With Phenetic and Cladistic Analyses
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mversity o ications ENTOMOLOGY Volume 113 Systematics and Biology of the Genus Macrocneme Hiibner (Lepidoptera: Ctenuchidae) Robert E. Dietz IV University of California Press SYSTEMATICS AND BIOLOGY OF THE GENUS MACROCNEME HUBNER (LEPIDOPTERA: CTENUCHIDAE) lRH6-J9F-THy|Sopyrighted matBrial Systematics and Biology of the Genus Macrocneme Hubner (Lepidoptera: Ctenuchidae) Robert E. Dietz IV UNIVERSTTY OF CAUFORNIA PRESS Berkeley • Los Angeles • London Copyrightecl material UNIVERSTTY OF CALIFORNIA PUBUCATIONS IN ENTOMOLOGY Editorial Board: Howell V. Daly, John Doyen, Henry Hespcnhcidc, Robert S I ane, John Pinto, Jerry A. Powell, Arthur Shapiro, Philhp S. Ward, John Smiley Volume 113 Issue Date: April 1994 UNIVERSITY OP CALIFORNIA FItBSS BERKELEY AND LOS ANGELES, CALIFORNIA UNIVERSITY OFCALIFORNIA PRESS, LTD. LONDON, ENGLAND O 1994 BY THE REGENTS OP THE UNIVERSmr OF CALIFORNIA PRINTED IN THE UNITED STATES OF AMERICA Libraiy of Congress Cataloging-in-Publication Data Dietz. Robert E. (Robert Edwin). 1941- Systematics and biology of the genus Macrocneme Httbner (Lepidoptera: Clenuchidae) / Robert E. Dietz IV. p. cm. — (University of California publications in entomology; v. 113 Includes bibliographical references (p. ). ISBN 0-520-09780-7 (pbk. ; alk. paper) K Macrocneme. I. Htle. n. Series. QL561.C8D54 1994 595.78 1—dc20 93-49869 OP The paper used in this publication meets the miniinuin requirements of American National Standard for Information Sdenoes^Fnmanence of Paper for Printed Ubraty Materials. ANSI Z39^1984, S Copyrighted matBrial Contents Acknowledgments and Abbreviations, vii Abstract, x Intrndiictinn 1 Materials and Meitiod^v Z Nomenclature and l axonomic History 5 Genus Marrncncnip Hiihner 7 Diagnosis and Systematic Placement, 7 Description, 8 Variation, 10 Biology 11 Geographical Distribution 21 Key to Species 30 Svsiematic Treatnient 33 M. maja (F.), 33 M. adonis Druce. 34 M. c/irysitis (Guerin-M6neville), 37 M. iole Dnice. 40 A/. st't)iiviridis Druce, 43 Af. cabimensis Dyar, 45 M. oponiensb, n. sp., 47 M. immanis Hampson, 4<^ M. tarsispecca, n. sp.. 51 A/, luihroc cladon, n. sp., 52 M. lades (Cramer), 54 M. thvra Moschler. 60 M. tliyridia Hampson, 67 M. coendescem Dognin, 71 A/. Lim averdia. n. sp.. lU Af. awifera Hampson, 78 Af. durcata, n. sp.. 81 V vi Contents M. bodoquero, n. sp., 84 M. imbelliSs n. sp.. 86 A/, inelunopeza, n. sp.. 87 M. orichalceay n. sp., 88 M. zongonata, n. sp., 90 M. hestia, n. sp.. 92 M cupreipennis Walker, 93 M. cyanea (Butler), 95 M ferrea Butler, 97 M. leucostifjnia (Perty), 99 M. memcybe, n. sp.. 104 A/, monno, n. sp., 106 A/, pelotas, n. sp., 108 Species Transferred from Macrocneme 110 Checklist of Species 111 Appendix, 113 Literature Cited, 115 Maps and Fibres, 123 Acknowledgments and Abbreviations Much of the material on which this study is based was borrowed from or examined at museums in the United States. Liitin America, and Europe. I thank the curators who assisted me in arranging loans, answering questions, and/or providing facilities for the study of types and specimens in their charge. Listed below are the institutioiis and individuals who cooperated in this study, along with the abbreviation used for each throughout the text: AMNH American Museum of Natural History. New Yorlc. (F.H. Rindge) BMNH British Museum (Natural History), London. (A, Watson; in Glasgow, R.A. Crowson) CAS California Academy of Sciences, San Francisco. (PJH. Amaud) CM Carnegie Museum, Pittsburgh, PA. (H.K. Clench) CNC Canadian National Collection. Ottawa. (I'.Ci. Muiiroe) CU Cornell University, Ithaca, New York. (L.L. Pechuman) HM Zoolo^sches Museum, Universitit Hamburg, Germany. (H. Weidner) IML Fundaci<3n Miguel Lillo, Tucumto, Argentina. (A. Willink) LACM Los Angeles County Museum of Natural History, Los Angeles, California. (J. P. Donahue) MCZ Museum ot Comparative Zoology, Harvard Lniversily, Cambridge, Massachusetts. (J.M. Bums) MNHU Museum fibr Naturkunde der Humboldt-Umversitat, Berlia (HJ. Hannemann) MSU Michigan State University, Liinsing. (R.L. Fischer) NMB Naturhistorisches Museum, Bern, Switzerland. (H.D. Volkart) NMG Musdum d'Histoire Naturelle, Geneva. (C. Besuchet) NRS Naturhistoriska Riksmuseet. Stockholm. (B. Gustafsson) OX Oxford University Museum, Oxford. Fngland. (E. Taylor) PM Museum National d'Histoire Nalurcllc, Pan.