Systematics, Biology, and Behavior of Fruit-Piercing and Blood- Feeding Moths in the Subfamily Calpinae (Lepidoptera: Noctuidae)
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SYSTEMATICS, BIOLOGY, AND BEHAVIOR OF FRUIT-PIERCING AND BLOOD- FEEDING MOTHS IN THE SUBFAMILY CALPINAE (LEPIDOPTERA: NOCTUIDAE) By JENNIFER MICHELLE ZASPEL 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 2008 1 © 2008 Jennifer M. Zaspel 2 To Dr. Hans Bänziger for assistance with this project and for his discovery of blood-feeding moths in the genus Calyptra. 3 ACKNOWLEDGMENTS First and foremost, I thank my advisor and chair of my graduate committee, Dr. Marc A. Branham and the members of my graduate committee, Dr. Marjorie A. Hoy, Dr. Jacqueline Miller, and Dr. David Reed for their professional advice, scientific guidance, and financial support. I also thank Dr. Hans Bänziger and Michael Fibiger for many helpful discussions about Calyptra. I would like to thank Drs. A. Jeyaprakash and J. Meyer for their technical advice and laboratory training in molecular biology. Vladimiar S. Kononenko was instrumental in organizing the expeditions to far eastern Russia and for the acquisition of the specimens used in several studies in my dissertation. I would also like to thank my field guide on both expeditions in Russia, Boris Popkov, the staff of the Hunting Area, and the research scientists at Gornotayeznaya Biological Station. I also greatly appreciate the assistance of Ms. Valentina Kolesnikova from the Russian Academy of Sciences Far Eastern Branch for her assistance in obtaining permits for collecting. I also thank Susan Weller and Harald Krenn for suggestions on the comparative mouthpart survey of calpine noctuids (Chapter 2); Hans Bänziger, Roland Hilgartner, and Harry Fay kindly provided adult feeding images figured in the chapter. Drs. R. Rougerie, M. Hajibabaei, D. Janzen, W. Hallwachs, and P. Hebert assistanced in obtaining barcode sequences for some taxa used in the molecular phylogenetic study. I also acknowledge all individuals and institutions listed in Chapter 3 for their assistance in obtaining specimens for the morphological and molecular phylogenetic study. I gratefully acknowledge the Florida Department of Agriculture, Division of Plant Industries, Gainesville, FL for the use of their SEM and Dr. Paul Skelly for his assistance with the imaging equipment as well as Branden Apitz for assistance with SEM and figure formatting. The following people are acknowledged for their assistance in data collection for chapter 7: M.E. Sharf and his graduate students for the use of the QT-RT PCR machine in his 4 laboratory, Paul Shirk for supplying specimens, and D.G. Boucias for the use of his spectrophotometer. Finally, I would like to thank past and present members of Branham, Hoy, McGuire Center, Reed, and Weller laboratories over the years for their support and numerous scientific discussions. This work would not have been possible without the support of my family and their encouragement throughout my graduate studies. This work was supported in part by the American Philosophical Society, Davies, Fischer, and Eckes Endowment in Biological Control to Marjorie A. Hoy at the University of Florida, Explorer’s Club, Florida Entomological Society, National Geographic Society’s Fund for Research and Exploration, National Park Service Inventories and Monitoring Program, National Science Foundation (DDIG, DEB-0807975), Systematics Research Fund, and the Vam York Scholarship Fund. 5 TABLE OF CONTENTS page ACKNOWLEDGMENTS ...............................................................................................................4 LIST OF TABLES...........................................................................................................................9 LIST OF FIGURES .......................................................................................................................11 ABSTRACT...................................................................................................................................13 CHAPTER 1 INTRODUCTION ..................................................................................................................15 Literature Review ...................................................................................................................15 Overview of Fruit-Piercing and Blood-Feeding Moths in Tribe Calpini........................15 Systematics of Calpini.....................................................................................................18 Proboscis morphlogy in piercing moths..........................................................................20 Blood feeding and endosymbionts in Calpini .................................................................21 Research Objectives................................................................................................................24 2 A COMPARATIVE SURVEY OF PROBOSCIS MORPHOLOGY AND ASSOCIATED STRUCTURES IN FRUIT-PIERCING AND BLOOD-FEEDING MOTHS IN THE SUBFAMILY CALPINAE (LEPIDOPTERA: NOCTUIDAE) ...............31 Introduction.............................................................................................................................31 Materials and Methods ...........................................................................................................34 Terminology ....................................................................................................................34 Characterization of Functional Feeding Catergories.......................................................35 Specimen Preparation......................................................................................................39 Results.....................................................................................................................................41 Description of the piercing structures of the proboscis visible by light microscopy ......41 Description of the structures visible by scanning electron microscopy ..........................43 Discussion...............................................................................................................................51 3 RECONSTRUCTING THE EVOLUTIONARY RELATIONSHPS OF THE VAMPIRE MOTHS AND THEIR FRUIT-PIERCING RELATIVES USING MORPHOLOGICAL AND MOLECULAR DATA (LEPIDOPTERA: NOCTUIDAE: CALPINAE: CALPINI) .........................................................................................................74 Introduction.............................................................................................................................74 Materials and Methods ...........................................................................................................76 Taxon Sampling...............................................................................................................76 Morphological Data.........................................................................................................78 Molecular Data ................................................................................................................80 6 Phylogenetic Analyses.....................................................................................................81 Evolution of Feeding Behaviors and Complementary Analyses.....................................82 Results and Discussion ...........................................................................................................83 Summary of Morphological Character Variation............................................................83 Phylogenetic Analysis of Morphological Data................................................................88 Molecular Data and Combined Analyses........................................................................89 Evolution of Feeding Behaviors and Complementary Analysis .....................................91 Conclusions.............................................................................................................................92 4 WORLD CHECKLIST OF TRIBE CALPINI (LEPIDOPTERA: NOCTUIDAE: CALPINAE) .........................................................................................................................125 Introduction...........................................................................................................................125 Checklist ...............................................................................................................................127 Calpini ...........................................................................................................................127 Genus and tribal placement undetermined ....................................................................140 5 ANOTHER BLOOD FEEDER? EXPERIMENTAL FEEDING OF A FRUIT- PIERCING MOTH SPECIES ON HUMAN BLOOD IN THE PRIMORYE TERRITORY OF FAR EASTERN RUSSIA (LEPIDOPTERA: NOCTUIDAE: CALPINAE) .........................................................................................................................141 Introduction...........................................................................................................................141 Materials and Methods .........................................................................................................142 Description of observation sites ....................................................................................142 Experimental methods...................................................................................................144 Results...................................................................................................................................145