Handbuch der Zoologie Handbook of Zoology Band/Volume IV Arthropoda: Insecta Niels P. Kristensen (ed.) Lepidoptera, Moths and Butterflies Volume 1: Evolution, Systematics, and Biogeography Teilband/Part 35 Handbuch der Zoologìe Eine Naturgeschichte der Stämme des Tierreiches Handbook of Zoology A Natural History of the Phyla of the Animal Kingdom Gegründet von / Founded by Willy Kükenthal Fortgeführt von / Continued by M. Beier, M. Fischer, J.-G. Helmcke, D. Starck, H. Wermuth Band / Volume IV Arthropoda: Insecta Teilband / Part 35 Herausgeber / Editor Maximilian Fischer W DE G Walter de Gruyter · Berlin · New York 1999 Niels P. Kristensen (ed.) Lepidoptera, Moths and Butterflies Volume 1: Evolution, Systematics, and Biogeography W DE G Walter de Gruyter · Berlin · New York 1999 Autor und Herausgeber Schriftleiter und Herausgeber Author and Scientific Editor Managing and Scientific Editor Professor Dr. Niels P. Kristensen Hofrat Univ.-Doz. Zoological Museum Mag. Dr. Maximilian Fischer University of Copenhagen Direktor am Universitetsparken 15 Naturhistorischen Museum Wien DK-2100 Copenhagen Burgring 7 Denmark A-1014 Wien Austria Verlag / Publishers Walter de Gruyter GmbH & Co. Walter de Gruyter, Inc. Genthiner Straße 13 200 Saw Mill River Road D-10785 Berlin Hawthorne, N. Y. 10532 Germany USA Library of Congress - Cataloging-in-Publication Data Handbook of Zoology agr 23-1436 © Printed on acid-free paper which falls within the guide- Die Deutsche Bibliothek - Cataloging-in-Publication Data lines of the ANSI to ensure permanence and durability. Handbuch der Zoologie : eine Naturgeschichte der Stämme des Tierreiches / gegr. von Willy Kükenthal. Copyright © 1998 by Walter de Gruyter GmbH & Co., Fortgef. von M. Beier ... — Berlin ; New York : de Berlin. All rights reserved, including those of translation Gruyter into foreign languages. No part of this book may be Teilw. mit Paralleli.: Handbook of zoology reproduced in any form - by photoprint, microfilm or Bd. 4. Arthropoda: Insecta / Hrsg. Maximilian Fischer any other means nor transmitted nor translated into a Teilbd. 35. Lepidoptera, moths and butterflies machine language without written permission from the Vol. 1. Evolution, systematics, and biogeography / publisher. - Printed in Germany. Niels P. Kristensen (ed.). - 1998 ISBN 3-11-015704-7 Typesetting and Printing: Arthur Collignon GmbH, Ber- lin. - Binding: Lüderitz & Bauer GmbH, Berlin. Editor's preface Thirty years have elapsed since the publication ever, in the case of some chapters where the of the first issues in this second edition of the responsibility for different section has been Insecta section of the Handbuch der Zoologie! clearly divided between authors, this has been Handbook of Zoology. Not surprisingly there- explained. fore, editorial principles and standards are now As is usual in works of this kind an approxi- changing to ensure that the series can continue mate total length of the volumes was agreed to fulfill its intended role in the biological refer- from the outset. The subsequent page allocation ence literature. to individual chapters was evidently fraught with While treatments of the majority of the small problems. As far as the systematics chapters are and mediumsized insect groups conventionally concerned a balance was sought between criteria ranked as 'orders' have been issued already, only concerning species richness, diversity (there is a single treatment of a 'megadiverse' order has not necessarily a simple correlation of the two), been completed, viz., Willi Hennig's 1973 volume importance in a phylogenetic context, biological on the Dipera. By the 1980s it had become clear phenomena of general interest, and knowledge that accounts of the larger insect groups of the available. Unquestionably different traditions in breadth and depth required in the Handbuch/ the study of different major lineages, as well as Handbook series were far beyond the capacity of contrasting maturity of their classifications, con- tribute to any imbalance that may remain in the any single author or even of a small team of treatment. For instance, while all tortricoid authors. The present Lepidoptera volume, then, moths are included currently in just a single fam- is the first example of the genuine multi-author ily, a globally applicable tribal taxonomy is now treatment which will be adopted in forthcoming available for this lineage; the detailed treatment volumes on the large insect orders. Moreover, it down to this level is likely to be welcomed by is our aim that future volumes in the series focus- users. By contrast, the preliminary state of the sing on systematic entities will be published in higher classification of the much more species- English. rich family Geometridae largely explains the rel- Two Lepidoptera volumes are planned. The atively more brief treatment of this taxon. Since present volume on systematics, evolution and most authors probably considered alloted space biogeography will be followed by one on mor- inadequate for a proper treatment of their sub- phology and physiology. The latter is in an ad- jects), exesses were the rule rather than the ex- vanced stage of preparation. Plans for a third vol- ception! Some manuscripts were shortened by ume dealing with Lepidoptera