Table Of ContentThe Genetics
of Social Evolution
Westview Studies in Insect Biology
Michael D. Breed, Series Editor
The Genetics of Social Evolution, edited by Michael D. Breed and
Robert E. Page, Jr.
The Entomology of Indigenous and Naturalized Systems in Agriculture,
edited by Marvin K. Harris and Charlie E. Rogers
lnterindividual Behavioral Variability in Social Insects, edited by Robert
L. Jeanne
Fire Ants and Leaf-Cutting Ants: Biology and Management, edited by
Clifford S. Lofgren and Robert K. Vander Meer
Management of Pests and Pesticides: Farmers' Perceptions and Prac-
tices, edited by Joyce Tait and Banpot Napompeth
Integrated Pest Management on Rangeland: A Shortgrass Prairie Per-
spective, edited by John L. Capinera
The "African" Honey Bee, edited by David J.C. Fletcher and Michael
D. Breed (forthcoming)
The Genetics
of Social Evolution
EDITED BY
Michael D. Breed
and Robert E. Page, Jr.
First published 1989 by Westview Press
Published 2019 by Routledge
52 Vanderbilt Avenue, New York, NY 10017
2 Park Square, Milton Park, Abingdon, Oxon OX14 4RN
Routledge is an imprint of the Taylor & Francis Group, an informa business
Copyright ©1989 Taylor & Francis
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Product or corporate names may be trademarks or registered trademarks, and are used only
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Library of Congress Cataloging-in-Publication Data
The Genetics of social evolution / edited by Michael D. Breed and
Robert E. Page, Jr.
p. cm.—(Westview studies in insect biology)
1. Insects—Genetics. 2. Insects—Evolution. 3. Insects—
Behavior. 4. Insect societies. 5. Social behavior in animals.
6. Behavior evolution. I. Breed, Michael D. II. Page, Robert E., Jr.
III. Series.
QL493.G46 1989
595.7'0524—dc19 88-12229
CIP
ISBN 13: 978-0-367-29248-5 (hbk)
CONTENTS
Preface vii
1. Introduction: Theory and Empirical Results in
Sociogenetics, Michael D. Breed 1
2. Genetic Structure, Division of Labor, and the
Evolution of Insect Societies, Robert E. Page, Jr.,
Gene E. Robinson, Nicholas W. Calderone, and
Walter C. Rothenbuhler 15
3. The Genetics of Colony-Level Selection,
Robin E. Owen 31
4. Genetic Basis for Division of Labor in an Insect
Society, Gene E. Robinson and Robert E. Page, Jr. 61
5. Ecological Determinants of Social Evolution,
Joan E. Strassmann and David C. Que/fer 81
6. Measuring Inclusive Fitness in Social Wasps,
David C. Que/fer and Joan E. Strassmann 103
7. Genetic and Social Changes Associated with Ant
Speciation, Philip S. Ward 123
8. Reproductive and Social Structure in Polygynous
Fire Ant Colonies, Kenneth G. Ross 149
.
9. Incompatibility Between Colonies of Acacia Ants:
Genetic Models and Experimental Results,
Alex Mintzer 163
10. Evolutionary Genetics of a Primitively Eusocial
Halictine Bee, Dialictus zephyrus,
Penelope F. Kukuk 183
Index 203
PREFACE
Our primary goal in organizing this book was to initiate a
synthesis of thought on how genetics structures the behavior of
individual animals that live within complex social systems. To do
this we have brought together leading theorists and empiricists
who apply genetics to the study of eusocial insect evolution.
The eusocial insects, with differentiated worker and
reproductive castes, pose particularly important problems in
evolutionary genetics. The most apparent of these is the question
of how extreme altruism (sterile worker castes} might have
evolved. Twenty-five years later, the elegant genetic kin selection
arguments of Hamilton still lack conclusive tests. Strassmann and
Queller (Chapter 5} and Queller and Strassmann (Chapter 6}
consider the current status of kin selection explanations. In doing
so they focus on the relative importance of ecology and genetics in
understanding the evolution and maintenance of extreme altruism.
