SYSTEMATIC STUDIES OF THE INDO-AUSTRALIAN CROWNED WEEVILS (COLEOPTERA: CURCULIONIDAE: CRYPTORHYNCHINAE) A DISSERTATION SUBMITTED TO THE FACULTY OF THE GRADUATE SCHOOL OF THE UNIVERSITY OF MINNESOTA BY Gregory Peter Setliff IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY Susan J. Weller, George D. Weiblen, Co-Advisers August 2009 © Gregory P. Setliff 2009 ACKNOWLEDGEMENTS I thank the many curators and colleagues that loaned material to me from their respective collections. I wish to acknowledge the assistance of my friends and colleagues at the New Guinea Binatang Research Center during my field studies in Papua New Guinea. I especially thank Vojtech Novotny, Jiri Hulcr, Scott Miller, George Weiblen and the parataxonomists. I also wish to acknowledge the PNG National Agricultural Research Institute, PNG National Research Institute, and the PNG Department of Environment and Conservation for their assistance in my field work. I am grateful to the people of Baitabag, Mis, Ohu, and Wannang villages for their efforts to preserve their forests and for allowing me to collect on their land. Margaret Borg, Linda Eells, Kristina Halfmann, and Anne Rojas of the Entomology Fisheries and Wildlife library (University of Minnesota) labored through hundreds of entomological journals on my behalf, never failing to find references, no matter how obscure or incomplete the citation. Mara Egger (USNM volunteer), John Luhman (Minnesota Department of Agriculture), and Steffen Pauls (UMSP) provided valuable translation services. Janet Moe (University of Minnesota) assisted with the printing and assembling of draft versions of this thesis. University of Minnesota undergraduate assistants, Peter Larson, Crystal Boyd, and Benjamin Dill helped record and process data for the project. I am very much indebted to my “academic family”, Michelle DeCosta, Amanda Roe, Becky Simmons, Jen Zaspel and Peter Larson and Crystal Boyd (Weller lab), Wendy Clement and Summer Silvieus (Weiblen lab), Desi Robertson, Lourdes Chamorro de Lacayo, Roger Blahnik, Anne Wasmund, Steffen Paules, and Patina Mendez (Holzenthal lab), and Philip Calusen and John Luhman (UMSP insect collection). These lab mates, co-workers, collaborators and friends have all provided support and encouragement and each have contributed to my education as an entomologist and a systematist. I wish to express my sincerest appreciation and deepest regard for my co-advisors, Susan Weller and George Weiblen i and committee members, Ralph Holzenthal and Sharon Jansa. I am thankful for the guidance that they provided me in virtually every aspect of this project and in my personal development as a scientist. They provided me with lab space, equipment, funding, manuscript reviews, and training and advice on every imaginable topic. Moreover, they welcomed me into their homes as well as their labs and for this, I am extremely grateful. Finally, I thank my parents, Pete and Faye Setliff, my mother-in-law Susan Chamberlain, and my wife Julie for their enduring love, unwavering support, and patience during this time. ii DEDICATION I dedicate this work to the memory of Faye Setliff, whom we lost during its writing, to Lexi and Vivien, whom we found, and to Julie and Susan who were with me all along. iii ABSTRACT This dissertation focuses on the systematics of weevils (Coleoptera: Curculionidae) from the Indo-Australian tropics. Special consideration is given to a novel group of seven genera of Cryptorhynchinae that comprise the crowned weevil group, so named for the crown-like carina or glabrous semicircular bulge on the vertex on the head of these weevils, which is one of the more salient features shared by these genera. Herein, the phylogeny of the largest crowned weevil genus, Asytesta Pascoe, is reconstructed with parsimony and Bayesian analyses based on a data set of 82 adult morphological characters (187 states) for 40 ingroup taxa. One species of Cyamobolus Schönherr and three species of Cyamotrox Heller are used as outgroups and all six genera and 14 species of the crowned weevil group as redefined here (Cyamomistus Heller, Eudyasmus Pascoe, Glochinorhinus Waterhouse, Nothotragopus Zimmerman, Panopides Pascoe, and Zygara Pascoe), are used to test the monophyly of Asytesta. The results did not support the monophyly of the Asytesta due to the nesting of monotypic Zygara within an apical subclade of Asytesta. Accordingly, Zygara becomes a new junior synonym of Asytesta and Zygara doriae (Kirsch) is returned to its original combination with Asytesta; A. doriae Kirsch status revised. With the inclusion of Zygara, Asytesta as redefined here is monophyletic. Resolution within Asytesta was poor; however, three subclades with strong support were recovered and are formally recognized as species