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A MONOGRAPHIC REVISION OF EUCHAETES HARRIS (LEPIDOPTERA: EREBIDAE: ARCTIINAE) A THESIS SUBMITTED TO THE FACULTY OF UNIVERSITY OF MINNESOTA BY Heather Marie Hendrickson IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE Dr. Susan J. Weller, Advisor Dr. Ralph W. Holzenthal, Co-Advisor December 2014 © Heather Marie Hendrickson 2014 ACKNOWLEDGMENTS The work presented here would not have been possible without the help, guidance, and encouragement granted me by many people along the way. I would like to first express the deep gratitude and appreciation I have for my primary advisor, Dr. Susan J. Weller. She has been an incredible mentor and friend through her willingness to welcome me into her lab and world of research with open arms almost four years ago, her countless encouragements, suggestions, guidances, and help along the way, her ever optimistic spirit in those trying moments, and so much more. The time she has made to teach me about the incredible world of Lepidoptera, and more specifically tiger moths, guide me in dissections and specimen exploration at the microscope, and to review posters, talks, and now this thesis and the aid she has granted there in her constructive criticism has been one of the greatest factors in molding me into an entomologist. Such an investment of her time in me fostered an environment which has helped me to become a better scientific writer, orator, and general communicator. I would also like to thank Susan and her family for their wonderful friendship, inclusion, and support over the years. Much thanks is also due my co-advisor, Dr. Ralph W. Holzenthal, for his willingness to take me under his mentorship and grant me space to work in his lab throughout my time as a master’s student. His ever-open door and willingness to help with literary searches, illustrating, imaging, collecting trips, and so much more has been a tremendous support to me and has contributed to making me a well-rounded scientist by equipping me with applicable skills beyond the microscope. I am also grateful to Ralph and his family for their great friendship, inclusion, and support over the years. I am grateful for the time, encouragement, and guidance granted me by the remaining members of my graduate committee: Dr. Sharon A. Jansa and Dr. Jennifer M. Zaspel (Purdue University). I would like to specifically thank Sharon for her support and guidance in research and her encouragement along the way. I would also like to thank Jennifer for her support and guidance in my research, her assistance in gathering and imaging specimens for me at the AMNH, assisting with analyses, including me in her i field work, and for her priceless friendship over the years, she has become an esteemed colleague and a dear friend, and I thank her for the time she has invested in me. I extend great thanks to all of the funding sources I received in my time as a graduate student. The research completed here would not have been possible without this financial support. Funding was granted by the National Science Foundation (Grant No. 0531639), Minnesota Experiment Station (Grant No. MN-17-022), University of Minnesota Bell Museum of Natural History (Dayton & Wilkie Funds), and the University of Minnesota Department of Entomology (Allan Peterson & Graduate Student Fellowship, Graduate School Block travel grants). I would like to express my gratitude to the many curators, colleagues, and staff of the following natural history collections for their aid in providing me with information, space to work, and loan material: David A. Grimaldi, Suzanne Rab-Green (AMNH), Martin Honey and Geoff Martin (BMNH), Shawn Clark (BYU), John Rawlins (CMNH), Chris Schmidt and Jocelyn Gill (CNC), Richard Hoebeke (CUIC), David Bettman (DMNS), Jerry A. Powell (EME), James H. Boone, M. K. Thayer and S. Huynh (FMNH), Julian Donahue and Weiping Xie (LACM), J. Y. Miller, Jim Hayden, Charles V. Covell and Andy Warren (MGCL), Gary L. Parsons, A. Cognato and M. C. Nielsen (MSUC), Donald Harvey, John W. Brown and Patricia Gentili-Poole (NMNH), Chris J. Marshall (OSAC), Luciana Musetti (OSUC), Edward G. Riley (TAMU), Danny Schpeley and Felix A. H. Sperling (UASM), Doug Yanega (UCRC), Mark O'Brien (UMMZ), Ralph Holzenthal (UMSP), M. J. Paulsen and Brett Ratcliffe (UNSM,). I would also like to thank colleagues Chris Grinter and Ray Nagle for providing me with material from their personal collections. I am also incredibly grateful to the many faculty members, staff, colleagues, and fellow students with whom I have had the privilege of interacting with in my time as a student and, most especially to those with whom I have worked most closely and who have provided me with assistance and guidance along the way: Dr. Robin Thomson, Avery Peace, Taylor