^. (P. Victtc) RML Rijksmuseum van NaiuuiUjkc Hislonc, Leiden, the Netherlands. (R. de Jong) SMM Zoologische Sammlung des Bayerischen Staates, MunidL (W. FOrster) UCB University of California, Berkeley. (J.A. Powell) UCD University of California, Davis. (R.O. Schuster) vu Copyrighted matsrial Acknowledgments UCV Universidad Central de Venezuela, Maracay. (F. Fern^dez-Yepez) UFP Unhrersidade Federal do Parani, Ouitiba, Brazil (O.H.H. Mieike) ULP Universidad Nadonal de La Plata, Argentina. (L. De Santis) UP? University of Pennsylvania, Philadelphia. (D.H. Janzen) USNM United States National Museum [National Museum of Natural HistoryJ, Smithsonian institution, Washington, D.C. (E.L. Todd) USP Museu de Zoologia, Universidade de S8o Faulo, Bra^. ( N. Bemardi) VM Natttrtustorisches Museum, Vienna. (F. Ka^) ZIH Martin Luther Universitftt, Halle, Germany. (HQsing) Without the assistance and encouragement of numerous friends and colleagues during the long course of this study, this project might never have been completed. I owe special thanks to Jerry Powell, majur advisor, tutor, and friend, for his patienee and unstinting assistance in seeing this study through to completion. His guidance was responsible for numerous improvements in the content and presentation of this paper. Robert OmdufiF and Evert Schlinger, as additional members of the dissertation committee, reviewed the manuscript Hieir suggestions and criticisms are gratefiilfy aduiowledged. W. Donald Duckworth and [the lalej Harold Moore gave me valuable experience in the tropics by including me as an assistant on their collecting expeditions. This project was begun under their tutelage, and their unfailing support and interest in my progress will always be remembered and appreciated. Allan Watson was especially helpfid in returning types to me for further study and in answering those inevitable questions that arise in a protracted project where early details have a way of fading from memory. Frederick Rindge kindly reexamined the typt of spitUvatm Fleming for me. [The late] Francisco Femtodez \€peiz helped arrange for my visit to Maracay in 1974-75. His innumerable kindnesses and assistance during that year are warmly remembered. Michael Adams. George Bernard, and Michael Murtaugh suttered the vagaries and frustrations of collecting in the Tropics with me. Their forbearance, humor, and good company in the face of plagues of swarming insects made all bearable. Among those who ccmtributed specimens to this study from their own collecting efforts or searched for material on my behalf are: Yves Barbotfn (Cayenne, French Guiana); Andre Blanchard (Houston, Texas); John Chcmsak (Berkeley); Rosser Garrison (San Juan, Puerto Rico); George Godfrey (Champaign, Illinois); John Hafemik (San Francisco); Daniel Janzen (Riiladelphia); Gerardo Lamas (Lima, Peru); [the late] Harold Moore (Ithaca, New York); Tom Pliske (Coral Gables, Florida); Jerr\ Powell (Berkeley); Don Viers (Pullman, Washington); Philip Ward (Davis, California). Dale Habeck provided inldrmaiion about rearing ctenuchids, and Norman Leppla helped to reassess the artificial diet Their suggestions were us^ul in improving the rearing results. Special thanks are due the following for their assistance during various phases of the study: friends Phillip Guins and Alden Kamikawa helped with labeling: Richard Clarke collated locality data, helped prepare maps, and proofread various versions Copyrighted material Acknowledgments iz of the manuscript: Hinrich Seeba and Martha Bell Dietz translated articles from German; John DeBcncdictis took the SEM photographs; Dean and Chalmers Luckhart took the photographs of the adults; Karen Bailey helped with the typing; Maiy Anne Qarke assisted me in converting hand-drawn griqphs to computer- generated ones. To my mother and late father. Ann and Robert Dietz, I owe special thanks for their generous support and enc()ura<^ement over the course of this study. Their unflagging taith and confidence in me were invaluable morale boosters. This study was aided in part by the following grants and awards: National Science Foundation Grant 6813X (to Jerry A. Powell, 1969-70); National Institutes of Health Training Grant #AI-218, UCB; Chancellor's Patent Fund Award. UCO: Predoctoral Fellowship Award, Office of Academic Studies. Smithsonian Institution; Organization of American Slates Research Fellowship #PRA-41437. Final publication costs were defrayed, in part, by a generous gift from Ann Espe Dietz. Note; Present address of author; Dr. Robert E. Dietz IV, 6939 Edith Boulevard, NE, Albuquerque, New Mexico 87113. Copyrighted matBrial Abstract Thirty species considered monoptiyletic and congeneric with Macrocneme maja (F.) are characterized, illustrated, and discussed. Fourteen new species are described from South America: ancaverdia (Peru); bestia (Brazil); bodoquero (Colombia); durcata (Venezuela); habroceladon (Bolivia); imbellLs (Peru); megacybe (Brazil); melanopeza (Peru); monno (Brazil); oponiensis (Colombia); orichalcea (Brazil); pelotas (Brazil); tarsispecca (BoVw hi); zongonata (Brazil). Three names are removed from synonymy, one subspecific name is reinstated as a valid species, and nine names are iwwly placed In synoaynqr. Three