Marine Insects Edited by LannaCheng Scripps Institution of Oceanography, University of California, La Jolla, Calif. 92093, U.S.A. NORTH-HOLLANDPUBLISHINGCOMPANAY, AMSTERDAM- OXFORD AMERICANELSEVIERPUBLISHINGCOMPANY , NEWYORK © North-Holland Publishing Company - 1976 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise,without the prior permission of the copyright owner. North-Holland ISBN: 0 7204 0581 5 American Elsevier ISBN: 0444 11213 8 PUBLISHERS: NORTH-HOLLAND PUBLISHING COMPANY - AMSTERDAM NORTH-HOLLAND PUBLISHING COMPANY LTD. - OXFORD SOLEDISTRIBUTORSFORTHEU.S.A.ANDCANADA: AMERICAN ELSEVIER PUBLISHING COMPANY, INC . 52 VANDERBILT AVENUE, NEW YORK, N.Y. 10017 Library of Congress Cataloging in Publication Data Main entry under title: Marine insects. Includes indexes. 1. Insects, Marine. I. Cheng, Lanna. QL463.M25 595.700902 76-17123 ISBN 0-444-11213-8 Preface In a book of this kind, it would be difficult to achieve a uniform treatment for each of the groups of insects discussed. The contents of each chapter generally reflect the special interests of the contributors. Some have presented a detailed taxonomic review of the families concerned; some have referred the readers to standard taxonomic works, in view of the breadth and complexity of the subject concerned, and have concentrated on ecological or physiological aspects; others have chosen to review insects of a specific set of habitats. Nevertheless, each has presented a general picture of the group of insects under discussion, their nature, ecology, life histories, special adaptations to marine environments, and a comprehensive review of the literature. It is difficult to draw a line between the sea and dry land, or between marine and fresh waters. The word ‘marine’ is broadly interpreted here as referring to any more or less saline waters, and a ‘marine insect’ as one that spends at least part of its life in association with the marine environment. Chapters in the first part of this book are predominantly ecological in the broad sense, while the rest of the chapters deal with major groups of insects found in marine environments. It has been impossible to cover all the orders or families of insects with marine representatives. There is no chapter here dealing with Hymenoptera, which include several genera of tiny wasps parasitic on marine Canaceidae, etc., as well as the more familiar ants which help to clear up crumbs and debris on the beaches. There are no chapters dealing with the Dermaptera, which include a marine earwig, Anisolabis littorea (White), known so far only from New Zealand; or the Homoptera, though some discussions of saltmarsh aphids can be found in Ch. 2; or the Neuroptera, although the larvae of at least two species of Sisyridae, Climacia areolaris (Hagen) and Sisyra vicaria (Walker), are quite common in two brackish water sponges (Vincent Resh, personal communication). Nor are there chapters on the Orthoptera or the Lepidoptera. Several families of Diptera with marine representatives, notably Dolichopodidae, Canaceidae, and [ V] VI Tipulidae, are discussed only with respect to their respiratory adaptations (see Ch. 3). A number of topics of general interest have been excluded because of insufficient knowledge. In particular, we might mention the evolutionary and genetic aspects of marine insect distributions, which present challenging problems relating, for instance, to dispersal to new territories, gene exchange from ocean to ocean across or around continental land masses, and patterns of gene flow in coastal species whose territories are virtually linear or one-dimensional - several thousand kilometres long yet only a few metres wide. (However, some genetic studies on Coelopa frigida (Fabricius) are given in Ch. 16.) We hope such omissions will not be regarded as demerits of this book, but rather as incentives to others to study some of the less familiar kinds of marine insects. We have considered in this volume not only true insects, including the Collembola (Ch. 7), and insect parasites of marine birds and mammals (Ch. 4), but also other kinds of intertidal air-breathing arthropods (Ch. 6), notably spiders, scorpions, mites, centipedes and millipedes, which live and feed with, or even on, the insects of marine habitats. We have also included a chapter on migratory and other insects at sea (Ch. 5), even if they are blown seaward only sporadically and probably involuntarily! Clearly there is vast scope for studies on marine insects at present. List of contributors Andersen, N. Møller Universitetets Zoologiske Museum, Universitetsparken 16, Kobenhavn, Den- mark Axtell, Richard C. Department of Entomology, North Carolina State University, Raleigh, N.C. 26707, U.S.A. Bowden, John Rothamsted Experimental Station, Harpenden, Herts, England Brown, Wynne L. Department of General Biology, University of Arizona, Tucson, Ariz. 85721, U.S.A. Cheng, Lanna Scripps Institution of Oceanography, University of California at San Diego, La Jolla, Calif. 