Lycaenidae): Phylogeny, Ecology, and Conservation John Mathew Old Dominion University
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Old Dominion University ODU Digital Commons Biological Sciences Theses & Dissertations Biological Sciences Summer 2003 Aphytophagy in the Miletinae (Lycaenidae): Phylogeny, Ecology, and Conservation John Mathew Old Dominion University Follow this and additional works at: https://digitalcommons.odu.edu/biology_etds Part of the Ecology and Evolutionary Biology Commons, Entomology Commons, and the Genetics Commons Recommended Citation Mathew, John. "Aphytophagy in the Miletinae (Lycaenidae): Phylogeny, Ecology, and Conservation" (2003). Doctor of Philosophy (PhD), dissertation, Biological Sciences, Old Dominion University, DOI: 10.25777/v7rh-mb21 https://digitalcommons.odu.edu/biology_etds/74 This Dissertation is brought to you for free and open access by the Biological Sciences at ODU Digital Commons. It has been accepted for inclusion in Biological Sciences Theses & Dissertations by an authorized administrator of ODU Digital Commons. For more information, please contact [email protected]. APHYTOPHAGY IN THE MILETINAE (LYCAENIDAE): PHYLOGENY, ECOLOGY, AND CONSERVATION by John Mathew B.Sc. June 1990, Madras Christian College M.Sc. June 1992, Madras Christian College M.Phil. May 1994, Madras University A Dissertation Submitted to the Faculty of Old Dominion University in Partial Fulfillment of the Requirement for the Degree of DOCTOR OF PHILOSOPHY ECOLOGICAL SCIENCES OLD DOMINION UNIVERSITY August 2003 Approved by: Deborah A. Waller (Co-Director) »mi E. Pierce (Co-Director) H. Savitzky (Member) Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ABSTRACT APHYTOPHAGY IN THE MILETINAE (LYCAENIDAE): PHYTOGENY, ECOLOGY AND CONSERVATION John Mathew Old Dominion University, 2003 Co-Directors of Advisory Committee: Dr. Deborah A. Waller Dr. Naomi E. Pierce Less than 1% of all Lepidoptera are aphytophagous; of these, a considerable proportion is found in the family Lycaenidae. The aphytophagous Lycaenidae are believed to have arisen from a mutualistic template involving ant attendance. With this association firmly in place, it is a relatively simple shift to exploitation, either of the ants themselves, through active camivory on the brood/trophallactic feeding from adults, or by camivory on ant-tended homopterans, with little to no interference by the ants. Among lycaenids, aphytophagy has arisen several times; most spectacularly in the subfamily Miletinae, where all of the approximately 150 species are presumed or known to be aphytophagous. With the exception of the North American speciesFeniseca tarquinius, the subfamily is restricted to the Old World, in particular, Africa and South-East Asia. The focus of this study was a comprehensive review of aphytophagy in the Miletinae, viewed in light of phylogenetic and ecological patterns. A representative genus,Thestor, endemic to southern Africa, was chosen for intense phylogenetic study, where the relationships of nearly all 29 morphological species and subspecies were analyzed, employing the molecular genes Cytochrome Oxidase 1 (COl) and subunit 5 of nicotinamide dinucleotide (ND5). The resultant phylogenies generated were used to inform life-history characters where known. The enigmatic Nearctic species, Feniseca tarquinius, was studied for life-history traits in Massachusetts, New Hampshire, Maine and New Bmnswick, with particular emphasis on diet and vibrational signaling. It was shown that the species produces the longest known acoustic pulse train of any lycaenid. SEM examination of final instar larvae was also undertaken for purposes of comparison with a close relative, the Palaearctic speciesTaraka hamada. An aphytophagous habit adds an extra dimension of complexity to a feeding habit, involving both the prey item Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. and its host-plant resource, rendering its practitioners immediately susceptible to extinction pressure. Conservation concerns for the Lycaenidae, with special emphasis on aphytophagous forms, were hence considered, from the perspective of IUCN Red Data listing, and suggested causes for decline, and appropriate conservation measures discussed, and applied to specific case studies Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. V Dedicated to my family, for accompanying me the length of the way; Deborah Waller, for permitting me to go, and Naomi Pierce, for allowing me to stay. