CONTRIBUTORS Alfred Dietz David H. Kistner Douglass H. Morse Shoichi F. Sakagami Social Insects Volume III Edited by HENRY R. HERMANN Department of Entomology University of Georgia Athens, Georgia 1982 ACADEMIC PRESS A Subsidiary of Harcourt Brace Jovanovich, Publishers New York London Paris San Diego San Francisco Sao Paulo Sydney Tokyo Toronto COPYRIGHT © 1982, BY ACADEMIC PRESS, INC. ALL RIGHTS RESERVED. NO PART OF THIS PUBLICATION MAY BE REPRODUCED OR TRANSMITTED IN ANY FORM OR BY ANY MEANS, ELECTRONIC OR MECHANICAL, INCLUDING PHOTOCOPY, RECORDING, OR ANY INFORMATION STORAGE AND RETRIEVAL SYSTEM, WITHOUT PERMISSION IN WRITING FROM THE PUBLISHER. ACADEMIC PRESS, INC. Ill Fifth Avenue, New York, New York 10003 United Kingdom Edition published by ACADEMIC PRESS, INC. (LONDON) LTD. 24/28 Oval Road, London NW1 7DX Library of Congress Cataloging in Publication Data Main entry under title: Social insects. Includes bibliographies and indexes. 1. Insect societies. I. Hermann, Henry R. QL496.S6 595.7'0524 78-4871 ISBN 0-12-342203-5 (v.3) AACR2 PRINTED IN THE UNITED STATES OF AMERICA 82 83 84 85 9 8 7 6 5 4 3 2 1 To My Wife, Lisa, and Son, Brad List of Contributors Numbers in parentheses indicate the pages on which the authors' contributions begin. Alfred Dietz (323), Department of Entomology, University of Georgia, Athens, Georgia 30602 David H. Kistner (1), Department of Biological Sciences, California State University, Chico, California 95929 Douglass H. Morse1 (245), Division of Biology and Medicine, Brown Uni- versity, Providence, Rhode Island 02912 Shoichi F. Sakagami (361), The Institute of Low Temperature Science, Hokkaido University, Sapporo 060, Japan 1 Present address: Department of Zoology, University of Uppsala, S-75122 Uppsala, Sweden Preface Chapter 1 of this volume represents the culmination of discussion on social insect phenomena in this four-volume treatise, and leads into subsequent chapters on the biology of the groups of eusocial Insecta themselves. As portrayed in Chapter 1, insect symbionts represent a very complex group of organisms of quite diverse habits. Knowledge of the number of symbiotic species and their unusual associations with eusocial insects is stag- gering—hence the very lengthy treatment. Nonetheless, new symbionts are still being recognized almost daily, and the relationships are increasingly fascinating. The remainder of this volume is dedicated to the Apidae, including the Apinae (Meliponini and Apini) and the Bombinae. As pointed out in Volume I, Chapter 3, by Carpenter and Hermann, the patterns of social behavior in the Apidae are considerably more diverse than in the Vespidae, as well as in the Formicidae. The bees obviously have developed eusociality independently many times, and eusociality in the bees has reached its climax in the Apidae. Since social behavior is found in other groups of bees as well, especially in some Halictidae and Anthophoridae, the reader is invited to peruse Volume II, Chapter 3, by Eickwort, to round out the picture of sociality in the bees in general. Henry R. Hermann Contents of Other Volumes Volume I Insect Sociality—An Introduction, H. R. Hermann Origin and Evolution of Insect Sociality: A Review of Modern Theory, C. K. Starr Antiquity of Sociality in Insects, F. M. Carpenter and H. R. Hermann Territoriality in Social Insects, Cesare Baroni Urbani Caste Differentiation and Division of Labor, M. V. Brian Genetics of Sociality, R. H. Crozier Larvae of the Social Hymenoptera, G. C. Wheeler and J. Wheeler Social and Evolutionary Significance of Social Insect Symbionts, David H. Kistner Volume II The Enemies and Defense Mechanisms of Termites, Jean Deligne, Andre Quennedey, and Murray S. Blum Defensive Mechanisms in the Social Hymenoptera, Henry R. Hermann and Murray S. Blum Presocial Insects, George C. Eickwort Sociality in the Arachnida, Ruth E. Buskirk Systematics of Social Hymenoptera, Roy R. Snelling Volume IV (in preparation) Social Wasps, Roger D. Akre Ants: Foraging, Nesting, Broad Behavior, and Polyethism, John H. Sudd Army Ants, W. H. Gotwald, Jr. Fungus Ants, Neal A. Weber Xlll 1 9 The Social Insects Bestiary DAVID H. KISTNER I. Introduction 2 A. Definition of Terms 2 B. Scope of the Chapter 3 II. Arthropods Other Than Insects 4 A. Mites 4 B. Spiders 10 C. Pseudoscorpions 13 D. Solfugida(Sun Spiders) 14 E. Diplopoda 15 F. Crustacea 16 III. Insects 19 A. Collembola or Springtails 19 B. Thysanura 22 C. Orthopteraor Crickets 29 D. Blattariaor Roaches 31 E. Hemiptera, Homoptera, and Thysanoptera 33 F. Psocoptera or Book Lice 39 G. Neuroptera 41 H. Coleoptera or Beetles 42 I. Strepsiptera 146 J. Hymenoptera 147 K. Lepidoptera 162 L. Diptera 177 IV. General Considerations 218 References 221 1 SOCIAL INSECTS, VOL. Ill Copyright (£) 1982 by Academic Press, Inc. All rights of reproduction in any form reserved. ISBN 0-12-342203-5 2 David H.Kistner I. INTRODUCTION A. Definition of Terms Symbionts of social insects are usually called myrmecophiles, ter- mitophiles, melittophiles, or sphecophiles depending upon whether they are found with ants, termites, bees, or wasps, respectively. The terms, as they are used in the literature, merely denote that an insect is usually found in associa- tion with its host. They do not imply anything about