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Species Boundaries, Biogeography, and Intra-Archipelago Genetic Variation Within the Emoia Samoensis Species Group in the Vanuatu Archipelago and Oceania" (2008)

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Species Boundaries, Biogeography, and Intra-Archipelago Genetic Variation Within the Emoia Samoensis Species Group in the Vanuatu Archipelago and Oceania Louisiana State University LSU Digital Commons LSU Doctoral Dissertations Graduate School 2008 Species boundaries, biogeography, and intra- archipelago genetic variation within the Emoia samoensis species group in the Vanuatu Archipelago and Oceania Alison Madeline Hamilton Louisiana State University and Agricultural and Mechanical College, [email protected] Follow this and additional works at: https://digitalcommons.lsu.edu/gradschool_dissertations Recommended Citation Hamilton, Alison Madeline, "Species boundaries, biogeography, and intra-archipelago genetic variation within the Emoia samoensis species group in the Vanuatu Archipelago and Oceania" (2008). LSU Doctoral Dissertations. 3940. https://digitalcommons.lsu.edu/gradschool_dissertations/3940 This Dissertation is brought to you for free and open access by the Graduate School at LSU Digital Commons. It has been accepted for inclusion in LSU Doctoral Dissertations by an authorized graduate school editor of LSU Digital Commons. For more information, please [email protected]. SPECIES BOUNDARIES, BIOGEOGRAPHY, AND INTRA-ARCHIPELAGO GENETIC VARIATION WITHIN THE EMOIA SAMOENSIS SPECIES GROUP IN THE VANUATU ARCHIPELAGO AND OCEANIA A Dissertation Submitted to the Graduate Faculty of the Louisiana State University and Agricultural and Mechanical College in partial fulfillment of the requirements for the degree of Doctor of Philosophy in The Department of Biological Sciences by Alison M. Hamilton B.A., Simon’s Rock College of Bard, 1993 M.S., University of Florida, 2000 December 2008 ACKNOWLEDGMENTS I thank my graduate advisor, Dr. Christopher C. Austin, for sharing his enthusiasm for reptile diversity in Oceania with me, and for encouraging me to pursue research in Vanuatu. His knowledge of the logistics of conducting research in the Pacific has been invaluable to me during this process. His desire for me to succeed as a graduate student in his lab has been crucial to my development as a scientist and independent researcher. I also thank the other members of my graduate committee, Dr. Robb T. Brumfield, Dr. Mark S. Hafner, Dr. Michael E. Hellberg, and Dr. Dorothy Prowell for their assistance in this project. Aside from my graduate committee members, I received assistance and advice from Dr. Bryan Carstens, Dr. Joseph Hartman, and Dr. George Zug on this research project, and I thank them for their time and generosity. I thank the members of the Environment Unit of the Republic of Vanuatu, especially the head of the unit, Mr. Ernest Bani, and Donna Kalfatak, Catherine Malosu, and Leah Nimoho for permission to conduct research from 2000-2005 and collecting and export permits. I am also especially thankful to Donna Kalfatak for her willingness to assist me with logistics associated with fieldwork, including convincing her friends and relatives to provide me occasional support and housing. Thanks to Catherine Malosu, Rolenas Tavue, and Donna Kalfatak for assistance with fieldwork in Vanuatu. Most of all I owe thanks to numerous ni-Vanuatu chiefs and villagers for access to land and assistance in Vanuatu in many ways, especially Karl and Dorrie Waldeback and Susan and Robinson Henry for their friendship, kindness, and storage space. Thank yu tumas long ol man Vanuatu, especially Charles and Daisy Lamlam and family (Telvet, MotaLava), Chief Timothy and family (Wiawi, Malakula), Mary, Bong, Oswell, and the rest of their family (Craig Cove, Ambrym), Chief Harry and Susan Wayback, little Jessica, Reneatte, and Harrison (Pangi, Pentecost), Napwhat, Monique, Janet, Jennifer, Terry, and the ii rest of their family (Lobukas, Tanna), Solomon and Purity Tavue and Roy (Matantas, Santo), Stanley Wayne and his family (Isiai, Tanna), Mackin, Leisau and the rest of the faculty and staff of Lamen Bay Secondary School (Epi), the members of the Melanesian Brotherhood and Florance Mata (Saratamata, Ambae), Father Robert Wobur and his family (Gaua), Lillian and Caleb Wilkins, Charles Smith, and Janet Wona (Sola, Vanua Lava), Reynolds and Hilton and their families (Loh Island), Chief Godfrey Manar and his family (Tenan, Vanua Lava), Willie Snow (Wintua, Malakula), Johnclayton Lini and Florence Gleny (Kerepei, Maewo), Pastor Phillip Baniuri (Anelguhuat, Aneityum), and people from the following villages: Anelguhuat, Port Patrick, and Umtech (Aneityum Island), Marae and Mission Bay (Futuna Island), Saratamata and Navunda (Ambae Island), Lamalana, Lawoni, Pangi, Raguru, Pantor, Salap, and Panwoki (Pentecost Island), Ipota (Erromango Island), MeleMat, Eton, Pango Port Vila, Errakor Island, Epau, Tanoliu, Ulei Secondary