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Phylogeny & Revision of the Genus Heraclides

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Phylogeny & Revision of the Genus Heraclides PHYLOGENY AND REVISION OF THE GENUS HERACLIDES HÜBNER, 1819 (LEPIDOPTERA: PAPILIONIDAE: PAPILIONINAE: PAPILIONINI) By DELANO ST. AUBYN LEWIS 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 2010 1 © 2010 Delano St. Aubyn Lewis 2 To my lovely wife, Dadria, and our first child 3 ACKNOWLEDGMENTS I thank the members of my Graduate Committee for putting up with the many challenges that have presented themselves in the process of me completing and presenting this research and dissertation. I would like to thank Dr. Thomas C. Emmel, Chair of my Graduate Committee, for seeing potential in me, and for his constant patient and thoughtful counsel. His genuine interest in my overall wellbeing, not just my academic wellbeing, has enabled me to pursue my dreams. Dr. Jaret Daniels’ guiding hand in helping me to learn the nuances of grant writing, and his encouragements and advice on various aspects of my research were without equal. Dr. Jacqueline Miller and Dr. Keith Willmott were instrumental in rescuing me from sure disaster when, almost a year into my topic, I discovered that my previous research and dissertation topic was published. They offered many suggestions and assistance in gathering material and helped to make sure no one else was doing the same project. My external member, Dr. Charles Baer, has a good eye for detail and helped to resolve any unanswered questions while effectively representing the interests of the University of Florida Graduate School. Special mention needs to be made of Dr. Charles V. Covell Jr. The techniques I learned from him during my Master’s work on Cyllopoda moths proved to be more valuable than I would ever imagine, once I started to break swallowtail butterflies apart. He has also been very considerate of the changes I had to make when I needed to restructure my graduate committee due to significant changes in the nature of my research. The many brief but frank conversations held in the hallways and offices of the McGuire Center with Dr. Andrei Sourakov and Dr. Deborah Matthews-Lott opened my eyes to the hurdles I would possibly face as I attempted to tackle the oxymoronic state of this well known but poorly studied group. Special thanks to Dr. Andrew Warren, whose expert eye and knowledge of the butterfly fauna of the neotropics, and especially Mexico, proved indispensable in deciphering the 4 intricacies of relationships between these species. His constant willingness to lend an extra pair of eyes, and at times hands and feet, when trying to wander through this maze, made the work less stressful if not easier. The assistance of Dr. George T. Austin (1943 – 2009) in his capacity as McGuire Center Collection Manager, answering my numerous questions, and discussing difficult topics with me at a moment’s notice, were most valuable. I will always cherish the many moments I lurked at his office door in anticipation of his words of wisdom, or went searching for him throughout the collections to seek his advice. The passing of a great scientist, a strong mentor, and friend has been a great loss. It would be remiss of me to not thank my parents and siblings who have constantly supported my interest in science, and my wife Dadria who, though wrapped up in her own graduate journey to doctoral dissertation ‘la la land’ while being pregnant with our first child, found the time to encourage and strengthen me when it was most needed. The assistance of my family, friends, and colleagues have helped to ‘tie up the loose ends’ created by the meandering path this dissertation has taken on its journey to apparent academic profitability, and I am most grateful to all. 5 TABLE OF CONTENTS page ACKNOWLEDGMENTS ...............................................................................................................4 TABLE OF CONTENTS .................................................................................................................6 LIST OF TABLES ...........................................................................................................................8 LIST OF FIGURES .........................................................................................................................9 ABSTRACT ...................................................................................................................................16 CHAPTERS 1 INTRODUCTION AND LITERATURE REVIEW ..............................................................18 Introduction .............................................................................................................................18 Historical Classification and Phylogenetics ...........................................................................19 Phylogenetic Placement of Heraclides ...................................................................................21 Historical Taxonomic Treatment of Heraclides .....................................................................24 Research Objectives and Questions ........................................................................................26 2 SPECIES AND SUBSPECIES ...............................................................................................28 The Gordian Knots of Taxonomy ...........................................................................................28 What is a Species? ..................................................................................................................30 The Subspecies .......................................................................................................................33 3 MATERIALS AND METHODS ...........................................................................................37 Sources of Material Examined ................................................................................................37 Specimen Preparation .............................................................................................................37 Taxon Sampling ......................................................................................................................39 Genes and Alignment of Molecular Sequence Data ...............................................................40 Phylogenetic Reanalysis of Molecular Data ...........................................................................40 Phylogenetic Analysis of Morphological Data .......................................................................42 Morphological Characters ......................................................................................................45 4 RESULTS ...............................................................................................................................81 Phylogenetic Reanalysis of Molecular Data ...........................................................................81 Phylogenetic Analysis of Morphological Data .......................................................................84 5 HERACLIDES PHYLOGENY ...............................................................................................99 6 Molecular Phylogeny ..............................................................................................................99 Morphological Phylogeny ....................................................................................................101 Group Structure and Biogeography ......................................................................................102 6 SPECIES ACCOUNTS FOR THE GENUS HERACLIDES................................................110 Heraclides Hübner, 1819 ......................................................................................................110 Key to Heraclides Adults .....................................................................................................110 The “thoas” Subgroup ..........................................................................................................113 Heraclides thoas (Linnaeus, 1771) ...................................................................................113 Heraclides cresphontes (Cramer, 1777) ...........................................................................118 Heraclides homothoas (Rothschild & Jordan, 1906) ........................................................120 Heraclides melonius (Rothschild & Jordan, 1906) ...........................................................121 Heraclides paeon (Boisduval, 1836) ................................................................................122 Heraclides aristor (Godart, 1819).....................................................................................124 Heraclides caiguanabus (Poey [1852]) ............................................................................125 The “thersites” Subgroup .....................................................................................................126 Heraclides thersites (Fabricius, 1775) ..............................................................................126 Heraclides androgeus (Cramer, 1775) ..............................................................................128 Heraclides astyalus (Godart, 1819) ..................................................................................130 Heraclides ornythion (Boisduval, 1836) ...........................................................................135 Heraclides andraemon Hübner, [1823] ............................................................................136
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