SYSTEMATICS OF THE MEGADIVERSE SUPERFAMILY GELECHIOIDEA (INSECTA: LEPIDOPTEA) DISSERTATION Presented in Partial Fulfillment of the Requirements for The Degree of Doctor of Philosophy in the Graduate School of The Ohio State University By Sibyl Rae Bucheli, M.S. ***** The Ohio State University 2005 Dissertation Committee: Approved by Dr. John W. Wenzel, Advisor Dr. Daniel Herms Dr. Hans Klompen _________________________________ Dr. Steven C. Passoa Advisor Graduate Program in Entomology ABSTRACT The phylogenetics, systematics, taxonomy, and biology of Gelechioidea (Insecta: Lepidoptera) are investigated. This superfamily is probably the second largest in all of Lepidoptera, and it remains one of the least well known. Taxonomy of Gelechioidea has been unstable historically, and definitions vary at the family and subfamily levels. In Chapters Two and Three, I review the taxonomy of Gelechioidea and characters that have been important, with attention to what characters or terms were used by different authors. I revise the coding of characters that are already in the literature, and provide new data as well. Chapter Four provides the first phylogenetic analysis of Gelechioidea to include molecular data. I combine novel DNA sequence data from Cytochrome oxidase I and II with morphological matrices for exemplar species. The results challenge current concepts of Gelechioidea, suggesting that traditional morphological characters that have united taxa may not be homologous structures and are in need of further investigation. Resolution of this problem will require more detailed analysis and more thorough characterization of certain lineages. To begin this task, I conduct in Chapter Five an in- depth study of morphological evolution, host-plant selection, and geographical distribution of a medium-sized genus Depressaria Haworth (Depressariinae), larvae of ii which generally feed on plants in the families Asteraceae and Apiaceae. Host-plant use is commonly studied in this group because of physiological and behavioral responses exhibited by Depressaria pastinacella to furanocoumarins produced by their host plants, yet no species level phylogeny is available. The phylogeny of Nearctic Depressaria is constructed using a morphological data matrix analyzed under the parsimony criterion. This study is the only modern phylogeny of the genus, and includes all North American species but one, and about half the Old World species. I redescribe these species. In Chapter Six I describe nine new species of Scythris Hübner (Scythridinae) from the Galápagos Islands, Ecuador, and provide a key and illustration of genitalia and abdominal modifications. Finally, Chapter Seven represents an application of moth taxonomy to address questions of sampling protocols used for studies of biodiversity and conservation. I use Gelechioidea in eastern North America as indicators of diversity, with attention to the effectiveness of different sampling protocols with respect to active versus passive sampling, and plot-based versus plotless sampling. A list of Gelechioidea was produced from trap sites from an Appalachian forest in southern Ohio. The composition and diversity of Ohio Gelechioidea captured in a passive, plot-based protocol compares favorably to more exhaustive sampling, and reinforces recent (and counterintuitive) recommendations that it is more efficient and repeatable to focus surveys on target groups in focal localities rather than to conduct extensive sampling programs. iii Dedicated to Randle, my best friend And we don't notice any time pass we don't notice anything we sit side by side in every class teacher thinks that I sound funny but she likes the way you sing Tonight I'll dream while I'm in bed when silly thoughts go through my head about the bugs and alphabet and when I wake tomorrow I'll bet that you and I will walk together again cause I can tell that we're going to be friends We are Going to be Friends – The White Stripes iv ACKNOWLEDGMENTS Graduate school is a strange place. I owe many people my gratitude for their patience and support during this time: mentors, colleagues, friends, family, and students. Without their help and encouragement, I would not be the scholar that I am today. I would like to thank my advisor, Dr. John W. Wenzel, for his investment in me. His commitment to my dissertation work and belief in me as a scientist have given me ability to continue my work. I am grateful for the many hours of discussion, writing, proofreading, and editing he has provided. There has not been one day in the last seven years that Dr. Wenzel has been in the museum and has not been available for his graduate students. Dr. Christopher Randle, Husband and Botanist, has been with me since the first day of my post-high school education. With him I