SYSTEMATICS OF LADYDIRD BEETLES (COLEOPTERA: COCCINELLIDAE) by JOSÉ ADRIANO GIORGI (Under the Direction of Joseph V. McHugh) ABSTRACT Despite the familiarity and economic significance of Coccinellidae, the family has thus far escaped analysis by rigorous phylogenetic methods. As a result, the internal classification remains unstable and there is no framework with which to interpret evolutionary events within the family. We analyzed coccinellid phylogeny using a combined dataset of seven genes: 12S rDNA, 16S rDNA, 18S rDNA, 28S rDNA, Cytochrome oxidase I, Cytochrome oxidase II, and Histone 3. The entire dataset consists of 6565 aligned nucleotide sites, 1305 of which are parsimony informative. Our study included 20% of the generic-level diversity and 80% of the tribal-level diversity and 100% previously recognized subfamilies. We analyzed the dataset using parsimony and Bayesian methods. Our study supports the monophyly of Coccinellidae; however, most of the traditional subfamilies are not supported as monophyletic. Three recently proposed, but not widely accepted, subfamilies are recognized. A new subfamily is proposed to accommodate Monocoryni. We recognize eight subfamilies of Coccinellidae: Microweiseinae, Monocorynae (new subfamily), Coccinellinae, Chilocorinae, Sticholotidinae, Scymninae, Exoplectrinae, and Hyperaspidinae. The circumscription of Hyperaspidinae (Hyperaspidini Mulsant, Brachiacanthini Mulsant and Selvadiini Gordon) is extended to accommodate Platynaspidini Redtenbacher and Aspidimerini Mulsant. The tribe Coccinellini Latreille is paraphyletic with respect to Tytthaspidini Mulsant (syn. nov.) and Halyziini Mulsant (confirmed status). The tribes Noviini Mulsant, Cryptognathini Casey, Poriini Mulsant, and Diomini Gordon are treated as incertae sedis. The relationship between some of the subfamilies and the placement of several tribes remain ambiguous. We also utilized the phylogenetic hypothesis to provide an evolutionary perspective on the feeding preferences of coccinellids. Coccinellids exhibit a wide range of preferred food types, spanning kingdoms and trophic levels. Our study suggests that the ancestral feeding condition for the family is coccidophagy and that polyphagy served as an evolutionary stepping stone for primarily predaceous groups to adopt new feeding habits. The Australian members of the ladybird beetle tribe Chilocorini are revised. Identification keys for genera and species are provided. Habitus and diagnostic characters from mouthparts, legs, and genitalia are illustrated. Major conclusions of the present study include recognition of the following: (1) five new species; (2) eleven junior synonyms at the species level; one junior synonym at the generic level; and (3) sixteen new lectotypes. According to our study, the Chilocorini in Australia consists of 23 species classified in 6 genera: Brumoides, Chilocorus, Exochomus, Halmus, Orcus and Trichorcus. INDEX WORDS: Coleoptera, Cucujoidea, Cerylonid Series, Coccinellidae, ladybird beetle, lady beetle, ladybug, systematics, phylogeny, evolution, food preference, prey, trophic shift, taxonomic revision, Chilocorinae, Chilocorini, Australia. SYSTEMATICS OF LADYDIRD BEETLES (COLEOPTERA: COCCINELLIDAE) by JOSÉ ADRIANO GIORGI B.S., Universidade Federal Rural do Rio de Janeiro, Brazil, 1996 M. S., Universidade de Sao Paulo, Brazil, 2001 A Dissertation Submitted to the Graduate Faculty of The University of Georgia in Partial Fulfillment of the Requirements for the Degree DOCTOR OF PHYLOSOPHY ATHENS, GEORGIA 2009 © 2009 JOSÉ ADRIANO GIORGI All Rights Reserved SYSTEMATICS OF LADYDIRD BEETLES (COLEOPTERA: COCCINELLIDAE) by JOSÉ ADRIANO GIORGI 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 December 2009 DEDICATION I dedicate this dissertation to Dona Norma, my mom. iv ACKNOWLEDGEMENTS I thank my advisor, Dr. Joseph V. McHugh for his encouragement, support and instruction. I thank my committee members, Dr. Ken Ross and Dr. Mark Farmer for their critical review and assistance. Thanks to my lab mates Chris Hartley, Floyd Shockley, James Robertson, Juanota Forrester, and Nathan Lord for their support and friendship over the past six years. I thank the graduate coordinators Dr. Darold Batzer and Dr. Mark Brown for taking care of the paperwork so I could enroll at UGA and for the fair and reasonable TA assignments; especially for not assigning me to teach Intro-bio! I thank the Department of Entomology at University of Georgia and the H.H. Ross Fund for the generous financial support for these studies and other activities associated to it. I am very grateful to the competence, kindness, and patience of many members of the Department of Entomology staff, especially to Danny Fendley, Jennie De la Vega, Marianne Shockley, Nancy Jordan, Sam Waychoff, and Terri All. I am particularly thankful to my collaborators at CSIRO (AU), Brigham Young University, University of New Mexico and USDA (Dr. Adam !lipi"ski, Dr. Michael Whiting, Dr. Kelly Miller, and Dr. Natalia Vandenberg, respectively). This study would have not been possible without the specimens kindly provided by the people acknowledged in the following chapters. v TABLE OF CONTENTS Page ACKNOWLEDGEMENTS.............................................................................................................v CHAPTER 1 INTRODUCTION .........................................................................................................1 2 LITERATURE REVIEW ..............................................................................................3 References .................................................................................................................4 3 THE EVOLUTION OF FOOD PREFERENCE IN COCCINELLIDAE .....................8 Abstract .....................................................................................................................9 Introduction .............................................................................................................10 Material and Methods..............................................................................................21 Results .....................................................................................................................24 Discussion ...............................................................................................................25 Acknowledgements .................................................................................................37 References ...............................................................................................................37 Figure captions ........................................................................................................60 4 PHYLOGENY OF LADYBIRD BEETLES (COLEOPTERA: COCCINELLIDAE) BASED ON MOLECULAR DATA ……………………………………………..69 Abstract ...................................................................................................................70 Introduction .............................................................................................................71 Material and Methods..............................................................................................74 vi Results .....................................................................................................................76 Discussion ...............................................................................................................77 Acknowledgements ...............................................................................................106 References .............................................................................................................106 Figure captions ......................................................................................................125 5 REVISION OF THE AUSTRALIAN COCCINELLIDAE (COLEOPTERA). PART 6. TRIBE CHILOCORINI ....................................................................................129 Abstract .................................................................................................................130 Introduction ...........................................................................................................131 Material and Methods............................................................................................132 Taxonomy..............................................................................................................134 Acknowledgments.................................................................................................199 References .............................................................................................................200 Figure captions ......................................................................................................206 6 SUMMARY AND CONCLUSIONS ........................................................................220 vii CHAPTER 1 INTRODUCTION This dissertation, written in article style, presents three separate research projects that aim to improve our knowledge about the evolution, phylogenetics and taxonomy of the ladybeetle family Coccinellidae. In the first study (chapter 3), the evolution of feeding patterns for the family Coccinellidae is interpreted under the phylogenetic framework using molecular data from two ribosomal genes. Coccinellidae is remarkable for many of its feeding habits. While all