School, Moala, Nasine, Panganisou, and Erratoka Island (Efate Island), Nasawa and Kerepei (Maewo), Matanatas, Vathe Conservation Area, Butmas, Lajmoli, Tanomom and Tasariki (Espiritu Santo Island), Port Stanley, Lakatoro, Litslits, Wintua, Nowointen, Nabelchel, Wiawi, Leviamp, Wiel, Asourah, Lamap, Port Sandwich and Lavalsal (Malakula Island), Port Resolution, Lowniel, Lobukas, Fetuki Primary School, Green Hill, Lamnatu, Lamlu, Ipak, Isiai, Kwataparen and Yakel (Tanna Island), Nerenigman and Telvet (Mota Lava Island), Sola and Tenan (Vanua Lava Island), Lunharevemenan village on Loh Island and Tegua Island (Torres Islands), Lembot and Lake Letas Conservation Area (Gaua Island), Lamen Bay (Epi Island), Penapo, Lonvert, Polianpul, Melpul, Ranu, Malvert, Craig Cove, Ranon and Ranvetlam (Ambrym Island), Shepherd (Aniwa Island). I am indebted to you all. Without your help, kindness, and knowledge this research would not have been possible. I thank Elaine Klein, Emily Hartfield, Mallory Ecktstut, Kara Blaha, and Kathy Grazyck for research assistance in Vanuatu, and for continued interest in and research on Vanuatu reptiles iii once back in the US. Thank you to Aaron Bauer, Robert Fisher, Steve Donnellan, and George Zug for tissues that were essential for the successful completion of this project. Thanks also to the following museums for loans of specimens: American Museum of Natural History, Texas Natural History Collections, California Academy of Sciences, British Museum of Natural History, Museum of Comparative Zoology, and the Field Museum, and to the American Museum of Natural History for permission to visit the collection. Funding for this project was provided by the National Science Foundation (DEB 0408010, DEB 0445213, and DBI 0400797), an EPSCoR Fellowship and grant s from the following funding sources: Graduate Women in Science, the American Society of Ichthyologists and Herpetologists, the Society for the Study of Amphibians and Reptiles, Sigma Xi (LSU chapter), the University of North Dakota (Graduate School, Office of research and program development and the Department of Biology), the Louisiana State University Museum of Natural Science and Louisiana State University Department of Biological Sciences Graduate Student Organization. I am particularly indebted to the graduate students and staff of the Louisiana State University Museum of Natural Science, including Tammie Jackson, Susan Murray, Nanette Crochett, and Liz Derryberry , as well as Prissy Milligan (Department of Biological Sciences at Louisiana State University) for all their assistance and support. The members of the Austin lab past and present: Nathan Jackson, John McVay, CJ Hayden, Jamie Oaks, Jesse Grismer, Greg Fuerst, and Eric Rittermeyer all deserve special thanks. I owe much thanks to my family (both by blood and by love) and for their encouragement and support throughout this process, and for always believing in me. Lastly, I thank Adam Freedman for his incredible patience, unequaled ability to make me laugh, delicious pasta sauce, and companionship. Again. Your love and support mean more than I can tell you. iv TABLE OF CONTENTS ACKNOWLEDGMENTS………………………………………………………………………...ii ABSTRACT…………………………………..………………………………………………….vi CHAPTER1: INTRODUCTION…………………………………………………………………1 CHAPTER 2: ISLAND AREA AND SPECIES DIVERSITY IN THE SOUTHWEST PACIFIC OCEAN: IS THE LIZARD FAUNA OF VANUATU DEPAUPERATE? ………………………9 CHAPTER 3: DISENTANGLING HISTORY IN OCEANIA: DIFFERENTIATING BETWEEN POLYNESIAN INTRODUCTIONS AND ENDEMIC SPECIES WITHIN THE LIZARD GENUS EMOIA………………………………………………………………………………….38 CHAPTER 4: ENDEMISM, MORPHOLOGICAL CONSERVATISM, AND EVIDENCE FOR ADAPTIVE DIVERSIFICATION IN A CLADE OF LIZARDS FROM OCEANIA……….…71 CHAPTER 5: EMOIA SANFORDI IN THE VANUATU ARCHIPELAGO ………………….112 CHAPTER 6: CONCLUSIONS………………………………………………………………..148 LITERATURE CITED…………………………………………..……………………………..153 APPENDIX 1: SPECIES LISTS FOR ISLAND GROUPS INCLUDED IN CHAPTER 2…...174 APPENDIX 2: PERMISSION FROM ECOGRAPHY…………….……………………..…….180 VITA………….……………….……………….……………….………………....……………181 v ABSTRACT Speciation, geographic variation, and genetic differentiation are fundamental processes that generate diversity, and understanding these processes are major goals of evolutionary biology. Evolutionary phenomena may be more observable on islands as compared to continental landmasses as a result of small population sizes, unoccupied niches, and the relative simplicity of island systems and their populations: physical isolation, shorter (and often well documented) geologic time scale, reduced faunal diversity, and lack of outside faunal influence. Yet, despite their incredible diversity, Pacific island faunas have received little research attention relative to other tropical regions. Using molecular data from several species of scincid lizards in the genus Emoia, I test hypotheses related to the generation and maintenance of biodiversity in Pacific oceanic systems,
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