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University of Florida Thesis Or Dissertation Formatting ECOLOGY AND CONSERVATION OF EUMAEUS ATALA POEY 1832 (LEPIDOPTERA; LYCAENIDAE) By SANDRA E. KOI A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2013 © 2013 Sandra E. Koi I dedicate this work to HaShem and all those Light beings who have played a significant role in my life: family and friends, human and other. ACKNOWLEDGMENTS I extend my heartfelt appreciation to my major professor, Dr. Jaret Daniels, who provided me with encouragement, materials, equipment and his expert guidance, giving me everything necessary to accomplish a complete and detailed life history and biological monograph of the Atala butterfly. I thank my committee members, Dr. Jacqueline Miller for sharing her in-depth and accomplished knowledge of tropical lepidoptera and excellent editing of my sometimes hurried and rough writing, and Dr. Christine Miller for her fresh ideas about monitoring behavior and keen advice in designing my experiments, terminology and scientific ethics. I also thank the leaders in Lepidoptera biology with whom I have been privileged to work, in conferences and/or in the field: Dr. Thomas Emmel, Dr. Donald Hall, Dr. Robert Michael Pyle, Dr. Marc Minno, and Dr. Dean and Sally Jue. I thank Dr. George Casella (z”l) for his superb statistical contribution to my experiment design, and Jonathan Colburn for his thoughtful feedback and help with statistical analyses. I thank Dr. Daniel Hahn for his generosity with laboratory materials, helpful practical advice, use of equipment and the expertise of his postdocs, Gian-Carlo Lopez and Caroline Williams. Dr. James Maruniak is acknowledged for his kind assistance with pathogen identification and Dr. Lyle Buss for his beetle and ant identifications as well as guidance using automontage. Dr. James Nation has been a great source friendship, support and information. Drs. Drion Boucias and Verena Lietze are thanked for help identifying possible pathogens. I thank Dr. Oscar Liburd for the use of his larger balance to weigh plant material. I am very appreciative of Dr. Deborah Matthews-Lott for guidance regarding preservation protocols for genitalic dissections and Dr. Charlie Covell is deeply thanked 4 for giving me access to his historical documents and for sharing his many memories of the early days of lepidoptera studies. I thank Jim Schlachta for his assistance in acoustic monitoring at McGuire Center for Lepidoptera and Biodiversity. I am deeply indebted to Susan Wright and the Biocontrol labs at our USDA/ARS facility for permission to use the environmental chambers to document life stage development of the Atala under controlled conditions. The help and guidance of USDA scientists Dr. Paul Skelley and Lou Soma for teaching me how to use the SEM camera. Drs. Richard Mankin, Seth McNeill, and Mirian Hay-Roe were very helpful setting up experiments to monitor Atala pupae and larvae for putative stridulation. Dr. James Hayden has been very helpful and often seemed to be the only other living soul on campus when questions arose on weekends and holidays. I am grateful to Ga-Eun Lee for months of dedicated help in all aspects of Atala rearing, as well as Matt Thom, Lukasz Barczak, and Marissa Streifel for help rearing the colony, and volunteers Jane Fowler, Bethene Wilkinson, Ryan Huether, Catherine White, Debby Gluckman, Michelle Gray, Ashleigh Price, Erin Kalinowski and Carlos Iglesias for temporary volunteer help. I thank Christine Eliazar, Debbie Hall, Nancy Sanders, Ruth Brumbaugh, and all of the staff in McGuire Center and the Entomology/Nematology Department for their ongoing kind assistance in the myriad of details associated with graduate school! I thank Bud and Jackie Klein at Duck Lake Coontie Farm in Dade City for a substantial host plant donation and Botanics Wholesale in Homestead, Florida for donating a non-native cycad species for herbivory choice tests. 5 I thank with love and appreciation the close friends who literally fortified me materially, spiritually, emotionally and intellectually during the years this project has been in process: Dr. Joshua Feingold and Laurie Flebotte, Dr. Sarah Meltzoff, Dr. Robin Sherman, Ericka Helmick and Dr. Thomas Chouvenc, and Thomas and Lisset Genung. I thank many associates for their friendship: Dr. Elane Nuehring, Janice Malkoff, Barbara DeWitt and members of the North American Butterfly Association in Palm Beach, Broward and Miami-Dade County, as well as biologists and staff in the city, county, state and federal parks in the tri-county area, who have helped collect data over many years. I thank the rabbis and members of Temple Adath Or in Fort Lauderdale, Temple Beth El in Bradenton, Congregation P’nai Or in Tampa, and Temple Shir Shalom in Gainesville for their support, friendship and spiritual nourishment as life’s journeys led me from place to place. I thank my friends and colleagues Marilyn Griffiths, Dr. Tighe Shomer and Susan Shapiro from Fairchild Tropical Botanic Garden for their massive contributions to as yet unpublished research, as well as Martin Feather, Mary Collins, and Georgia Tasker. I also thank the biologists and staff at Montgomery Botanical Center for sharing their on- going research about cycads native to Central and South America and their specialized herbivores as well as our Caribbean and Floridian species. 6 TABLE OF CONTENTS page ACKNOWLEDGMENTS .................................................................................................. 4 LIST OF TABLES .......................................................................................................... 10 LIST OF FIGURES ........................................................................................................ 11 LIST OF OBJECTS ....................................................................................................... 15 LIST OF ABBREVIATIONS ........................................................................................... 16 ABSTRACT ................................................................................................................... 17 CHAPTER 1 HISTORY AND LITERATURE REVIEW OF EUMAEUS ATALA POEY 1832 (LEPIDOPTERA: LYCAENIDAE: THECLINAE) ...................................................... 19 Taxonomy ............................................................................................................... 19 Description .............................................................................................................. 21 Historical and Current Range and Distribution ........................................................ 23 Conservation Status ............................................................................................... 42 Larval Host Plant: Zamia integrifolia (Zamiaceae: Cycadales) ................................ 44 Conclusions ............................................................................................................ 49 2 NEW AND REVISED LIFE HISTORY OF THE HAIRSTREAK EUMAEUS ATALA (LEPIDOPTERA: LYCAENIDAE) WITH NOTES ON CURRENT CONSERVATION STATUS .................................................................................... 65 Introduction ............................................................................................................. 65 Materials and Methods............................................................................................ 68 Biological Rearing: Livestock and Progeny ...................................................... 68 Immature Measurements .................................................................................. 72 Adult Biology and Reproductive Behavior ........................................................ 73 Results .................................................................................................................... 74 Oogenesis, Oocyte Development, Ovipositing and Ova Production ................. 74 Larval Development ......................................................................................... 75 Pupal Development .......................................................................................... 80 Total Development Time .................................................................................. 81 Adult Measurements and Longevity ........................................................................ 81 Adult Dissections and Ova Development ................................................................ 84 Lifespan and Sex Ratio ........................................................................................... 85 Diseases, Pathogens and Abnormalities ................................................................ 86 Discussion .............................................................................................................. 86 Conservation Notes ................................................................................................ 89 7 3 ETHOLOGY (BEHAVIORAL STUDY) OF EUMAEUS ATALA (LEPIDOPTERA: LYCAENIDAE) ...................................................................................................... 115 Insect Behavior ..................................................................................................... 116 Materials and Methods.......................................................................................... 117 Atala Adults .................................................................................................... 118 Learned behaviors
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