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. vi ACKNOWLEDGMENTS As the inspired author, Vikram Seth, was once wont to say, “My debts are manifold and various,” and I take his words as my own in framing this heartfelt statement of gratitude. Seeing as the debt transcends borders both political and oceanic, I place my sentiments in order of temporal priority, beginning in India in the year 1995, continuing through Norfolk, Virginia (though prefixed with a detour to other parts of the United States), and concluding 8 years thence in Cambridge, Massachusetts. India My first acknowledgment of thanks must be laid at the feet of my parents, the geneticist Dr. Chona Mathew and the once historian and indefatigable advocate of the creative arts Mrs. Elizabeth Mathew, who made my path to academic achievement possible. To the first I attribute my love of biological process (the tantalizing possibilities of dihybrid test crosses) apart from an ingrained passion for cricket and chess, and to the latter, theatre, music, writing, literature, and spirituality, which have in such large measure, become a way of life. Their lasting support, even when they have not always found it easy to understand the choices their son has made, leaves me more grateful than I can say. Thank you. To my maternal grandmother, Mrs. K. V. John, the most cheerful nonagenarian I have ever known, I pay tribute. The unflinching faith she has displayed in the face of recurring adversity remains a lasting inspiration and I have imbibed much strength at her watering trough of generosity and love. Likewise, I thank my maternal aunt, Mrs. P. J. John, for her unwavering support when my own sense of self-belief flickered. Dr. P. Dayanandan, Dr. Leela Madhavan, Dr. K. Sunder Raj, and Mr. Samuel Sukumar, all professors at my Alma Mater, Madras Christian College, composed letters of recommendation that would eventually contribute to my gaining admittance to Old Dominion University. For their support, I remain beholden. The friends I made in India, too numerous to name, remain, outside of my family and select individuals in the United States, the bulwark upon which so much of my security has depended. I thank them. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. To Kerala, Tamil Nadu, India, and South Asia, I reaffirm my fealty. I have drawn upon these my native and adopted regions so much during my years in the United States for my sense of identity, and in turn I hope that I have been, in some wise, an ambassador, for them and for the developing world as a whole. They are the subjects of my visions, now and for the future, and It is my privilege to belong to them. The United States To my beloved sister, my confidante, and among my closest of friends, Rachel Mathew, I can never say thank you sufficiently. She has walked the hardest roads with me, supporting me at every turn. I could not have completed this journey without her. I love her and am beholden for the unconditional love that she has always made available to me. I am incredibly grateful to Raj Mysore, my brother-in-law, for his quiet, instrumental, and constant support over the years, and to Sangeeta Mysore, his sister, who paved the way to my arrival in the United States and spurred me on when my steps flagged. Eapen John, who is as much family today as friend, has, in similar vein, been a tower of strength and counsel. I shall never forget the generosity of those named here. I thank my cousin Champa Saigal for making possible for me so many varied glimpses of this incredible nation. My cousin Johnny John has outfitted me with support and shoes alike, over the years, and for both, and more, I am obliged. To the many family friends I have made in Connecticut, New York and New Jersey, I thank them for enduring moments of sunshine and laughter. My sojourn has consistently been brightened by friends who came before and after me from India and who, in hard places, kept the promise of shared experiences to come, worthwhile. I especially thank Siddharth Balachandran, Shylaja Rao, Peter and Susan D’Souza, Sudha Selva Fatima, Cynthia Isaac, Yohaan John, Meena Natarajan, Sheena Matheiken, and Deepa Ramasinghani. Norfolk, VA My heartfelt thanks redound to the Department of Biological Sciences, Old Dominion University for affording to me admission to the doctoral program in Ecological Sciences with a Special Doctoral Research Assistantship to make the opportunity financially possible. I remain indebted to Dr. Deborah Waller, my advisor, who took a chance on an unknown quantity from halfway across the world, and ensured that my presence at Old Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Dominion could indeed come to pass. Her unwavering backing and efforts on my behalf in registering me semester after semester for five years, and her astute editorial eye have helped keep my feet and my flights of otherwise fancy firmly grounded, and for this, I am profoundly grateful. I thank the other members of my guidance committee, Dr. Alan Savitzky, Dr. Charles Bell, Dr. Kerry Kilbum, and Dr. Rebecca Bray for so kindly taking me through the arduous process of my qualifying examination. I fondly acknowledge Dr. Lytton Musselman for his generosity in making me feel an immediate part of the department and larger community. I am grateful to Dr. Frank Day for his early direction as Graduate Program Director, and express my admiration to Dr. Savitzky for his unstinting support in the same capacity, as well for his being, simply, among the finest teachers I have ever encountered. I thank Dr.