the
Lives
of
Bees
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the
LIVES
of
BEES
The Untold Story
of the Honey Bee
in the Wild
Th o ma s D. S e e l e y
PRINCETON UNIVERSITY PRESS
PRINCETON AND OXFORD
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Copyright © 2019 by Princeton University Press
Published by Princeton University Press
41 William Street, Princeton, New Jersey 08540
6 Oxford Street, Woodstock, Oxfordshire OX20 1TR
press.princeton.edu
All Rights Reserved
LCCN 2019930923
ISBN 978- 0- 691- 16676- 6
British Library Cataloging- in- Publication Data is available
Editorial: Alison Kalett and Kristin Zodrow
Production Editorial: Brigitte Pelner
Text and Jacket/Cover Design: Carmina Alvarez
Jacket/Cover Credit: Bees/iStock
Production: Jacqueline Poirier
Publicity: Sara Henning- Stout
Copyeditor: Amy K. Hughes
This book has been composed in Perpetua and Bembo
Printed on acid- free paper ∞
Printed in the United States of America
1 3 5 7 9 10 8 6 4 2
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Dedicated to Roger A. Morse (1927–2000), who for more than
forty years as a scientist, writer, and teacher at Cornell University
informed and entertained students about the honey bee, and whose
support of the author laid the foundation for this book
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Contents
Preface
vii
1 Introduction 1
2 Bees in the Forest, Still
17
3 Leaving the Wild
57
4 Are Honey Bees Domesticated?
79
5 The Nest
99
6 Annual Cycle
140
7 Colony Reproduction
155
8 Food Collection
187
9 Temperature Control
215
10 Colony Defense
243
11 Darwinian Beekeeping
277
Notes
293
References
317
Acknowledgments
337
Illustration Credits
341
Index
349
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Preface
We humans have always been fascinated by the honey bee— Apis mellifera,
the “honey- carrying bee.” For hundreds of thousands of years, our earliest
ancestors in Africa, Europe, and Asia surely marveled at this bee’s astonish-
ing industry in storing honey and making beeswax, two substances of great
value. More recently, during the last 10,000 years, we invented the intri-
cate craft of beekeeping and we began our scientific studies of honey bees.
It was, for example, the ancient philosopher Aristotle who first described
this bee’s practice of “flower constancy”: a worker bee generally sticks to
one type of flower throughout a foraging trip to boost the efficiency of
her food collection. And within the past few hundred years, we have writ-
ten tens of thousands of scientific articles about the honey bee, many on
the practical stuff of apiculture but also countless others on the funda-
mental biology of this endlessly enchanting bee. We have also authored
thousands of books on bees and beekeeping. In the United States alone,
nearly 4,000 titles on beekeeping, honey bee science, children’s tales
about bees, and the like have been published between the 1700s and 2010.
Given humanity’s enduring fascination with the honey bee, it is a curious
thing that until recently we have known rather little about the true natural
history of Apis mellifera—that is, about how colonies of this species live in
the wild. What explains our long delay in making a broad reconnaissance
of the natural lives of honey bees? I think the answer is simple: the most
ardent students of this industrious and intriguing insect—beekeepers and
biologists—have almost always worked with managed colonies inhabiting
man- made hives crowded in apiaries, not wild colonies occupying hollow
trees and rock crevices dispersed across the landscape. Managed colonies
are the ones that produce our honey and pollinate our crops, so it is not
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x
Preface
surprising that beekeepers have focused their attention on the colonies
living in their hives. Managed colonies are also the ones best suited for
scientific investigations, which require controlled experiments, so it is also
not surprising that biologists too have worked primarily with colonies liv-
ing in artificial homes. The Nobel laureate Karl von Frisch, for example,
would never have discovered the meaning of the honey bee’s waggle dance
if he had not worked with a colony living in a glass- walled observation hive,
labeled some of its foragers with paint marks for individual identification,
and then watched how these bees behaved inside his hive when they re-
turned from an artificial food source, a little dish of sugar syrup that he
had set up in the courtyard just outside his laboratory.