whether the relationship is facultative or obligatory or anything about the specificity. However, as more is known about the relationships of the arthropods involved, casual relationships can usually be excluded from the category and they then imply some sort of symbiosis. The origins of the terms are quite interesting. Myrmecophile is the oldest, first appearing in English literature in the late 1800s and transferring quite naturally into American English. Termitophile was used quite freely in the German literature as early as 1857 (Kraatz, 1857) but it did not appear in English literature until the American, Schwarz (1889), introduced it. Mellitophile and sphecophile were introduced into English by Wheeler (1928) and they have not found their way yet into standard English dictionaries. A myrmecophile (or other symbiont) is said to be integrated into the ant col- ony (or other colony) if it interacts with the ants so that the ants accept it into their society. This is in contrast to other myrmecophiles which may never achieve integration into the ant society but which may, nevertheless, live in the nest and exploit its natural resources. Such myrmecophiles are said to be nonintegrated. These definitions are greatly expanded and illustrated in Chapter 8 of Volume I of this series. Integration may be effected by the use of chemical cues or by tactile mimicry. These concepts are also expanded and illustrated in Volume I of this series. Tactile mimicry when used to effect social integration is called Wasmannian mimicry (Rettenmeyer, 1970). The meaning of Wasmannian mimicry was expanded by Kistner and Jacobson (1975) to include all mimicry of social releasers. Thus integration by the use of pheromones could also be a form of Wasmannian mimicry if the chemical mimicked a pheromone which functioned as a normal social releaser among members of the social insect col- ony. Such mimicry is still controversial among students of social insects, some of whom believe that tactile mimicry is the result of selection by predators out- side the social insect society. Social insects tend to be avoided by predators, since most have stings, can emit noxious substances, or have powerful man- dibles. Thus a symbiont might escape predation by its resemblance to undesirable prey. If this is true, then ant mimicry or termite mimicry would be a form of Batesian mimicry. Undoubtedly, elements of Batesian mimicry 1. The Social Insects'Bestiary 3 function in the New World army ant societies where some myrmecophiles such as Ecitophya and Ecitomorpha habitually accompany their Eciton hosts on raids. Where color changes throughout the range of Eciton burchelli Westwood, the color of the species of Ecitomorpha and Ecitophya varies ac- cordingly. The color of other myrmecophiles found with the same hosts which do not comingle on raids does not vary. Thus form and texture of Ecitophya and Ecitomorpha are probably aspects of Wasmannian mimicry, permitting acceptance by the ants, whereas color, which would be meaningless to the ants with their limited eyesight, would be a form of Batesian mimicry directed at outside predators such as the numerous ant birds who are attracted to the ant columns primarily to pick up insects which are flushed out by the ants (Willis andOniki, 1978). The foregoing types of phenomena are stressed in this chapter, however, any interesting aspects of social biology of the symbionts are reviewed even if they do not presently fit into any existing conceptual framework. It is only by keeping unexplained phenomena in the forefront that new concepts emerge. Absent from this discussion are concepts such as amical selection and sym- philic instincts which were elaborated by Wasmann (1920, and elsewhere). These concepts have been laid to rest many times, most thoroughly by K. Holldobler (1948), so that there is no need to resuscitate them here. The con- cept of Wasmannian mimicry does not need these erroneous props. B. Scope of the Chapter In Chapter 8 of Volume I of this treatise, the social and evolutionary significance of myrmecophiles and termitophiles was discussed. Data organized around such themes are necessarily selective and tend to give the im- pression that far fewer foreign insects inhabit the nests of social insects than are actually there. The purpose of this chapter is to review the social biology of all the groups of Arthropods found with the social insects. The mere presence of a foreign insect in a nest of social insects is a biological fact, but these myriads of facts are not reviewed unless something else is known about the in- teraction of the foreign insect with its hosts. There is a great need for a catalog of the social insect symbionts by host but this is not the format in which to do it. To my knowledge, the only previous attempts to do this were by Escherich (1909) and Donisthorpe (1927). Escherich's essay was limited to guests of ter- mites, while Donisthorpe's book was limited both to guests of ants and geographically limited to the United Kingdom. Wilson (1971) gave an adum- brated table in which he briefly reviewed some of the biology. All other monographs are limited to single taxonomic groups often found with far more limited hosts (Seevers, 1957, 1965). So many new discoveries have been