Phylogenetic Relationships within Heliodinidae and Systematics of Moths Formerly Assigned to Heliodines Stainton (Lepidoptera: Yponomeutoidea) Yu-Feng Hsu and Jerry A. Powell Phylogenetic Relationships within Heliodinidae and Systematics of Moths Formerly Assigned to Heliodines Stainton (Lepidoptera: Yponomeutoidea) Yu-Feng Hsu and Jerry A. Powell UNIVERSITY OF CALIFORNIA PRESS Berkeley • Los Angeles • London UNIVERSITY OF CALIFORNIA PUBLICATIONS IN ENTOMOLGY Editorial Board: Penny Gullan, Bradford A. Hawkins, John Heraty, Lynn S. Kimsey, Serguei V. Triapitsyn, Philip S. Ward, Kipling Will Volume 124 UNIVERSITY OF CALIFORNIA PRESS BERKELEY AND LOS ANGELES, CALIFORNIA UNIVERSITY OF CALIFORNIA PRESS, LTD. LONDON, ENGLAND © 2005 BY THE REGENTS OF THE UNIVERSITY OF CALIFORNIA PRINTED IN THE UNITED STATES OF AMERICA Library of Congress Cataloging-in-Publication Data Hsu, Yu-Feng, 1963– Phylogenetic relationships within Heliodinidae and systematics of moths formerly assigned to Heliodines Stainton (Lepidoptera: Yponomeutoidea) / Yu-Feng Hsu and Jerry A. Powell. p. cm. Includes bibliographical references. ISBN 0-520-09847-1 (paper : alk. paper) — (University of California publications in entomology ; 124) 1. Heliodinidae—Classification. 2. Heliodinidae—Phylogeny. I. Title. II. Series. QL561.H44 H78 595.78 22—dc22 2004058800 Manufactured in the United States of America The paper used in this publication meets the minimum requirements of ANSI/NISO Z39.48-1992 (R 1997) (Permanence of Paper). Contents Acknowledgments, ix Abstract, xi Introduction ...................................................... 1 Problems in Systematics of Heliodinidae and a Historical Review ............ 4 Material and Methods ............................................ 6 Specimens and Depositories, 6 Dissections and Measurements, 7 Wing Venation Preparation, 7 Scanning Electron Microscope Preparation, 8 Species Discrimination and Description, 8 Larval Rearing Procedures, 8 Phylogenetic Methods, 9 Phylogeny of Heliodinidae ......................................... 12 Monophyly of Heliodinidae, 12 Characters and States Used in the Cladistic Analyses, 13 Results of Phylogenetic Analyses, 19 Systematic Account ................................................ 27 Family Heliodinidae Heinemann, 27 Key to the Genera of Heliodinidae and Species Groups of Aetole, 29 Heliodines assemblage, 31 Heliodines Stainton, 31 H. roesella (Linnaeus), 32 Epicroesa assemblage, 35 Epicroesa Meyrick, 35 Philocoristis Meyrick, 35 Lamprolophus assemblage, 36 Lamprolophus Busck, 36 Aetole assemblage, 36 ‘Heliodines’ princeps subassemblage, 36 ‘Heliodines’ princeps lineage, 36 ‘Heliodines’ princeps Meyrick, 37 Lithariapteryx subassemblage, 38 Lithariapteryx Chambers, 38 Neoheliodines Hsu, new genus, 38 Key to species of Neoheliodines, 40 N. nyctaginella (Gibson), new combination, 40 N. eurypterus Hsu, new species, 43 N. hodgesi Hsu, new species, 43 N. megostiellus Hsu, new species, 46 N. melanobasilarus Hsu, new species, 47 N. vernius Hsu, new species, 48 N. cliffordi (Harrison and Passoa), new combination, 51 N. arizonense Hsu, new species, 53 N. albidentus Hsu, new species, 55 Aetole subassemblage, 56 Embola Walsingham, 56 Key to species of Embola of North and Central America, 58 E. ionis (Clarke), new combination, 59 E. ciccella (Barnes and Busck), new combination, 61 E. cyanozostera Hsu, new species, 63 E. albaciliella (Busck), new combination, 64 E. friedlanderi Hsu, new species, 66 E. melanotela Hsu, new species, 67 E. autumnalis Hsu, new species, 68 E. sexpunctella (Walsingham), new combination, 69 E. dentifer Walsingham, 70 E. powelli Hsu, 71 E. xanthocephala Walsingham, 73 Heliogemma Hsu, new genus, 74 H. gigantea Hsu, new species, 75 H. grandis Hsu, new species, 76 H. preclara Hsu, new species, 77 Pseudastasia Walsingham, 78 Euheliodines Hsu, new genus, 79 E. chemsaki Hsu, new species, 79 E. jaliscella Hsu, new species, 81 Aetole Chambers, revised status, 82 Eximia Group, 83 A. eximia Hsu, new species, 84 A. insolita Hsu, new species, 86 Bella Group, 87 A. bella Chambers, 88 A. schulzella (Fabricius), new combination, 93 A. fulgida Hsu, new species, 95 A. prenticei Hsu, new species, 96 Tripunctella Group, 99 A. tripunctella (Walsingham), new combination, 101 A. unipunctella (Walsingham), new combination, 103 A. inusitata Hsu, new species, 107 A. aprica Hsu, new species, 108 A. cera Hsu, new species, 108 A. favonia Hsu, new species, 110 Extraneella Group, 111 A. extraneella (Walsingham), new combination, 112 Calcifer Group, 116 A. calcifer (Walsingham), new combination, 117 A. calciferoides Hsu, new species, 119 Galapagoensis Group, 121 A. galapagoensis (Heppner and B. Landry), new combination, 122 Copocentra Meyrick, 123 Scelorthus Busck, 124 Incertae Sedis, 124 ‘Heliodines’ aureoflamma Walsingham, 124 ‘H.’ marginata Walsingham, 125 ‘H.’ urichi Busck, 126 Biology ........................................................ 