Social Insects. Social Insects V2

496 Pages·1981·11.004 MB·English

by  Henry Hermani (Eds.)

CONTRIBUTORS
Murray S. Blum
Ruth E. Buskirk
Jean Deligne
George C. Eickwort
Henry R. Hermann
Andre Quennedey
Roy R. Snelling
Social
Insects
Volume II
Edited by HENRY R. HERMANN
Department of Entomology
University of Georgia
Athens, Georgia
ACADEMIC PRESS
A Subsidiary of Harcourt Brace Jovanovich, Publishers
New York London Toronto Sydney San Francisco 1981
COPYRIGHT © 1981, 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 Da
Main entry under title:
Social insects.
Includes bibliographies and index.
1. Insect societies. I. Hermann, Henry R.
QL496.S6 595.7«052A 78-4871
ISBN 0-12-342202-7 (v. 2)
PRINTED IN THE UNITED STATES OF AMERICA
81 82 83 84 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.
Murray S. Blum (1, 77), Department of Entomology, University of
Georgia, Athens, Georgia 30602
Ruth E. Buskirk (281), University of Texas Marine Science Institute,
Galveston, Texas 77550
Jean Deligne (1), Laboratoire de Biologie Animale et Cellulaire, Universite
Libre de Bruxelles, Belgique, Belgium
George C. Eickwort (199), Department of Entomology, Cornell University,
Ithaca, New York 14853
Henry R. Hermann (77), Department of Entomology, University of
Georgia, Athens, Georgia 30602
Andre Quennedey (1), Laboratoire de Zoologie, Universite de Dijon, 21000
Dijon, France
Roy R. Snelling (369), Los Angeles County Museum of Natural History,
Los Angeles, California 90007
Preface
Just prior to and at the time of the printing of Volume I of "Social In-
sects," the field of insect sociality was undergoing a definite change. Prior
to that time many of the earlier reports as reviewed by Wilson (1971) were
of an investigative nature.* Current research of note immediately following
Wilson was largely theoretical, as indicated by the reports of Alexander
(1974), Eberhard (1975), Hamilton (1972), Lin and Michener (1972),
Orlove (1974), and Trivers and Hare (1976). Volume I of "Social Insects"
reviewed these publications and many of the reports that offered founda-
tion for such theory. Yet, most investigators of insect sociality realized at
that time that additional foundation was needed in basic research and that
we really knew very little about the behavior of eusocial insect species.
With Volume II we are standing on the threshold of a new period in the
study of insect sociality, a period of intensive research in an attempt to fill
the gaps in our knowledge of this special group of arthropods, a step
necessary for further speculation. Possibly some of this research will shed
new light on subjects such as altruism, haplodiploidy, the evolution of in-
sect sociality, kin selection, social parasitism, polygyny, and parental
manipulation.
Volume II represents a continuation of Volume I, which reviews known
concepts related to insect sociality. At the same time, Volume II treats the
systematics of eusocial hymenoptera and delves into the closely associated
realms of presocial insects and social non-insectan arthropods, setting the
stage for Volumes III and IV, which deal with the biology of eusocial
species. A comprehensive review of termite taxonomy may be found in
y Krishna and Weesner (1970, Vol. II, pp. 127-152).
With the completion of Volume II, the reader has enough review mate-
* Reference to the reports listed above may be found following Chapter 1 of Social Insects,
Volume I, Academic Press, 1979.
xi
Xll Preface
rial on the various subjects associated with insect sociality to piece together
a general description of what social insects do and how they do it. A com-
prehension of these concepts will facilitate an understanding of the biology
of eusocial Insecta as they are portrayed in Volumes III and IV, and it will
no doubt raise new questions that can be answered at a later date.
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 III (in preparation)
1. The Social Insects' Bestiary, D. H. Kistner
2. Behavior and Ecology of Bumble Bees, D. H. Morse
3. Honey Bees, A. Dietz
4. Stingless Bees, S. F. Sakagami
Volume IV (in preparation)
1. Social Wasps, R. D. Ackre
2. Ants: Foraging, Nesting, Broad Behavior, and Polytheism, John H. Sudd
3. Army Ants, W. H. Gotwald
4. Fungus Ant, N. Weber
Xlll
1
The Enemies and Defense
Mechanisms of Termites
JEAN DELIGNE, ANDRE QUENNEDEY,
AND MURRAY S. BLUM
I. Introduction 2
II. Termite Enemies 3
A. Invertebrata 3
B. Vertebrata 7
C. Conclusions 14
III. Defense by the Nest 14
IV. Defense by Larvae, Nymphs, and Reproductives 17
V. Defense by Workers 18
VI. Defense by Soldiers 19
A. Phragmotic Soldiers 20
B. Mandibular Weapons 21
C. Salivary Weapons 26
D. Frontal Weapons 29
E. Labral Weapons 41
F. Chemistry of Secretions 42
G. General Evolution of Weapons 58
VII. Integrated Defense 60
A. Relative Numbers of Soldiers 60
B. Polyethism Within the Soldier Caste 63
C. Replacement of Individuals 64
D. Coordination of Different Behaviors 64
VIII. Summary and Conclusions 66
References 67
SOCIAL INSECTS, VOL. II
Copyright © 1981 by Academic Press, Inc.
All rights of reproduction in any form reserved
ISBN 0-12-342202-7
2 J. Deligne, A. Quennedey, and M. S. Blum
I. INTRODUCTION
The defense mechanisms that a species uses may be defined as the
strategy—incorporating morphological, physiological, and behavioral
adaptations—by which that species limits or eliminates the action of its
parasites, predators, and competitors.
This definition is certainly not the only one possible for defense
mechanisms. It is possible, for instance, by defining defense in a wider
sense, to include reactions against certain undesirable, inanimate objects,
and various unfavorable physical and chemical factors in the environment.
In certain cases, such inanimate external agents provoke the same sort of
reactions as a live enemy. However, a wide definition of this sort, including
reactions against both biotic and abiotic elements in the environment, runs
the risk of becoming too heterogeneous. On the other hand, it is possible to
limit the definition of defense to that which concerns certain enemies, and
in particular predators, in order to discuss a more limited and therefore
more coherent number of phenomena. It must, nevertheless, be pointed out
that the often imprecise divisions of the various categories of enemies make
this definition slightly artificial at times.
For this chapter, we have adopted the first definition of defense—all ac-
tivity directed against an enemy. So defined, the concept of defense has an
evident biological unity; it is an important factor in the relations between
individuals and between populations in a biotic community and is one of
the factors governing survival and success in a given ecosystem.
Defense thus does not consist of one well-defined type of behavior or
physiological change, but may be described as a functional, ecological con-
cept. In termites, which are a rather diverse systematic group with a well-
developed polymorphism, defense mechanisms may vary greatly from one
species to another, one caste or subcaste to another, and even from one
situation to another.
Studies of defense mechanisms are complicated by the fact that adapta-
tions concerning defense are rarely totally independent of other types of
behavior. Stuart (1967) has shown that defensive behavior may involve the
communication of alarm, the laying of a trail, or building behavior. In
such cases, defensive behavior against an intruder appears to share com-
mon motor elements with other types of behavior, for instance, nest
building and food recruitment.
Finally, Howse (1975) has pointed out that it is not always easy to
distinguish defense from attack and threat, particularly when competitors
of the same or a similar species are concerned. The difficulty is, however,
more formal than fundamental, and will not be discussed further.
Information relative to defense behavior in termites is, at least in part,