Polistes wasps are the source of their empirical data. Kukuk
(Chapter 1 0} presents genetic data from populations of the
primitively eusocial sweat bee, Lasioglossum zephyrum (also
refered to as Dialictus zephyrus}, and discusses the application of
these data to kin selection hypotheses. Kin recognition may be a
prerequisite or a logical consequence of kin selection; in Chapter 9,
Mintzer presents genetic models for the kin recognition system of
Acacia ants.
Selection may act on a social group as a unit. Owen
(Chapter 3} considers the conditions under which the colony can
be considered a unit of selection and discusses the genetic
consequences of colony-level selection.
Empirical studies have found considerable genetic
variation within colonies of many species, despite the genetic
cohesion of eusocial groups that is normally assumed. Page et al.
(Chapter 2} and Robinson and Page (Chapter 4} look at
intracolonial genetic structure in honey bees and its impact on
division of labor. Ross (Chapter 8} documents the genetic
structure of polygynous ant colonies and presents data on
differential reproductive output of queens.
Social parasitism is particularly problematical. Social
parasites are often considered to be sister species to their hosts.
If this is the case, then special mechanisms of speciation might be
invoked to explain hosVparasite divergence. Ward (Chapter 7}
considers the mechanisms by which social parasites have evolved
and their role in shaping social diversity.
The new conceptual bases of thought in evolutionary
genetics, as presented in the chapters in this book, are largely
dependent on a new array of techniques. Included in the new
arsenal of evolutionary genetics are computer modeling,
biochemical techniques that allow access to different genetic
markers, and more sophisticated modes of ecological and
behavioral analysis. Further advances on the questions raised
here will depend on the application of these techniques to a set of
taxa that allows a broadly comparative understanding of eusocial
systems.
Most of the papers in this book were first given in a
symposium at the National Conference of the Entomological Society
of America, December 1987. We are grateful to the Society for
its financial support of the symposium. We would also like to
express our thanks to Westview Press, and particularly Kellie
Masterson, for advice and encouragement of this project. Finally,
we are indebted to the Psychobiology Program of the National
Science Foundation for its support of our research programs.
M.D. B. and R. E. P.
CHAPTER 1
INTRODUCTION: THEORY AND EMPIRICAL
RESULTS IN SOCIOGENETICS
Michael D. Breed
The domain of sociogenetics includes karyotype evolution,
sex determination and mating systems, genetics of social behavior,
kin recognition, and caste and levels of selection (Crozier 1987).
Eusocial insects are particularly fertile ground for studies in
sociogenetics; they have complex behavioral patterns which are
often poorly understood from an evolutionary point of view and
they are compatible with a variety of genetic approaches. The
purpose of this volume is to present a broad view of current work
that relates genetics to the special evolutionary features of ants,
bees, and wasps.
In this chapter I define some terminology that will be used
throughout the book. I also discuss some of the major conceptual
issues that are addressed in later chapters and give some views on
some of the evolutionary processes leading to the origin and
maintenance of sterile castes. Sociogenetics is an emerging area of
research; some of the major issues in the sociogenetics of eusocial
insects are discussed.
LEVELS OF EUSOCIAUTY
Batra (1966) introduced the term eusocial to describe
colonies of insects in which there is a reproductive division of
labor. Eusocial colonies have three defining features: 1.) castes
and division of labor, 2.) matrifilial colony organization with two
generations of adults present, and 3.) cooperative brood care. In
such colonies some second generation individuals have
reproductive capacity reduced or absent and raise the offspring of
the parental generation. Individuals with reduced fecundity are
called workers and the fecund parent is the queen.
Other possible social organizations include: communal,
Department of Environmental, Population, and Organismic Biology, The
University of Colorado, Boulder, CO USA 80309-0334
1
