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THE EVOLUTIONARY HISTORY of HAWAIIAN LEAF-MINING MOTHS in the GENUS Philodoria (GRACILLARIIDAE: LEPIDOPTERA)

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THE EVOLUTIONARY HISTORY of HAWAIIAN LEAF-MINING MOTHS in the GENUS Philodoria (GRACILLARIIDAE: LEPIDOPTERA) THE EVOLUTIONARY HISTORY OF HAWAIIAN LEAF-MINING MOTHS IN THE GENUS Philodoria (GRACILLARIIDAE: LEPIDOPTERA) By CHRISTOPHER AGUSTIN JOHNS 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 2017 © 2017 Christopher Agustin Johns To my friends, family, and the Islands ACKNOWLEDGMENTS I thank Charmian Dang, Betsy Gangé, and Cynthia King (Hawai‘i Department of Land and Natural Resources, Division of Forestry and Wildlife) for permitting; Keahi M. Bustamente (Leeward Haleakalā Watershed Restoration Project) and Natalia Tangalin (National Tropical Botanical Garden) for endangered plant identification, collection assistance, reports on field observations, and project planning; Margaret J. Sporck- Koehler (Hawai‘i Department of Land and Natural Resources) for assistance with fieldwork, land access, and gathering information on host plant conservation statuses; Pat Bily and Russel Kalstrom (Nature Conservancy) for land access; Pomaika‘i Kanaiaupio-Crozier, Lono Dunn, Daniel Tanaka, and Joe Ward (Pu‘u Kukui Watershed Preserve) for land access and field support; West Maui Mountain Watershed Partnership for logistical support; Avery Chumbley for West Maui access to land; Butch Haase (Moloka‘i Coastal Land Trust), Ane Bakutis (Hawai‘i Plant Extinction Prevention Program) for support of fieldwork on Moloka‘i; William P. Haines (Univ. of Hawai‘i, Mānoa), Karl Magnacca (Oahu Army Natural Resources Program), Hank Oppenheimer (Hawai‘i Plant Extinction Prevention Program), Susan Ching Harbin (Hawai‘i Plant Extinction Prevention Program), Joel Q. C. Lau, and Steve Montgomery for helpful suggestions on insect and host plant locations; Jesse Eiben (University of Hawai‘i Hilo), Mark Wasser (Hawai‘i Volcanoes National Park), and Melissa Dean (Hawai‘i Experimental Tropical Forest) for help coordinating fieldwork on Hawai‘i Island; Diana Crow (Ulupalakua Ranch) for assistance with access on East Maui; Marie VanZandt (Auwahi Wind Energy) and Fritz Klasner (Office of Mauna Kea Management) for permitting and access; Daniel Rubinoff (Univ. of Hawai‘i, Mānoa) for the initial development of the project; Sheperd Meyers and Barbara Kennedy for allowing 4 examination of insect and herbarium specimens at the Bernice Pauahi Bishop Museum; Donald R. Davis (Smithsonian National Museum) and Shigeki Kobayashi (Osaka Prefecture University) for assisting with specimen identification; Jesse W. Breinholt, Emmanuel F.A. Toussaint, David Plotkin, Francesca Ponce, Kelly Dexter, and Lei Xiao (Florida Museum of Natural History) for assistance with lab work and data analysis; and Jonathan Bremer (Florida Museum of Natural History) for assistance in herbarium analyses. Special thanks to Thomas C. Emmel (McGuire Center for Lepidoptera and Biodiversity) for guidance and early support of the project; University of Florida Department of Entomology for initial support of the research; faculty and staff of the McGuire Center for Lepidoptera and Biodiversity and the members of the Kawahara Lab for support and collaboration; Nico Cellinese, Gustav Paulay, and Jiri Hulcr (University of Florida) for their support of my academic and personal development; and the attendants of the 2015 (Japan) and 2016 (Hawai‘i) International Symposium on Gracillariidae for their comments and direction on research questions. This research was supported by the National Science Foundation (Graduate Research Fellowship to C. A. J.; DEB #1354585 to A. Y. K); the National Geographic Society (#C283-14 to C. A. J., and #9686-15 to A. Y. K.); the Entomological Society of America (2014 SysEB Travel Award to C. A. J); the University of Florida’s Tropical Conservation and Development Program (2014 Field Research Grant to C. A. J); the Florida Museum of Natural History (Lockhart Fellowship and Travel Award to C. A. J.); the International Biodiversity Foundation (Documentary Production Grant to C. A. J.); and the Society for Systematic Biologists’ (2012 SSB MiniARTS Grant to A. Y. K). 5 This body of work would not have been possible without the support of Thomas Johns, Elizabeth Agustin, DeeAnn Johns, Ryan Johns, Sophia Ramos, Geena Hill, Rick Merker, Emily Johns, Juli Burden, Niko Marshall, Parker Pflaum, Kai Moreb, Keahi Bustamente, Nora Beale, and Calvin Beale. Lastly, I thank Akito Y. Kawahara, for his tremendous friendship, guidance, and support. 6 TABLE OF CONTENTS page ACKNOWLEDGMENTS ...................................................................................................... 4 LIST OF TABLES ................................................................................................................ 9 LIST OF FIGURES ............................................................................................................ 10 LIST OF OBJECTS ........................................................................................................... 11 ABSTRACT ........................................................................................................................ 12 CHAPTER 1 MOLECULAR PHYLOGENY, REVISED HIGHER CLASSIFICATION, AND IMPLICATIONS FOR THE CONSERVATION OF ENDANGERED HAWAIIAN LEAF-MINING MOTHS (LEPIDOPTERA: GRACILLARIIDAE: Philodoria) .............. 13 Introduction ................................................................................................................. 13 Materials and Methods ............................................................................................... 17 Taxon Sampling, Amplification, and Sequencing ................................................ 17 Phylogenetic Analyses ......................................................................................... 20 Hypothesis Testing ............................................................................................... 21 Results ........................................................................................................................ 23 Discussion ................................................................................................................... 23 Subgeneric Classification ..................................................................................... 23 Philodoria: Implications for Ecology and Conservation ....................................... 25 2 EVIDENCE OF AN UNDESCRIBED, EXTINCT Philodoria SPECIES (LEPIDOPTERA: GRACILLARIIDAE) FROM HAWAIIAN Hesperomannia HERBARIUM SPECIMENS ........................................................................................ 32 3 CONSERVATION MEDIA AS A TOOL TO RAISE AWARENESS FOR Philodoria MOTHS AND THEIR CONSERVATION .................................................. 36 Background ................................................................................................................. 36 Engagement ................................................................................................................ 38 Future Considerations ................................................................................................ 38 4 EVOLUTIONARY HISTORY OF Philodoria ............................................................... 40 Introduction ................................................................................................................. 40 Hawaiian Geology ................................................................................................ 40 Origin of the Hawaiian Biota ................................................................................ 42 Calibration Strategy .............................................................................................. 42 7 Philodoria .............................................................................................................. 44 Materials and Methods ............................................................................................... 45 Taxon Sampling ................................................................................................... 45 Sample Preparation, Sequencing, Data Processing, and Dataset Construction ...................................................................................................... 51 Phylogenomic Analyses ....................................................................................... 53 Divergence Time Estimation ................................................................................ 54 Historical Biogeography and Ancestral Host Plant Reconstruction .................... 57 Phylogenomics ..................................................................................................... 59 Divergence Time Estimation ................................................................................ 64 Historical Biogeography and Ancestral Host Plant Association ......................... 65 Discussion ................................................................................................................... 70 Phylogenomics ..................................................................................................... 70 Dataset Subsampling ........................................................................................... 71 Calibration Strategy .............................................................................................. 72 Evolution of Philodoria and Their Host Associations .......................................... 74 Conservation of Philodoria Moths and Their Host Plants ................................... 80 APPENDIX A ACCESSION AND
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