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Revision and Phylogeny of the Limacodid-Group Families, with Evolutionary Studies on Slug Caterpillars (Lepidoptera: Zygaenoidea)

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Revision and Phylogeny of the Limacodid-Group Families, with Evolutionary Studies on Slug Caterpillars (Lepidoptera: Zygaenoidea) MARC E. EPSTEIN W9\ I SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 582 SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of "diffusing knowledge" was expressed by the first Secretary of the Smithsonian. In his formal plan for the institution, Joseph Henry outlined a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropology Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to the Marine Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Folklife Studies Smithsonian Studies in Air and Space Smithsonian Studies in History and Technology In these series, the Institution publishes small papers and full-scale monographs that report the research and collections of its various museums and bureaux or of professional colleagues in the world of science and scholarship. The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world. Papers or monographs submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover. I. Michael Heyman Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 582 Revision and Phylogeny of the Limacodid-Group Families, with Evolutionary Studies on Slug Caterpillars (Lepidoptera: Zygaenoidea) Marc E. Epstein SMITHSONIAN INSTITUTION PRESS Washington, D.C. 19% ABSTRACT Epstein, Marc E. Revision and Phylogeny of the Limacodid-Group Families, with Evolutionary Studies on Slug Caterpillars (Lepidoptera: Zygaenoidea). Smithsonian Contribu- tions to Zoology, number 582, 102 pages, 409 figures, 7 tables, 1996.—The limacodid group, composed of Limacodidae (+ Chrysopolomidae), Dalceridae, Megalopygidae, Aididae, and Somabrachyidae, is revised, and diagnoses, redescriptions, and keys to larval and adult stages are provided for each family. Comparative morphology and ontogeny of the larval stage are emphasized. Immature stages, including first instars, of a number of taxa are figured for the first time. New character data for adults and immatures are presented, including the presence of larval crochets on abdominal segments A2-A7 and A10 in species of Megalopygidae and Limacodidae. Computer-assisted phylogenetic analyses of limacodid-group families using separate character-data sets from adult and immature stages produces congruent cladograms of equal parsimony, with Zygaenidae selected as the outgroup. Megalopygidae (Megalopygi- nae + Trosiinae) is found to be the most basal family among the limacodid group, and Aididae, previously a subfamily of Megalopygidae, is the sister group to the Limacodidae + Dalceridae clade. Chrysopolomidae is placed as a subfamily of Limacodidae based on new character evidence from adult and immature stages. Relationships of Somabrachyidae, Epipyropidae, and Cyclotornidae also are discussed. Previous interpretations of character homologies for abdominal proiegs in the larvae are revised based on the phylogenetic analysis, resulting in a novel hypothesis concerning the evolutionary transformation from proiegs to a slug-like ventral surface. New representations of homology for larval body setae and for juxta and valva of male genitalia also are given. Behavioral observations of immatures and adults of species in the limacodid group are presented, along with a review of their biology. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus). Library of Congress Cataloging-in-Publication Data Epstein, Marc E. Revision and phylogeny of the Limacodid-group families, with evolutionary studies on slug caterpillars (Lepidop- tera : Zygaenoidea) / Marc E. Epstein. P. cm. — (Smithsonian contributions to zoology ; no. 582) Includes bibliographical references (p. 99). 