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Biology and Systematics of Grey a Busck and Tetragma, New Genus (Lepidoptera: Prodoxidae)

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Biology and Systematics of Grey a Busck and Tetragma, New Genus (Lepidoptera: Prodoxidae) Biology and Systematics of Grey a Busck and Tetragma, new genus (Lepidoptera: Prodoxidae) DONALD R. DAVIS, OLLE PELLMYR, and JOHN N. THOMPSON I SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 524 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 Astrophysics 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. Robert McC. Adams Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 524 Biology and Systematics of Greya Busck and Tetragma, new genus (Lepidoptera: Prodoxidae) Donald R. Davis, Olle Pellmyr, and John N. Thompson SMITHSONIAN INSTITUTION PRESS Washington, D.C. 1992 ABSTRACT Davis, Donald R., Olle Pellmyr, and John N. Thompson. Biology and Systematics of Greya Busck and Tetragma, new genus (Lepidoptera: Prodoxidae). Smithsonian Contributions to Zoology, number 524, 88 pages, 375 figures, 7 maps, 1992.—Greya is a genus of particular biological interest, in that it is among the genera closest related to the yucca moths, which are widely quoted in discussions of coevolution. Both Greya and the new genus Tetragma share some morphological and behavioral traits with the yucca moths. In this paper, the general morphology, classification, distribution, and biology of the western North American genera Greya and Tetragma new genus, are reviewed, and a phytogeny is proposed. Sixteen species of Greya are recognized, including seven new species: G. mitellae, G. obscura, G. enchrysa, G. variabilis, G. pectinifera, G. suffusca, and G. powelli. Greya piperella is resurrected as a valid species. The new genus Tetragma is described, with the single, new species T. gei. Keys are provided for all species, and diagnostic characters of all taxa are illustrated by line drawings and photographs. 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 Davis Donald R. (Donald Ray) Biology and systematics of Greya Busck and Tetragma, new genus (Lepidoptera: Prodoxidae) / Donald R. Davis, Olle Pellmyr, and John N. Thompson. p. cm.-(Smithsonian contributions to zoology ; no. 524) Includes bibliographic references. 1. Greya. 2. Tetragma. I. Pellmyr, Olle. II. Thompson, John N. III. Title. IV. Series. QL1.S54 no. 524 [QL561.P57] 591 s-dc20 [595.781!] 91-28433 ® 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 Abbreviations 1 Acknowledgments 2 Life History 2 Host Plants 2 Adult 3 Flight Period 3 Activity Pattern and Foraging 4 Courtship 4 Oviposition 4 Egg 5 Larva 5 Pupa 10 Natural Enemies 10 Larva 10 Adult 10 Egg and Pupa 10 Geographic Distribution 10 Phylogeny 13 Interrelationships between Greya and Tetragma Species 13 Greya and Tetragma in Relation to Other Prodoxid Genera 16 Key to the Genera of Prodoxidae 18 Tetragma Davis and Pellmyr, new genus 18 Tetragma gei Davis and Pellmyr, new species 19 Genus Greya Busck 24 Key to the Species of Greya 27 Greya punctiferella (Walsingham) 28 Greya piperella (Busck) 33 Greya mitellae Davis and Pellmyr, new species 36 Greya obscura Davis and Pellmyr, new species 39 Greya obscuromaculata (Braun) 40 Greya sparsipunctella (Walsingham) 43 Greya politella (Walsingham) 43 Greya enchrysa Davis and Pellmyr, new species 51 Greya variabilis Davis and Pellmyr, new species 56 Greya pectinifera Davis and Pellmyr, new species 58 Greya variata (Braun) 59 Greya subalba Braun 63 Greya solenobiella (Walsingham) 64 Greya suffusca Davis and Pellmyr, new species 66 Greya reticulata (Riley) 68 Greya powelli Davis and Pellmyr, new species 70 Literature Cited 72 Figures 251-375 74 Biology and Systematics of Grey a Busck and Tetragma new genus (Lepidoptera: Prodoxidae) Donald R. Davis, Olle Pellmyr, and John N. Thompson Introduction of eight species. Eight new taxa are described in this report. The yucca-pollinating moths, Tegeticula and Parategeticula, With the exception of a recorded single flower visit (Taylor, arc probably the best known group of non-ditrysian moths. 1965) nothing was known about the early instars or the biology Their status arises from the obligate pollination mutualism that of any member until oviposition in G. subalba was studied in has evolved between most species of these genera and their the 1980's (Thompson, 1986,1987). The onset of a large-scale larval host plants (Riley, 1892a; Powell and Mackie, 1966; study of the phylogeny and biology by two of the authors in Davis, 1967; Powell, 1984). Female yucca moths collect pollen 1987 has led to rapid growth of our knowledge in this respect, with specialized mouthparts prior to oviposition, then pollinate and much more material has also become available for flowers while visiting them for oviposition. Thus, they insure a systematic studies. Although more information is likely to food source for their progeny, which feed on seeds or become available to resolve certain issues (e.g., generic status vegetative parts of the fruit. of two species currently unclear), we have chosen to publish at In contrast to the yucca moths and associated Agavaceae- this point to make the new names available. Although a feeders, their probable sister group Greya has received far less member a different subfamily than Greya, the new genus attention. No known morphological synapomorphy delimits Tetragma likewise is included to provide a name and to present the Agavaceae-feeding genera from Greya, and it is likely that important biological information. The systematic parts of this an understanding of the phylogeny, systematics, host associa- revision have been prepared by Davis and Pellmyr, while the tions, and interactions of this genus would shed light on the biological information was brought together by Pellmyr and evolutionary origin of the yucca-yucca moth mutualism. Thompson. In the biological sections, information without Although the first species now placed within Greya was specific references to other works is based on unpublished data described in 1880 (Walsingham, 1880), the genus was not collected by the authors. recognized until 1903 (Busck, 1903). It has never been subject ABBREVIATIONS.—Abbreviations for the museums, institu- to a modem revision, and Davis (1983) brought together tions, and private collections where the material examined is several taxa previously placed within other genera to list a total deposited. Donald R. Davis, Department of Entomology, National Museum of AMNH American Museum of Natural History, New Natural History, Smithsonian Institution, Washington, D.C. 20560. Olle Pellmyr, Department of Biological Sciences, University of York, NY, USA Cincinnati, Cincinnati, Ohio 45221 and Departments of Botany and ANSP Academy of Natural Sciences in Phila- Zoology, Washington State University, Pullman, Washington 99164. delphia, Philadelphia, PA, USA John N. Thompson, Departments of Botany and Zoology, Washington State University, Pullman, Washington 99164. BMNH Collections of the former British Museum Review Chairman: Ronald J. McGinley, National Museum of Natural (Natural History), now renamed as the History, Smithsonian Institution. Natural History Museum, London, United Reviewers: Ebbe S. Nielson, Division of Entomology, CSIRO, Kingdom Canberra City, Australia; Jerry A. Powell, University of California, Berkeley, California; and David L. Wagner, University of Connecti- CAS California Academy of Sciences, San Fran- cut, Storrs, Connecticut. cisco, CA, USA SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY CNC Canadian National Collection, Ottawa, ON, and individuals for loans, gifts, or access to inspection of Canada material in their charge or possession: Paul H. Arnaud Jr., CAS, CU Cornell
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