groups. The analyses also recovered monophyletic Nothotragopus, Panopides, and Glochinorhinus. Relationships among the genera were not resolved. The monophyly of Eudyasmus was not supported. In a strict consensus of all trees recovered, Eudyasmus collapsed into a polytomy with Glochinorhinus. A revision of Asytesta is also presented. Of the 41 species recognized here, 23 are previously described and 18 are new: A. alexanderiae, A. alexriedeli, A. allisoni, A. iv biakana, A. cheesmanae, A. concolora, A. emarginata, A. fayae, A. frontalis, A. gressitti, A. julieae, A. marginalis, A. morobeana, A. sedlaceki, A. thompsoni, A. tuberculata, A. vivienae, and A. woodlarkiana, new species. Asytesta bidentata Voss new status, is elevated to species status from a subspecies of A. lugubris Heller. Four Asytesta species are synonymized: A. circulifera Lea, 1928 = A. rata Heller, 1910, A. definita Faust, 1898 = A. humeralis Pascoe, 1865, A. granulifera Lea, 1928 = A. aucta Faust, 1898, and A. setipes Lea, 1928 = A. lugubris Heller, 1895 new synonyms. Two previously described species were misplaced in Asytesta and are transferred to other genera as follows: A. maura Pascoe to Microporopterus Lea and A. ypsilon Heller to Meroleptus Faust, new combinations. Three species groups, the doriae, dorsalis, and emarginata groups are newly recognized based on the phylogenetic results and are described. Lectotypes are designated for 18 species. A checklist and key for all crowned weevil genera, key to species groups and species of Asytesta, adult habitus illustrations, distribution maps, and line drawings of diagnostic characters are provided. v TABLE OF CONTENTS ACKNOWLEDGEMENTS ............................................................................................ i DEDICATION ................................................................................................................ iii ABSTRACT .................................................................................................................... iv TABLE OF CONTENTS ................................................................................................ vi LIST OF TABLES .......................................................................................................... x LIST OF FIGURES ........................................................................................................ xi DISCLAIMER ............................................................................................................. xviii PHYLOGENETIC ANALYSIS AND REVISION OF ASYTESTA PASCOE, 1865 (COLEOPTERA: CURCULIONIDAE: CRYPTORHYNCHINAE).......................... 1 INTRODUCTION ................................................................................................... 1 Ecology of Asytesta ..............................................................................................4 Taxonomic History .............................................................................................. 5 Subfamily, tribes, and subtribes ....................................................................... 6 The crowned weevil group ............................................................................... 6 Asytesta Pascoe ................................................................................................ 8 MATERIALS AND METHODS ............................................................................. 10 Collections Consulted .......................................................................................... 10 Material Examined ............................................................................................... 12 Morphological Terminology ................................................................................ 13 Specimen Preparation .......................................................................................... 13 Species Descriptions and Illustrations ................................................................. 14 Character Systems and Coding ............................................................................ 15 Phylogenetic Analyses ......................................................................................... 16 vi RESULTS ................................................................................................................ 18 Characters and Character States ........................................................................... 18 Additional Comments on Character Performance ............................................... 30 Phylogenetic Analyses ......................................................................................... 31 Taxonomic Actions .............................................................................................