INVESTIGATING THE INTEGRATION OF SMALL HIVE BEETLES (AETHINA TUMIDA MURRAY, COLEOPTERA: NITIDULIDAE) INTO WESTERN HONEY BEE (APIS MELLIFERA L., HYMENOPTERA: APIDAE) COLONIES By EDWARD BLAKE ATKINSON A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2011 1 © 2011 Edward Blake Atkinson 2 To my wife and kids 3 ACKNOWLEDGEMENTS I thank my advisor Dr. Jamie Ellis and other members of my committee, Drs. H. Jane Brockmann, Andy Cline, Heather McAuslane, and Peter Teal, for guidance and assistance with what was a wonderful introduction to my life of entomological research. Also, I thank the University of Florida for funding me through my doctoral career. I acknowledge the following members of the University of Florida Honey Bee Research and Extension Laboratory who provided technical assistance with my projects: Katie Buckley, Meredith Cenzer, Renee Cole, Jonnie Dietz, Mark Dykes, Katy Evans, Dr. Kamran Fakhimzadeh, Jason Graham, Dr. Alec Gregorc, Pablo Herrera, Michelle Kelley, Ben King, Jeanette Klopchin, Hannah O’Malley, Mike O’Malley, Catherine Zettel Nalen, Dr. Akers Pence, Andy Sheffler, Marissa Streifel, Cindy Tannahill, Melissa Teems, Liana Teigen, Tricia Toth, Anthony Vaudo, Sparky Vilsaint, and Larry Wise. I am grateful to Jane Medley of the University of Florida Entomology and Nematology department for creating the wonderful diagrams used in this dissertation, Drs. Mark Carroll, Nicole Benda, Adrian Duehl, and Richard Arbogast of the USDA-ARS Center for Medical and Veterinary Entomology (CMAVE) for offering their scientific expertise during the experimental process, and Tredina Davis, Dr. Matt Lehnert, and Curtis Murphy of CMAVE for providing beetles when I was in need. Also, I thank Alex Bolques of the Gadsden County Extension Office, Dr. Keith Delaplane and Amanda Tedrow of the University of Georgia, Jim McBride of Jim’s Farm, Jim Boyer and Buck Nelson of the University of Florida Plant Research and Education Unit, and Kyle Straughn of Straughn Farms for aiding me in my search for beetle species. Jerry Hayes of the Florida Department of Agriculture and Consumer Services, Division of Plant Industry (DPI) also was very accommodating when I needed more bee colonies, and Drs. Paul Skelley and Mike Thomas of DPI never hesitated to offer me help in identifying beetles and using the scanning electron microscope. Finally, I thank my parents, sister, nephew, 4 and my in-laws for being excited for me, my wife for being so very kind to and supportive of me, our babies for making me the happiest daddy around, and my grandfather-in-law, Dr. Robert Woodruff, for introducing me to the field of Entomology. 5 TABLE OF CONTENTS page ACKNOWLEDGEMENTS......................................................................................................... 4 LIST OF TABLES ...................................................................................................................... 9 LIST OF FIGURES .................................................................................................................. 10 ABSTRACT ............................................................................................................................. 11 CHAPTER 1 INTRODUCTION ............................................................................................................. 13 Methods Used by Symbionts to Integrate into Non-Apis mellifera Social Bee Nests ........... 14 Morphological ............................................................................................................. 14 Passive defense .................................................................................................... 15 Phoresy ................................................................................................................ 16 Other .................................................................................................................... 16 Behavioral ................................................................................................................... 17 Phoresy ................................................................................................................ 17 Feeding/reproduction ............................................................................................ 18 Chemical ..................................................................................................................... 20 Methods Used by Symbionts to Integrate into Apis mellifera Nests .................................... 20 Morphological ............................................................................................................. 21 Passive defense .................................................................................................... 21 Phoresy ................................................................................................................ 21 Other .................................................................................................................... 22 Behavioral ................................................................................................................... 22 Phoresy ................................................................................................................ 22 Feeding/reproduction ............................................................................................ 22 Other .................................................................................................................... 24 Chemical ..................................................................................................................... 25 Methods Used by SHBs to Integrate into Apis mellifera Nests ............................................ 25 Morphological ............................................................................................................. 26 Behavioral ................................................................................................................... 26 Chemical ..................................................................................................................... 27 Introduction to the Research Presented in this Dissertation ................................................. 28 2 ATTRACTION OF MULTIPLE BEETLE SPECIES TO HONEY BEE HIVE ODORS .... 47 Materials and Methods ....................................................................................................... 49 Beetles ........................................................................................................................ 49 Honey Bees ................................................................................................................. 50 Olfactometer ............................................................................................................... 50 6 Choice Bioassay .......................................................................................................... 51 Statistical Analysis ...................................................................................................... 51 Results ............................................................................................................................... 52 Discussion .......................................................................................................................... 53 3 SHB ADAPTIVE LEG MORPHOLOGY .......................................................................... 58 Materials and Methods ....................................................................................................... 61 Results ............................................................................................................................... 61 Discussion .......................................................................................................................... 62 4 ADAPTIVE BEHAVIOR OF HONEY BEES TOWARD BEETLE INVADERS EXHIBITING VARIOUS LEVELS OF COLONY INTEGRATION ................................. 68 Materials and Methods ....................................................................................................... 70 Beetles ........................................................................................................................ 70 Observation Hives ....................................................................................................... 70 Behavioral Assays ....................................................................................................... 71 Statistical Analysis ...................................................................................................... 72 Results ............................................................................................................................... 72 Discussion .......................................................................................................................... 74 5 DIETARY EFFECTS ON THE CUTICULAR PROFILE OF SHBS AND ITS EFFECT ON BEHAVIORAL TREATMENT BY HONEY BEES .................................................... 82 Materials and Methods ....................................................................................................... 84 Beetles ........................................................................................................................ 84 Observation Hives ......................................................................................................