have grown as a person and a scientist. Above all, his intellect and admiration inspire me to persevere. Just listen to your heart, that's what I do. I would like to thank my committee members, Dr. Hans Klompen, Dr. Daniel Herms, and Dr. Steven Passoa, who have provided me with vital evaluation of this document. In particular, Dr. Passoa has been critical to the completion of this project. A mentor in my studies of Lepidoptera, he has aided in identification, supplied and assisted v in the collection of specimens, and provided discussion. He has also been a magnificent supplier of items useful to my dissertation, many of which I had no idea I needed. I would like to thank Dr. Jean-François Landry for his attention: it was he who revealed to me the subtle allure of microlepidoptera. My graduate career began (with a study of Coleophora) and ended (with a study of Scythris) under his tutelage. Many other scientists have shaped this work, as well. Jerry Powell, Daniel Rubinoff, David Wahl, Hugo Kons, Lauri Kaila, and David Horn have provided specimens. Ronald Hodges, Richard Brown, Jean-François Landry, David Wagner, John Rawlins, Joël Minet, Patrice Leraut, Lauri Kaila, Jerry Powell, David Adamski, Paul Goldstein, and Fred Stehr have provided comments through the years regarding this work. David Adamski and John Brown were always helpful during museum study at the NMNH. Brian Scholtens provided data from the GSMNP ATBI, and David Wagner provided unpublished data for Lepidoptera of Connecticut. John Herbert, Foster Purrington, Chad Schone, and Eric Dotseth provided helpful comments on the LAWCO manuscript. John Herbert and Foster Purrington helped with the initial moth sorting. Christopher Randle was immensely helpful in DNA alignment, phylogenetic analysis, and discussion. John Wenzel translated text written in French, and Christopher Randle translated text written in German. Mary Daniels provided technical assistance in the initial stages of the molecular projects. My friends have had an equally significant influence. Conversations with my lab mates of the past and present have hugely benefited my work as a graduate student. In particular, Todd Blackledge, Kurt Picket, Hojun Song, Joe Raczkowski, and Ryan Caesar have played a central role in entomological and phylogenetic discussions. Hojun Song, vi with whom I have shared an office for several years, has not only provided stimulating discussion regarding the evolution of insect genitalia but also helped to keep me sane (or at least minimized a drastic increase in madness). I thank all of my friends at the Museum of Biological Diversity, especially members of the Phylogenetic Discussion Group, who helped to polish my phylogenetic skills. I also thank all of my friends in the departments of Entomology and EEOB who have helped me through the years. Mark Mort and Christopher Randle shared their limited space with me while I was in Kansas. I thank my family, the Randles (Biz, Noël, Jinx (my own private editor!), Shonkie, William, Alex, Lizzie, Jackson, Dan-O, Fiona, Bridget, Jack, and Violet) and the Buchelis (Momma, Dad, Grammy, Pat, Brad, Patrice, Baby Joey, and Emma), for their love and support. Finally, I thank my mother, Christina Arlia, who I believe is responsible for my love of insects. When I was a small child, I would bring to her tokens of my love: hideous six-legged tokens that bit and stung and made terrible smells. Instead of freaking out and killing them, she would tell me that they were "mommy-bugs" that had to go home to their babies. I grew up thinking that all insects were wonderful and caring creatures, just like my mother. vii VITA May 22, 1973……………………………..………..……… Born – Tampa, Florida 1996………………………………………………..………. B.A. Biology, Hiram College 1999………………………………………………………... M.S. Entomology, The Ohio State University PUBLICATIONS Bucheli, S. R., J.-F. Landry, and J. W. Wenzel. 2002. "Cladistic Analysis of Larval Case Architecture and Implications of Host-Plant Associations for North American Coleophora (LEPIDOPTERA: COLEOPHORIDAE)", Cladistics 18, 71-93. Bucheli, S. R. and J. W. Wenzel. 2005. Gelechioidea (Insecta: Lepidoptera) systematics: A reexamination using combined morphology and mitochondrial DNA data. Molecular Phylogenetics and Evolution 35, 380-394. Bucheli, S. R., D. Horn, and J. W. Wenzel. In Press. Biodiversity of Gelechioidea (microlepidoptera): An assessment of a re-established Appalachian forest in southern Ohio. Biodiversity and Conservation. FIELDS OF STUDY Major Field: Entomology Specialization: Systematics of Lepidoptera, especially Gelechioidea and other microlepidoptera, phylogenetics, morphological evolution, and host-plant evolution. viii TABLE OF CONTENTS Abstract………………………………………………………………………………ii Dedication…………………………………………………………………………...iv