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Systematics of Neotropical Satyrine Butterflies Louisiana State University LSU Digital Commons LSU Historical Dissertations and Theses Graduate School 2001 Systematics of Neotropical Satyrine Butterflies (Nymphalidae: Satyrinae: Euptychiina) Based on Larval Morphology and DNA Sequence Data and the Evolution of Life History Traits. Debra Lynne Murray Louisiana State University and Agricultural & Mechanical College Follow this and additional works at: https://digitalcommons.lsu.edu/gradschool_disstheses Recommended Citation Murray, Debra Lynne, "Systematics of Neotropical Satyrine Butterflies (Nymphalidae: Satyrinae: Euptychiina) Based on Larval Morphology and DNA Sequence Data and the Evolution of Life History Traits." (2001). LSU Historical Dissertations and Theses. 424. https://digitalcommons.lsu.edu/gradschool_disstheses/424 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 Historical Dissertations and Theses by an authorized administrator of LSU Digital Commons. For more information, please contact [email protected]. INFORMATION TO USERS This manuscript has been reproduced from the microfilm master. UMI films the text directly from the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter face, while others may be from any type of computer printer. The quality of this reproduction is dependent upon the quality of the copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandard margins, and improper alignment can adversely affect reproduction. In the unlikely event that the author did not send UMI a complete manuscript and there are missing pages, these will be noted. Also, if unauthorized copyright material had to be removed, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, charts) are reproduced by sectioning the original, beginning at the upper left-hand comer and continuing from left to right in equal sections with small overlaps. Photographs included in the original manuscript have been reproduced xerographically in this copy. Higher quality 6* x 9” black and white photographic prints are available for any photographs or illustrations appearing in this copy for an additional charge. Contact UMI directly to order. ProQuest Information and Learning 300 North Zeeb Road. Ann Arbor, Ml 48106-1346 USA 800-521-0600 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. NOTE TO USERS This reproduction is the best copy available. UMI' Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. SYSTEMATICS OF NEOTROPICAL SATYRINE BUTTERFLIES (NYMPHALIDAE: SATYRINAE: EUPTYCHDNA) BASED ON LARVAL MORPHOLOGY AND DNA SEQUENCE DATA AND THE EVOLUTION OF LIFE HISTORY TRAITS 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 Entomology by Debra L. Murray B.A., University of North Carolina at Chapel Hill, 1987 December 2001 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. UMI Number. 3042640 UMI’ UMI Microform 3042640 Copyright 2002 by ProQuest Information and Learning Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code. ProQuest Information and Learning Company 300 North Zeeb Road P.O. Box 1346 Ann Arbor, Ml 48106-1346 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. To the Murray clan: may you still fly the skies, tend the earth, nurse the creatures, manage the accounts, watch the birds, stain the glass, and research the mysteries of this world. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ACKNOWLEDGMENTS I am deeply indebted to the many people who helped bring this project to fruition. The development of my project began in Ecuador while I was serving as a Peace Corps Volunteer. For encouragement, support and assistance during this period, special thanks go to Alejandro Suarez. Without his help, I could never have accomplished as much as I did. Equally helpful was the support staff of Fundacion Jatun Sacha and the many volunteers at the biological station. In particular, thanks to David Neill for plant identifications and support. For assistance in the field, thanks to my two sisters Marion and Melissa Murray, Barb Swartz, Saara DeWalt, Rose Hanrahan, Celia Harvey, and Gabriel Tapuy. I am also indebted to Francisco Panches for astutely placing me as a volunteer in a small biological station, and for the excellent support given by The United States Peace Corps. The few early lepidopteran visitors to the biological station, Stan Nicoly, Dave Arenholtz, Carla Penz, and Bob Srygley, provided critical direction, advice, and support. I am also greatly indebted for the support of Phil DeVries, for placing faith in someone he’s never met and for the stimulating discussions on butterflies. During those infrequent visits to the United States, I will forever appreciate the generosity and support of Bob Robbins during my visits to the Smithsonian Institution. Equally important to the advancement of this project was the support and advice of Lee and Jackie Miller during my visits to the Allyn Museum. At Louisiana State University, special thanks to my advisors Dorothy Prowell and Chris Carlton and to my committee members Tom Riley and Fred Sheldon. This has been a great learning experience for me and hopefully for my advisors as well. I have iii Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. certainly enjoyed the lengthy discussions on homologous structures with a non- lepidopterist, Chris Carlton. The Organization of Tropical Studies, Rio Bravo Research Station, Yasuni Biological Station, and Maquipucuna Research Station provided excellent support during my additional field collecting trips. I also had the opportunity to visit several museums during my field trips and special thanks for assistance to Gerardo Lamas and Giovanni Onore. Many organizations provided monetary support during my time at Louisiana State University, the Organization for Tropical Studies, American Women in Science, Sigma Xi, and LSU Graduate School. Larvae donated or loaned to me by Andres Suarez, Miguel Suarez, Andrei Sourakov, and Texas A&M Museum, Department of Entomology were invaluable to this project. I also greatly appreciate the generosity of Andre Freitas for allowing me to examine his wonderful collection of larvae. Many people also provided DNA material and I am grateful to Jim Tuttle, Dan Lang, Andy Brower, Chris Carlton, and Victoria Moseley. For assistance in molecular work, there were many people who provided advice and use of their computers. Thanks go to Susan Pell, Fred Sheldon, Lowell Urbatsch, Roland Roberts, Michael Hellberg, Frank McKinney, and Meredith Blackwell. I am indebted to Susan Pell for invaluable assistance in assembling the many figures in this dissertation. And finally, thanks for fruitful discussion on satyrines, rearings, and butterfly systematics go to Andres Fretias, Gerardo Lamas, Bob Robbins, Lee Miller, Susan Weller, Andy Brower, David Lees, Dick Vane-Wright, and Keith Brown. iv Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. TABLE OF CONTENTS ACKNOWLEDGMENTS.................................................................................................. iii ABSTRACT........................................................................................................................vi CHAPTER 1 INTRODUCTION........................................................................................ 1 2 DESCRIPTIONS OF EUPTYCHIINE BUTTERFLY LARVAE.......... 21 3 SYSTEMATICS AND EVOLUTION OF THE EUPTYCHIINE BUTTERFLIES BASED ON LARVAL MORPHOLOGY.................109 4 PHYLOGENETIC RELATIONSHIPS OF THE SUBTRIBE EUPTYCHHNA: COMBINED MOLECULAR AND MORPHOLOGICAL ANALYSES.......................................................262 5 EVOLUTION OF LIFE HISTORY TRAITS WITHIN SATYRINE BUTTERFLIES...................................................................................... 326 6 CONCLUSIONS......................................................................................358 VITA................................................................................................................................ 367 v Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ABSTRACT Although satyrine butterflies (Lepidoptera: Nymphalidae: Satyrinae) are highly diverse and found worldwide in most terrestrial habitats, they have received little attention from systematists. This research focuses on on* of the largest satyrine groups, the euptychiines, found from central United States to Argentina, with the greatest diversity in the Amazon. The subtribe Euptychiina has a troubled taxonomic history. Most genera were erected without diagnoses and placed within the subtribe without being examined. Before this project
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