92093, U.S.A. Dobson, Terry Department of Genetics, University of Aarhus, Aarhus 8999, Denmark Doyen, John T. Department of Entomology, University of California, Berkeley, Calif. 94720, U.S.A. Foster, William D. Department of Zoology, University of Cambridge, Downing Street, Cambridge, England Hashimoto, Hiroshi General Education Department, Shizuoka University, Shizuoka, Japan Hinton, Howard E. Department of Zoology, Bristol University, Bristol, England Johnson, C.G. Rothamsted Experimental Station, Harpenden, Herts, England Joosse, Els N.G. Biologisch Laboratorium, Vrije Universiteit, Amsterdam, The Netherlands [VII] VIII Leader, John Department of Physiology, Otago Medical School, P.O.Box 913, Dunedin, New Zealand Legner, E.F. Department of Entomology, University of California, Riverside, Calif. 92502, U.S.A. Linley, John R. Florida Medical Entomology Laboratory, P.O. Box 520, Vero Beach, Fla. 32960, U.S.A. Moore, Ian Department of Entomology, University of California, Riverside, Calif. 92502, U.S.A. Murray, M.D. Division of Animal Health, McMaster Laboratory, CSIRO, Glebe, N.S.W. 2037, Aus- tralia O’Meara, George F. Florida Medical Entomology Laboratory, P.O.Box 520, Vero Beach, Fla. 32960, U.S.A. Polhemus, John T. 3115 South York, Englewood, Colo. 80110, U.S.A. Roth, Vincent D. Southwestern Research Station, American Museum of Natural History, Portal, Ariz. 85632, U.S.A. Scudder, Geoffrey G.E. Department of Zoology, University of British Columbia, Vancouver 8, B.C., Canada Simpson, Karl W. Division of Laboratories and Research, State of New York Department of Health, New Scotland Avenue, Albany, N.Y. 12201, U.S.A. Treherne, John E. Department of Zoology, University of Cambridge, Cambridge, England Acknowledgements No multi-authored book could be compiled without cooperation between the contributors and the editor. I would like to thank the contributors not only for the time and effort they devoted to their chapters, but also for generously considering and accepting many suggestions from their colleagues as well as from the editor. To all those who have helped in one way or another during the editorial process of this book, I offer my sincere thanks; in particular to Donna DeCamp, Virginia L. DeCamp and Vickie Gillespie for their skillful transformation of scribbled and cut-and-pasted pages into flawless typescripts; to Chi-ling Liu and Gayle M. Kidder for assistance in compiling the indexes; and to the following for all kinds of helpful suggestions and unpublished observations and comments: Dru Binney, Martin C. Birch, Linda Birch, Ted H. Bullock, Eric B. Edney, K.C. Emerson, John D. Isaacs, B.C. Nelson, Dietrich Neumann, Lester J. Newman, William A. Newman, Vincent Resh, Richard A. Schwartzlose, Eric Shulenberger, Gerry L. Wick and, of course, Ralph A. Lewin. The preparation of this book was partly supported by the National Sea Grant Program NOAA, Department of Commerce, under grant number 04-5-158-20 to the Scripps Institution of Oceanography. I should also like to thank the following for permission to reproduce figures: 4.3, Pacific Coast Entomological Society; 7.14 & 7.16, C.N.R.S., Paris; 7.4–7.13, Akademische Verlagsgesellschaft Geest & Portig, Leipzig; 7.28, 7.29, Faun.-Okol.¨ mitt., Kiel.; 9.1, 9.2, 9.6, 9.7, 9.9, 9.10, University of California Press; 9.5 & 9.11, Bishop Museum Press; 9.8, Smithsonian Institution; 9.13, Entomological Society of America; 9.3, Royal Entomological Society, London. [IX] Contents For a detailed list of contents the reader is referred to the opening page of each chapter Preface V List of contributors VII Acknowledgements IX Contents Chapter 1 Insects in marine environments, by Lanna Cheng 1 Topics of general interest Chapter 2 Insects of marine saltmarshes: problems and adaptations, by 5 William A. Foster and John E. Treherne Chapter 3 Respiratory adaptations of marine insects, by Howard E. 43 Hinton Chapter 4 Insect parasites of marine birds and mammals, by M.D. 79 Murray Chapter 5 Migrating and other terrestrial insects at sea, by John 97 Bowden and C.G. Johnson Chapter 6 Other intertidal air-breathing arthropods, by Vincent D. Roth 119 and Wynne L. Brown Reviews of major groups of marine insects Chapter 7 Littoral apterygotes (Collembola and Thysanura), by Els 151 N.G. Joosse XI XII Chapter 8 Water-striders (Hemiptera: Gerridae, Veliidae, etc.) by Nils 187 Møller Anderson and John T. Polhemus Chapter 9 Shore bugs (Hemiptera: