, consistent with “fair use” as prescribed in the U.S. Copyright Law. Requests for permission for extended quotation from or reproduction of this thesis in whole or in parts may be granted only by the copyright holder. Ian Andrew Foley May 2006 iv ACKNOWLEDGEMENTS This thesis would not have been possible without the guidance, understanding, and assistance of Michael Ivie, as well as funding for this project through the Montana State Agricultural Experiment Station. Richard Hurley provided much needed relief and support. My committee, Matt Lavin, Richard Miller, and Kevin O’Neill, offered useful suggestions and discussion on drafts of this document. I am thankful to the graduate students, staff, and faculty of the Entomology Graduate program, as well as the Department of Plant Sciences and Plant Pathology at Montana State University, especially past members of the Ivie Lab Katharine Marske, Sardis-Medrano Cabral, J. Joseph Giersch, Alistair Ramsdale and Jessica Fultz, Sarah Wallace and Rebecca Baril. Michael Caterino afforded me the opportunity to learn molecular techniques and use of the SEM at the Santa Barbara Natural History Museum, Chuck Triplehorn, Alfred Newton, and Warren Steiner kindly offered unpublished data on specimens, literature and thoughts on the group. This research would also not have been possible without the timely response to loan request from all of the curators and collection managers who facilitated specimen loans, particularly Maxwell Barclay. Translations of Russian type labels were kindly provided by Alex Konstantinov. Finally, I am grateful for the constant encouragement and support from my parents, John and Mary Ellen, and siblings Aaron, Jenna, and Patrick as well as my friends. v TABLE OF CONTENTS 1. THE FAMILY ZOPHERIDAE .......................................................................................1 Introduction......................................................................................................................1 Diagnostic Morphology ...................................................................................................2 Taxonomic History of the Family Zopheridae.................................................................2 2. REVISION OF THE GENUS PHELLOPSIS..................................................................9 Introduction......................................................................................................................9 Taxonomic History ..........................................................................................................9 Materials ........................................................................................................................12 Methods -- Morphology.................................................................................................16 Methods-- Phylogenetic Analysis..................................................................................18 Taxonomy of the Genus Phellopsis LeConte ................................................................23 Biology of the Genus .....................................................................................................27 Key to the Species of Phellopsis....................................................................................28 Species Descriptions ......................................................................................................30 Phellopsis obcordata (Kirby) ........................................................................................30 Phellopsis porcata (LeConte)........................................................................................35 Phellopsis suberea Lewis ..............................................................................................41 Phellopsis amurensis (Heyden) ....................................................................................45 Phellopsis chinensis Semenow ......................................................................................48 Phellopsis yulongensis Foley and Ivie NEW SPECIES ................................................52 Synoptic Catalog of Phellopsis Species ........................................................................54 Results of Phylogenetic Analysis...................................................................................55 Biogeography.................................................................................................................57 Habitat Conservation .....................................................................................................60 Future Research .............................................................................................................62 3. REVIEW OF THE ZOPHERINI...................................................................................64 Introduction....................................................................................................................64 Zopherini Solier .............................................................................................................65 Taxonomic History of the Zopherini .............................................................................65 Materials ........................................................................................................................68 Methods -- Morphology.................................................................................................68 Methods-- Phylogenetic analysis ...................................................................................70 Results -- Phylogenetic Analysis ...................................................................................76 Key to World Genera of Zopherini................................................................................79 Zopherus Gray ...............................................................................................................82 Synopsis of Zopherus Gray Species ..............................................................................83 vi TABLE OF CONTENTS-CONTINUED Zopherosis White...........................................................................................................85 Synopsis of Zopherosis White Species .........................................................................86 Zopher Ślipiński and Lawrence .....................................................................................86 Synopsis of Zopher Ślipiński and Lawrence Species ...................................................86 Noserinus Casey.............................................................................................................87 Key to Species of Noserinus Casey ..............................................................................88 Synopsis of Noserinus Casey Species............................................................................88 Scoriaderma Fairmaire ..................................................................................................89 Synopsis of Scoriaderma Fairmaire Species .................................................................91 Meralius Casey ..............................................................................................................91 Key to species of Meralius Casey..................................................................................93 Synopsis of Meralius Casey Species .............................................................................93 Meralius clavaplius Foley and Ivie NEW SPECIES.....................................................93 Sesaspis Casey NEW SENSE........................................................................................96 Key to Species of Sesaspis Casey................................................................................100 Synopsis of Sesaspis Casey Species ............................................................................102 Sesaspis denticulata (Solier) NEW COMBINATION ................................................102 Sesaspis emarginatus (Horn) NEW COMBINATION................................................104 Sesaspis doyeni (García-París et al.) NEW COMBINATION ....................................104 Sesaspis lutosus (Champion) NEW COMBINATION................................................108 Sesaspis