Table Of ContentThe Dark Side of the Hive
The Dark Side of the Hive
THE EVOLUTION OF THE IMPERFECT HONEY BEE
Robin Moritz and Robin Crewe
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Library of Congress Cataloging- in- Publication Data
Names: Moritz, Robin F. A., and Crewe, Robin M.; authors.
Title: The dark side of the hive / Robin Moritz and Robin Crewe.
Description: New York, NY : Oxford University Press, 2018. |
Includes bibliographical references and index.
Identifiers: LCCN 2018010888 | ISBN 9780190872281
Subjects: LCSH: Honeybee—Life cycles.
Classification: LCC QL568.A6 M594 2018 | DDC 595.79/9156—dc23
LC record available at https://lccn.loc.gov/2018010888
1 3 5 7 9 8 6 4 2
Printed by Sheridan Books, Inc., United States of America
{ CONTENTS }
Preface vii
Acknowledgments ix
List of Image Credits xi
1. Introduction 1
2. Out of the Dark 7
3. A Difficult Diet 19
4. The Chemistry of Social Regulation 29
5. The Reproductive Machine 37
6. The Worker Bee in a Variety of Guises 53
7. Diseases, Pests, and Parasites 83
8. The Idiosyncrasies of Sex and Reproduction 97
9. Apiculture and Long- Suffering Bees 121
10. Dark Sides of Honey Bee Science 145
11. A Silver Lining for the Future of Bees? 153
References 159
Index 179
{ PREFACE }
Producing a monograph on the life history of the honey bee, Apis mellifera,
may seem somewhat quixotic at first sight in view of the fact that there are a
series of excellent monographs on this topic that have been published during
the past three decades. However, we believe that these volumes have a bias that
is too strongly focused on the harmony and the perfection of cooperation in the
colony. Tom Seeley’s excellent treatises Honeybee Ecology (1985), The Wisdom
of the Hive (1995), and, most recently, Honeybee Democracy (2010) have had
a major influence on students of honey bee biology. Indeed, they stand out as
seminal works in organismal biology in the mold of the work of von Frisch,
who won the Nobel Prize for his work on honey bees. Mark Winston’s book,
The Biology of the Honey Bee (1987), as well as Robin Moritz and Edward
E. Southwick’s Bees as Superorganisms— An Evolutionary Reality (1991) have
dealt in great detail with the marvels of cooperation inside the colony. Yet as
with any complex social system, honey bee societies are prone to error, rob-
bery, cheating, and social parasitism. The honey bee colony is thus far from
being a harmonious, cooperative whole. It is full of individual mistakes, ob-
vious maladaptations, and evolutionary dead ends. Conflict, cheating, worker
inefficiency, and curious reproduction strategies all occur. The perfection that is
perceived to exist in their social organization is a function of a particular exper-
imental focus on the colony as a whole rather than exploring the idiosyncrasies
of its individual members.
The fact that honey bee colonies get by remarkably well despite many seem-
ingly odd biological features that are often dismissed as aberrations requires us
to focus attention on these very “aberrations” because they are central to un-
derstanding all aspects of social organization. Since we have worked together
for more than two decades on the chemical ecology, genetics, and evolution of
parasitic honey bee workers, we believed it is now overdue to report on the plas-
ticity of social organization in the honey bee colony with a view to achieving a
more nuanced understand of the evolution of sociality in these insects.
What we cover in this volume is not designed to suggest that the work of our
colleagues and our own previous work require revision or reconsideration but,
rather, provides a richer understanding of the real life of a honey bee in the
colony. Our work thus focuses on the role of the individual within the colony
rather than studying the colony as a biological entity (superorganism). We try
to dissect the various careers a male and a female honey bee can have and their
roles in colony organization.
viii Preface
Colonies of honey bees are made up of a number of families because the
queens are multiply mated by a number of drones (males). In order to under-
stand the effects of this genetic diversity within the colony, we provide a thor-
ough discussion of how the variance of individual worker phenotypes drives
self- organized processes in a colony.
In addition, we show how competition using both physical force and chem-
ical signaling drives colonial organization. Here, we deal with handicaps that
limit the use of physical force and the chemical arms races that drive competi-
tion in relation to worker reproduction.
This monograph focuses on bees as individuals in the colony. The story that
we tell spans the full range of biological disciplines ranging from genomics
to systems biology. We explore the situations in which individual interests are
pursued often at the expense of the colony, and we show that the solutions that
have evolved are often less than optimal.
{ ACKNOWLEDGMENTS }
This volume would not have been possible without Anne and Mary allowing
us to drone on endlessly about honey bees when indeed we should have been
much more polite and addressed other more important topics. We are deeply
indebted for this indulgence and have no really convincing ideas about how to
repay it, particularly because we are not certain that this may not happen again
despite solemn undertakings.
We are also grateful to the late Ingemar Fries, Christian Pirk, Anja Buttstedt,
and two anonymous reviewers for providing many important comments that
markedly improved the text. We considered most of those, but not all; therefore,
any errors and misconceptions remain ours and cannot be blamed on others.
The generous funding of many different public institutions and funding or-
ganizations that in turn received their money from the taxpayers in Germany,
South Africa, and also all European Union member states allowed us to study
honey bee biology over several decades. It was this long period of familiarity
with honey bee biology that made us bold enough to face the challenge of
writing this volume.
During the actual writing phase, this project was significantly facilitated
by the receipt of the Harry Oppenheimer Fellowship Award from the
Oppenheimer Memorial Trust that allowed RMC to take periods of extended
leave in Germany each year from 2014 to 2017. This leave allowed us to work
together on the preparation of the manuscript and also to encourage doctoral
students to pursue new lines of research.