ecology and behav- the editor. But variation between authors in ob- iour were abandoned after consultation with a serving space limitations remains another source number of potential editors: While ecological and of imbalance in chapter length. behavioural research on moths and butterflies has long been vigorous, there is greater motivation Differences between individual chapters in for reviewing findings emerging therefrom in style and presentation have been left unharmo- contexts other than one centred just on the order nized, as long as they are unlikely to create con- Lepidoptera. Needless to say, a certain body of fusion. For instance, the use of both Roman and information on ecology and behaviour is indeed Arabic numerals to designate abdominal segment being provided in relevant places in both volume numbers has been accepted. Similarly, examples 1 and 2. Moreover, volume 1 was broadened to of both English and American spelling will be accommodate a chapter on larval food relation- found. ships, a topic which certainly can be appropri- While works of this kind are intended primar- ately summarized in a Lepidoptera context, i.e., ily to provide state-of-the-art reviews, the prepa- within a large-scale evolutionary framework. ration of contributions for the present volume Cross references between chapters in this two- prompted some authors to undertake a substan- volume Handbook treatment of the Lepidoptera tial amount of original research. Hence the is eased by means of emboldened volume-and- content of some chapters is truly innovative. chapter numbers; thus 1-8 and 2—8 refer to Necessarily, subjective decisions had to be made chapter 8 in volume 1 and 2 respectively. concerning what could appropriately be included Name sequence of contributors to coauthored in the treatment published here. For instance, the chapters is alphabetical (except in the case of sizable original analyses of butterfly inter-rela- chapter 1—20, where this principle was aban- tionships upon which chapter 1 — 16 is largely doned at an express request from within the based were found to exceed the scope of a Hand- author team), hence it does not necessarily indi- book chapter, and they were accordingly pub- lished as a separate journal article (de Jong, R., cate who was the principal contributor (if the Vane-Wright, R. I. & Ackery, R R.: Ent. scand. workload was indeed unevenly shared). How- VI Editor's preface 27: 65-101, 1996). But it was decided that the As an editor I am greatly indebted to the team similarly innovative analysis of Gelechioid phy- of authors for their commitment, faith and pa- logeny, which forms the basis of 1—9, should be tience. I similarly thank the numerous specialists included, with the primary material printed as who provided reviews of individual chapters. Fi- appendices. nally I thank Dr Maximilian Fischer (former sole While some contributions were completed just editor, Insecta section of the Handbook series) before type-setting of the entire colume was initi- as well as PD Dr Mario Noyer-Weidner and Ms Christiane Bowinkelmann (Walter de Gruyter, ated, others had been completed considerably Biosciences Division) and Ms Ingeborg Klak earlier. However, most authors were able to take (Walter de Gruyter, Production Department) for the opportunity to make additions to their con- their collaboration on this enterprise. tributions in 1997, so the volume as a whole will appear reasonably updated by the time of its Copenhagen, April 1998 publication. Niels P. Kristensen Contents Editor's preface V 12. The Tortricoidea. Marianne Horak 199 Contributors IX 1. Historical Introduction. 13. The Smaller Microlepidopteran-Grade Niels P. Kristensen 1 Superfamilies. John S. Dugdale, Niels P. Kristensen, 2. Phylogeny and Palaeontology. Gaden S. Robinson & Malcolm J. Scoble 217 Niels P. Kristensen & Andrzej W. Skalski 1 14. The Pyraloidea. 3. Classification and Keys to Higher Taxa. Eugene Munroe & M. Alma Solis . . .. 233 David J. Carter & Niels P. Kristensen. . 27 15. The Axioidea and Calliduloidea. 4. The Non-Glossatan Moths. Joël Minet 257 Niels P. Kristensen 41 16. The Butterflies: Hedyloidea, 5. The Homoneurous Glossata. Hesperioidea and Papilionoidea. Niels P. Kristensen 51 Phillip R. Ackery, Rienk de Jong & 6. The Monotrysian Heteroneura. Richard I. Vane- Wright 263 Donald R. Davis 65 17. The Drepanoid/Geometroid 7. The Tineoidea and Gracillarioidea. Assemblage. Donald R. Davis & Gaden S. Robinson 91 Joël Minet & Malcolm J. Scoble 301 8. The Yponomeutoidea. 18. The Bombycoidea and their Relatives. John S. Dugdale, Niels P. Kristensen, Claude Lemaire & Joël Minet 321 Gaden S. Robinson & Malcolm J. Scoble 119 19. The Noctuoidea. 9. The Gelechioidea. Ian J. Kitching & John E. Rawlins . . . 355 Ronald W. Hodges 131 20. Evolution of Larval Food Preferences 10. The Zygaenoidea. in Lepidoptera. Marc E. Epstein, Henk Geertsema, Clas Jerry Powell, Charles Mitter & Brian M. Naumann & Gerhard M. Tarmann 159 Farrell 403 11. The Cossoid/Sesioid Assemblage. 21. Biogeography of the Lepidoptera. Edward D. Edwards, Patricia Gentili, Jeremy D. Holloway <6 Ebbe S. Nielsen 423 Niels P. Kristensen, Marianne Horak & Ebbe S. Nielsen 181 Index of Scientific Lepidoptera Names . . . 463
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