127 Larval Host Plant Relationships, 127 Life History, Voltinism, and Diapause, 129 Behavior, 129 Mating, 129 Hind Leg Posture, 130 Oviposition, 130 Larval Biology, 130 Pupal Biology, 131 Summary ...................................................... .133 Tables, 134 Literature Cited, 148 Figures 26–220, 159 Acknowledgments We are grateful to the following for assistance in field work, comments on the manuscript, the loan of specimens, and other contributions: John Brown, Systematic Entomology Laboratory, USDA, U.S. National Museum of Natural History, Washington, D.C.; John Chemsak, Essig Museum of Entomology, University of California, Berkeley; David Furth, formerly at the Museum of Comparative Zoology, Harvard University, Cambridge; Reinhard Gaeike, Deutsches Entomologisches Institut, Eberswalde; Terry Harrison, University of Illinois, Urbana; John Heppner, Florida State Collection of Arthropods, Gainesville; Ronald Hodges, Eugene, Oregon, formerly USDA, U.S. National Museum of Natural History, Washington, D.C.; Niels Kristensen, University of Copenhagen, Universitetsparken; Bernard Landry, Muséum d’histoire naturelle, Geneva; Jean-François Landry, Canadian National Collection, Ottawa; Martin Lodl, Naturhistorisches Museum Wien, Vienna, Austria; Carolina Martin, Museo Nacional de Ciencias Naturales, Madrid, Spain; Scott Miller, formerly of the Bishop Museum, Honolulu, Hawaii; Sigeru Moriuti and Toshiya Hirowatari, University of Osaka Prefecture, Sakai, Japan; Norman Penny, California Academy of Sciences, San Francisco; Michael Prentice, Jorge Santiago-Blay, and Robert Zuparko, University of California, Berkeley; Edward Riley, Texas A.&M. University, College Station; Michael Shaffer, Natural History Museum, London; Frederick Stehr, Michigan State University, East Lansing; David Wagner, University of Connecticut, Storrs; Shen-Horn Yen, Imperial College at Silwood Park, Ascot. Brown and Harrison provided detailed critical reviews of the manuscript that were exceptionally useful. Many plant identifications were made by Barbara Ertter, UC Herbarium, which retains the voucher specimens. YFH is especially indebted to John Doyen and Thomas Duncan, University of California, Berkeley, for many helpful comments and suggestions and Christopher Meacham and Brent Mishler for assistance with cladistic analysis. Finally, we are indebted to Gordon Nishida, who prepared the illustrations in electronic format, and provided other computer expertise enabling completion of the final copy. ix Abstract Heliodinidae traditionally have been characterized on the basis of forewing venation, color and scaling, and perch behavior, but none of these attributes defines monophyly. We identify four uniquely derived autapomorphies for the family: (1) M vein of forewing two-branched, presumably with M3 lost; (2) tegumen greatly expanded posteriorly, forming a sclerotized, hollow tube; (3) ventral branches of apophyses anteriores originating from a fused transverse bridge; and (4) pupa with long, stiff dorsal and lateral setae. Phylogenetic relationships among genera and species groups of world Heliodinidae are constructed using parsimony and character compatibility as optimality criteria, with representatives of six other families of Yponomeutoidea as outgroups. Results of the analyses show Heliodines Stainton, as formerly recognized (i.e., all the species with conspicuous red markings on the forewings), to be a polyphyletic assemblage. To accommodate the New World fauna, two old names, Aetole Chambers and Embola Walsingham, have been resurrected from synonymy, and three new genera are described: Neoheliodines Hsu (Type species: Heliodines nyctaginella Gibson, 1914), Heliogemma Hsu (Type species: H. gigantea Hsu), and Euheliodines Hsu (Type species: E. chemsaki Hsu). The South American genus Crembalastis Meyrick is synonymized with Embola. A descriptive taxonomy is provided for North and Central American and Caribbean species formerly assigned to Heliodines; 45 species are treated, 25 of which are described as new: Aetole fulgida (TL: Sinaloa, Mexico), A. prenticei (Calif.), A. eximia (Baja Calif., Mexico), A. insolita (El Salvador), A. cera (Calif.), A. favonia (Calif.), A. inusitata (Baja Calif., Mexico), A. aprica (Texas), A. calciferoides (Veracruz, Mexico); Embola autumnalis (Ariz.), E. cyanozostera (Nevada), E. friedlanderi (San Luis Potosí, Mexico), E. melanotela (Haiti); Euheliodines chemsaki (S. L. P., Mexico), E. jaliscella (Jalisco, Mexico); Heliogemma gigantea (Jalisco, Mexico), H. grandis (Tamaulipas, Mexico), H. preclara (Jalisco, Mexico); Neoheliodines albidentus (Ariz.), N. arizonense