1. Lepidoptera—Classification. I. Title. II. Series. QL1.S54 no. 582 [QL545] 591 s—dc20 [595.78M] 96-16840 ® The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials Z39.48 1984. Contents Page Introduction 1 Acknowledgments 3 Material and Methods 3 Abbreviations 4 History of Classification of the Limacodid Group 5 Biology 6 Larvae 6 Cocoons 8 Adults 10 Eggs 10 Morphology 10 Adults 13 Larvae 25 Pupae 56 Phylogenetic Analysis 61 Data Analysis 61 Results and Discussion 66 Keys to Limacodid-Group Families and Allies 70 Key to Adults 70 Key to Larvae 71 Limacodid-Group Families and Allies 72 MEGALOPYGIDAE Herrich-Schaffer, 1855 72 AlDlDAE Hopp, 1927, new status 75 DALCERIDAE Dyar, 1898 79 LIMACODIDAE Duponchel, 1844 81 CHRYSOPOLOMINAE Aurivillius, 1895, new status 84 SOMABRACHYIDAE Hampson, 1920 85 EPIPYROPIDAE Dyar, 1903, and CYCLOTORNIDAE Meyrick, 1912 86 Ontogeny and Function of Limacodid Spinnerets 87 Homology and Evolution of Larval Body Setae 91 DALCERIDAE 91 LIMACODIDAE 91 Evolution and Origin 93 Evolution of the Ventral Abdominal Surface 94 Prolegs and Crochets 94 Homology of Suckers 95 Function and Adaptation 96 Summary 97 Literature Cited 99 111 Dedication This contribution is dedicated to the memories of my father, Joseph Samuel Epstein, and fellow lepidopterists F. Martin Brown and B. Adrienne Venables. Revision and Phylogeny of the Limacodid-Group Families, with Evolutionary Studies on Slug Caterpillars (Lepidoptera: Zygaenoidea) Marc E. Epstein Introduction surface. There is a wide diversity of larval forms in the limacodid group (Figures 1-6), although it is best known for Limacodidae (=Eucleidae, Cochlidiidae) is the richest spiny larvae that cause dermatitis. The Limacodidae, Mega- (- 1000 species) and most widespread moth family among a lopygidae, and Dalceridae are economic pests of palms and monophyletic group of five families in the Zygaenoidea, which I term the "limacodid group." Distributed worldwide, although other plantation crops in the tropics (Genty et al., 1978; Cock et primarily in the tropics, the family Limacodidae is related to the al., 1987). New World limacodid-group families Dalceridae (84 spp.) Relationships among families in the limacodid group and (Miller, 1994), Aididae (=Aidinae, Megalopygidae, 6 spp.), their allies are poorly understood. In large part this is due to an and Megalopygidae (~ 230 spp.) (Hopp, 1934; Becker, 1995). incomplete knowledge of immature stages. Furthermore, there Two Old World relatives are Chrysopolomidae (35 spp.) and has been little detailed morphological study of immature stages Somabrachyidae (~ 10 spp.). Restricted to Africa and Madagas- or of adults. car (Hering, 1937), Chrysopolomidae is placed herein as a Fortunately the fog is lifting. During the past decade there subfamily of Limacodidae. The Somabrachyidae, often re- have been studies on the limacodid-group families Limacodi- garded as constituting a subfamily of Megalopygidae, are dae (Holloway, 1986; Holloway et al., 1987; Epstein, 1988) found in Africa and in Mediterranean Europe (Geertsema, pers. and Dalceridae (Stehr and McFarland, 1987; Miller, 1994) as comm.; Freina and Witt, 1990). well as on others associated with them, such as parasitic The limacodid group is characterized by external-feeding families Epipyropidae and Cyclotomidae (Davis, 1987; Com- larvae that have retractile heads and either prolegs on mon, 1990); however, comparative morphology from these abdominal segments A2-A7 or a flexible, slug-like ventral studies has been needed in order to gain a better understanding of the interfamilial relationships. I base my phylogenetic (= cladistic) analysis of the limacodid group on the compara- tive morphological data given herein and in other recent studies Marc E. Epstein, Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. cited above. Although I do not treat characters of the Zygaenidae extensively, I examine them in phylogenetic Review Chairman: John M. Burns, National Museum of Natural context of the limacodid group. History, Smithsonian Institution, Washington, D.C. 20560. Dyar (1895a, 1899b) believed ventral suckers in limacodid Reviewers: Jeremy D. Holloway, CA.B. International Institute of Entomology, 56 Queen's Gate, London, SW7 5JR, England; James S. larvae to be derived from fusion of the membranous pads found Miller, Department of Entomology, American Museum of Natural in megalopygids. Hinton (1955:516), however, indicated that History, New York, New York 10024-5192; John E. Rawlins, Section of there was no convincing explanation for the "loss of abdominal Invertebrate Zoology, Carnegie
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