Wardwell, Desi Robertson, John Luhman, Roger Blahnik, Dr. Mark Asplen, Trisha Franz, Joe Kaser, Corey McQueen, Roger Moon, William Hutchison, Felicia Christy, and Tomison Olayiwola. I am very grateful to Dr. Christine Dieckhoff, ii Anna Gerenday, and Dr. Robert Suranyi for their assistance in the translation of taxonomic and related ecological works. I would also like to thank the many friends who granted their support and encouragement along the way. In closing, I wish to express my deepest gratitude to my husband, Brad Hendrickson, my stepson David, my parents, Brian & Jayne Cummins, and my three brothers and sister-in-law, Sean & Shyla, Joshua, and Christopher. You have all stood by me continuously, have granted your love, support and encouragement, and so much patience and understanding during these long years—I cannot thank you all enough. This thesis would not have been possible without each and every one of you. iii DEDICATION I dedicate this thesis to Dr. Robert A. Wharton. You have been the greatest of mentors, the closest of friends, and the truest of confidants. You are the reason this work was begun and you are the reason it has been completed. Thank you for all that you have done in my life. iv DISCLAIMER The species names provided in the following text are provisional and are not published within the meaning of the International Code of Zoological Nomenclature (ICZN1999; Article 9). The names and nomenclatural acts established in this dissertation are, therefore, not available within the meaning of the Code (ICZN, Article 10). v ABSTRACT Euchaetes Harris, 1841 is a genus of subfamily Arctiinae, family Erebidae. The genus is distributed throughout the New World, with diversity spanning southern Canada, North America, and Central America. The following is a monographic treatment of the genus. A species-level revision of Euchaetes is completed examining 41 species representing 18 ingroup and 23 outgroup species, scoring a total of 75 morphological characters (191 states) including: head, appendages, male genitalia, female genitalia. The revision treats 18 species, including three newly described species: Euchaetes donahuei, Euchaetes gravastellus, and Euchaetes harveyi. All species are fully illustrated with adult habitus and male and female genitalia (when both were available for study). When known, the biology of a species is summarized. Range maps for each species are also provided based upon the material examined in this study. The type of Euchaetes scepsiformis Graef, 1887 is established as the neotype for Euchaetes bolteri Stretch, 1885. Pygoctenucha Grote, 1883, is placed as a junior subjective synonym of Euchaetes Harris, 1841, based on male and female genitalic characters and the placement of the type species P. terminalis within the Euchaetes clade in the phylogenetic analysis. Former species of Pygoctenucha are placed in Euchaetes as either sensu lato (E. enna, E. azteca) or incertae sedis (E. pyrrhoura and E. clitus). Euchaetes bicolor Rothschild, 1935 is transferred to Agaraea Herrich-Schäffer, 1855 based on its shared adult habitus—body shape, wing shape, and wing venation. Euchaetes fusca (Rothschild 1913) is transferred to Cycnia Hübner, 1818 based on the orb-like uncal processes on the male genitalia and presence of rayflower-like signa in female bursae. Additionally, Pagara Walker, 1856 is vi placed as a junior subjective synonym of Agaraea Herrich-Schäffer, 1855 based on the shared morphology of the male genitalic capsules. vii TABLE OF CONTENTS ACKNOWLEDGMENTS ............................................................................................... i DEDICATION ................................................................................................................. iv DISCLAIMER .................................................................................................................. v ABSTRACT ..................................................................................................................... vi TABLE OF CONTENTS ............................................................................................... viii LIST OF TABLES ............................................................................................................ x LIST OF FIGURES .......................................................................................................... xi LIST OF APPENDICES ................................................................................................. xiv A monographic revision of the Nearctic tiger moth genus Euchaetes Harris, 1841 (Lepidoptera: Erebidae: Arctiinae) ............................................................................... 1 INTRODUCTION ..............................................................................................