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Coleoptera: Curculionidae) Systematics, Morphology, and Evolution of the New World Conoderinae Schoenherr, 1833 (Coleoptera: Curculionidae) by Salvatore Anzaldo A Dissertation Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy Approved June 2019 by the Graduate Supervisory Committee: Nico Franz, Chair Emilia Martins Kathleen Pigg Christian Rabeling ARIZONA STATE UNIVERSITY December 2019 ABSTRACT Weevils are one of the most diverse groups of animals with thousands of species suspected to remain undiscovered. The Conoderinae Schoenherr, 1833 are no exception, being especially diverse and unknown in the Neotropics where they are recognizable for their unique behaviors and color patterns among weevils. Despite these peculiarities, the group has received little attention from researchers in the past century, with almost nothing known about their evolution. This dissertation presents a series of three studies that begin to elucidate the evolutionary history of these bizarre and fascinating weevils, commencing with an overview of their biology and classificatory history (Chapter 1). Chapter 2 presents the first formal cladistic analysis on the group to redefine the New World tribes Lechriopini Lacordaire, 1865 and Zygopini, Lacordaire, 1865. An analysis of 75 taxa (65 ingroup) with 75 morphological characters yielded six equally parsimonious trees and synapomorphies that are used to reconstitute the tribes, resulting in the transfer of sixteen genera from the Zygopini to the Lechriopini and four generic transfers out of the Lechriopini to elsewhere in the Conoderinae. Chapter 3 constitutes a taxonomic revision of the genus Trichodocerus Chevrolat, 1879, the sole genus in the tribe Trichodocerini Champion, 1906, which has had an uncertain phylogenetic placement in the Curculionidae but has most recently been treated in the Conoderinae. In addition to redescriptions of the three previously described species placed in the genus, twenty-four species are newly described and an identification key is provided for all recognized species groups and species. Chapter 4 quantitatively tests the similarity in color pattern among species hypothesized to belong to several different mimicry complexes. The patterns of 160 species of conoderine weevils were evaluated for 15 categorical and continuous characters. Non- i metric multidimensional scaling (NMDS) is used to visualize similarity by the proximity of individual species and clusters of species assigned to a mimicry complex in ordination space with clusters being statistically tested using permutational multivariate analysis of variance (PERMANOVA). ii ACKNOWLEDGMENTS There are many people who supported and assisted me over the past six years that I would like to thank, for I could not have completed this dissertation without them. From an early age, my parents, Maria and Stephen, and my sisters, Jackie and Christina, have encouraged my interest in insects. It is hard to imagine still being able to pursue a profession I love without growing up in such a supportive and caring environment. Pushing through the final months of these projects would not have been possible without the patience and support of my wife Hannah, who has made many sacrifices for us to be together even when I was at my busiest. My advisor, Nico Franz, helped me discover my interest in weevils and gave me the time to develop my projects. His mentorship has made me a profoundly better scientist and I will be forever grateful for the opportunity to be a student in his lab. I would also like to thank the members of the Franz lab for sharing many good times collecting beetles throughout the Southwest. I am especially indebted to Charlie and Lois O’Brien, whose hospitality always made visiting their collection and home a treat. Charlie’s generosity with specimens and the sharing of his weevil knowledge throughout all stages of these projects has made each one immeasurably better and contributed significantly to my increasing interest in weevils early in my graduate studies. Without the labors of previous weevil workers to uncover the mysteries of these fascinating insects there would have been no foundation to develop the projects of this dissertation on. Special thanks go to the Arizona State University library staff for their timely fulfillment of the hundreds of interlibrary loan requests I have made in the past six years, thus making this prior research available to me. iii TABLE OF CONTENTS Page LIST OF TABLES .................................................................................................................................... vi LIST OF FIGURES ................................................................................................................................ vii CHAPTER 1 INTRODUCTION TO THE CONODERINAE SCHOENHERR, 1833 ............. 1 2 SYSTEMATICS OF THE LECHRIOPINI LACORDAIRE, 1865 (COLEOPTERA: CURCULIONIDAE: CONODERINAE) .................................... 7 Introduction ...................................................................................................................... 8 Taxonomic History of the New World Conoderinae ............................................... 9 Materials and Methods .................................................................................................... 9 Results .............................................................................................................................. 13 Discussion ....................................................................................................................... 38 3 REVISION OF THE WEEVIL GENUS TRICHODOCERUS CHEVROLAT, 1879 (CURCULIONIDAE: CONODERINAE: TRICHODOCERINI) .............. 46 Introduction .................................................................................................................... 47 Materials and Methods .................................................................................................. 48 Phenotypic Uniqueness of Trichodocerus ...................................................................... 49 Taxonomic Treatment ................................................................................................... 54 T. spinolae Group ...................................................................................................... 63 T. brevilineatus Group ............................................................................................... 92 T. ochromae Group .................................................................................................. 107 T. antiquus Group .................................................................................................. 131 T. mecistotarbus Group ............................................................................................ 150 iv CHAPTER Page T. obrieni Group ...................................................................................................... 177 Key to Trichodocerus Species Groups and Species .................................................... 203 Discussion ..................................................................................................................... 210 4 A QUANTITATIVE CHARACTERIZATION OF CONODERINE MIMICRY COMPLEXES (COLEOPTERA: CURCULIONIDAE) ......................................... 218 Introduction .................................................................................................................. 219 Mimicry Complexes in the New World Conoderinae ........................................... 219 Materials and Methods ................................................................................................ 222 Results ............................................................................................................................ 230 Discussion ..................................................................................................................... 233 REFERENCES ..................................................................................................................................... 237 APPENDIX A CHARACTER MATRIX USED IN CLADISTIC ANALYSIS .......................... 245 B CHARACTER MATRIX USED IN MIMETIC ANALYSIS .............................. 248 C PAIRWISE PERMANOVA RESULTS FOR THE FULL DATASET ............. 253 v LIST OF TABLES Table Page 2.1. Summary Classification of the Lechriopini and Zygopini ........................................... 40 3.1. Checklist of Trichodocerus Species and Documented Distributions .......................... 209 4.1. Pairwise PERMANOVA Results for the Reduced Analysis .................................... 230 vi LIST OF FIGURES Figure Page 2.1. Modification to the Mesoventrite in the New World Conoderinae ..................... 10 2.2. Confocal Laser-Scanning Microscopy Images of the Maxilla, Part I ................... 16 2.3. Confocal Laser-Scanning Microscopy Images of the Maxilla, Part II .................. 17 2.4. Confocal Laser-Scanning Microscopy Images of the Labium ............................... 20 2.5. Scanning Electron Micrographs of the Prosternum ................................................ 22 2.6. Various Morphological Structures in the New World Conoderinae .................... 35 2.7. Preferred Cladogram of Six Equally Parsimonious Trees ...................................... 36 2.8. Strict Consensus of Six Equally Parsimonious Trees .............................................. 37 3.1. Line Illustrations of the Antennal Club and Funicular Article 7, Part I .............
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