SYSTEMATICS OF THE LICHEN MOTH TRIBE LITHOSIINI (LEPIDOPTERA: EREBIDAE: ARCTIINAE) INCLUDING A REVIEW OF THE GENUS LYCOMORPHA HARRIS By CLARE HILARY SCOTT 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 2012 1 © 2012 Clare H. Scott 2 To my family whose support throughout this long journey has been invaluable 3 ACKNOWLEDGMENTS This research would not have been possible without the support of numerous people and institutions. I would like to acknowledge the support and advice of my major advisor and committee chair, Dr. Marc Branham, and the other members of my graduate committee, Drs. Rebecca Simmons, Jackie Miller, David Reed, and Christine Miller. I would like to thank Drs. Susan Weller and Jennifer Zaspel for giving me the opportunity to participate in their molecular phylogeny of Arctiinae. Furthermore, I wish to acknowledge the National Science Foundation Award DEB#0919185 for the partial financial support of this research. Without the funding Drs. Weller and Zaspel received, the molecular phylogeny of Lithosiini would not have been completed. In addition, the work of Pablo Chialvo, Taylor Wardwell, and Elizabeth Phillippi on this study was invaluable. Pablo helped to amplify the COI and 28S gene fragments and willingly came into the lab on the weekend and holidays to help with this project. Taylor and Elizabeth helped to amplify the CytB and RpS5 gene fragments. In addition, Taylor dealt with submitting all fragments for sequencing. I would like to acknowledge all of the collections listed in Chapters 2-5 for the material that they provided. Without the use of these specimens, the morphological studies would not have been possible, and the taxon sampling of the molecular study would not have been as broad. Further thanks are offered to the American Museum of Natural History, the Carnegie Museum of Natural History, the Cornell University Insect Collection, the Florida Museum of Natural History, the Museum of Comparative Zoology (Harvard), the United States Museum of Natural History, and the Yale Peabody Museum. Each of these institutes was kind enough to allow me to visit and examine 4 their holdings of several families of Lepidoptera, while looking for specimens of Lycomorpha. The tribal phylogenetic analyses were conducted using the University of Florida Phyloinformatics Cluster for High Performance Computing in the Life Sciences and the CIPRES Science Gateway (Miller et al. 2010). Without access to these computer clusters, the time needed to run the tribal phylogenetic analysis would have been prohibitively long. This research was funded in part by the Museum of Comparative Zoology Ernst Mayr Travel Grant in Animal Systematics, the Systematics, Evolution, and Biodiversity Section’s Student Travel Award of the Entomological Society of America, the University of Florida Alumni Fellowship, and the University of Florida SPICE fellowship. 5 TABLE OF CONTENTS page ACKNOWLEDGMENTS .................................................................................................. 4 LIST OF TABLES .......................................................................................................... 10 LIST OF FIGURES ........................................................................................................ 11 ABSTRACT ................................................................................................................... 14 CHAPTER 1 INTRODUCTION .................................................................................................... 16 Literature Review .................................................................................................... 16 Larval Feeding Behavior................................................................................... 17 Chemical Defense ............................................................................................ 19 Courtship Behavior ........................................................................................... 20 Research Objectives ............................................................................................... 22 Chapter 2 .......................................................................................................... 22 Chapter 3 .......................................................................................................... 23 Chapter 4 .......................................................................................................... 24 Chapter 5 .......................................................................................................... 25 Chapter 6 .......................................................................................................... 26 2 A PRELIMINARY PHYLOGENY OF THE LICHEN MOTH TRIBE LITHOSIINI (LEPIDOPTERA: EREBIDAE: ARCTIINAE) BASED ON MORPHOLOGICAL CHARACTERS ....................................................................................................... 27 Background Information .......................................................................................... 27 Materials and Methods............................................................................................ 31 Taxon Sampling ............................................................................................... 31 Morphology ....................................................................................................... 32 Characters Examined ....................................................................................... 33 Characters and their Phylogenetic Usefulness ................................................. 33 Head .......................................................................................................... 33 Thorax ........................................................................................................ 34 Wings ......................................................................................................... 35 Abdomen.................................................................................................... 36 Male abdomen and genitalia ...................................................................... 37 Female abdomen and genitalia .................................................................. 42 Phylogenetic Analysis ...................................................................................... 44 Results .................................................................................................................... 45 Discussion .............................................................................................................. 49 6 3 MOLECULAR PHYLOGENY OF THE TRIBE LITHOSIINI (LEPIDOPTERA: EREBIDAE: ARCTIINAE) ....................................................................................... 83 Background Information .......................................................................................... 83 Materials and Methods............................................................................................ 87 Taxon Sampling ............................................................................................... 87 DNA Extraction, Amplification, and Sequencing ............................................... 88 Phylogenetic Methods ...................................................................................... 91 Results and Discussion........................................................................................... 92 4 PHYLOGENY OF THE LICHEN MOTH GENUS LYCOMORPHA HARRIS (LEPIDOPTERA: EREBIDAE: ARCTIINAE) ......................................................... 109 Background Information ........................................................................................ 109 Materials and Methods.......................................................................................... 112 Taxon Sampling ............................................................................................. 112 Collections Consulted ..................................................................................... 113 Morphology ..................................................................................................... 114 Characters Examined ..................................................................................... 115 Phylogenetic Analysis .................................................................................... 115 Results .................................................................................................................. 117 Phylogenetic Analysis and Taxonomic Implications ....................................... 117 Species Checklist of Lycomorpha .................................................................. 122 Checklist of Species Described as Lycomorpha ............................................. 123 Lithosiini ................................................................................................... 123 Arctiini ...................................................................................................... 124 Zygaenidae .............................................................................................. 125 Incertae sedis ........................................................................................... 126 Characters and their Phylogenetic Usefulness ............................................... 126 Head .......................................................................................................