Humankind’s long- standing focus of interest on honey bees housed in
hives—whether clay cylinders, woven baskets, wooden boxes, or (most
recently) polystyrene containers—continues to this day. Over the last few
decades, however, beekeepers and biologists have begun to investigate how
these engaging insects live when they do so without our supervision, and
this “return to nature” has opened our eyes to many new mysteries in the
lives of honey bees. This book is my attempt to review what has been
learned about how colonies of honey bees live in their natural world. We
will see that the free- living colonies residing in tree cavities and rock crev-
ices lead lives that differ substantially from the lives of beekeepers’ colonies
inhabiting the white boxes that we see parked in apple orchards and blue-
berry fields, jam- packed in apiaries, and nestled in backyards. Perhaps
most remarkably, we will see that the wild colonies are surviving and are
maintaining their numbers, while at the same time some 40 percent of the
colonies managed by beekeepers are dying each year.
The story of the wild honey bees is an important one, because it can
expand how we view ourselves in relation to Apis mellifera and how we
conduct the craft of beekeeping. We might, for example, start to think of
the honey
bee not only as a compliant, hardworking insect that we can ma-
nipulate to produce honey crops and fulfill pollination contracts but also
as a wondrous insect that we can admire, respect, and treat in truly bee-
friendly ways. In the coming chapters, I will show how numerous strands
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Preface xi
of work on honey bee colonies living in the wild—their nest architecture,
nest spacing, foraging range, mating system, disease resistance, colony
genetics, and more—have come together to reveal how these colonies
living all on their own are thriving. In the final chapter, “Darwinian Bee-
keeping,” I will discuss how we can use this growing body of knowledge to
address a critically important issue: how to be better partners with Apis
mellifera, a species that has sweetened the lives of humans for many thou-
sands of years, and on which humanity’s food supply depends more and
more every year.
My fascination with honey bees living in the wild began in the spring of
1963, when I was not quite 11 years old. I lived then, as I still do today, in
a little valley called Ellis Hollow, which lies a few miles east of Ithaca, New
York. It is a stream valley barely 1.6 kilometers (1 mile) wide and 3.2
kilometers (2 miles) long, and it lies between two steep- sided hills, Mount
Pleasant and Snyder Hill. These are parallel units of an ancient escarpment
of sandstone that runs through the Finger Lakes region of central New York
State and adds rugged beauty to the area. Ellis Hollow was a good place to
grow up, because the wooded hillsides and the valley bottom—with its
sloping fields bordered by dark groves of hemlock trees, sunny swamps
patrolled by dragonflies, and the gently flowing Cascadilla Creek meander-
ing through it all—seemed endless. It is where I first observed a magnifi-
cent pileated woodpecker chiseling into a tree for carpenter ants, first
watched a steely- eyed snapping turtle laying eggs deep in moist soil, and
first showed my pet raccoon how to hunt for crayfish under rocks in little
streams. Fortunately, there were no “No Trespassing” signs to limit my
explorations of this ever- fascinating place. Even today, when I drive home
along the Ellis Hollow Creek Road, I take note of spots that I still need to
investigate.
One day, back in early June 1963, I was walking along Ellis Hollow
Road, when I heard a loud buzzing sound and saw a bread- truck- size cloud
of honey bees circling the ancient black walnut tree that stands beside the
road about 100 meters (330 feet) east of my family’s house. I was fright-
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xii Preface
ened, so I crossed to the shady woods on the other side of the road and
watched from what felt like a safe distance. From there, I saw that the bees
were landing on a thick limb about 4 meters (12 feet) off the ground,
blanketing it with their thousands of leathery- brown bodies, and streaming
into a knothole the size of a golf ball. The bees were moving in! Suddenly
this huge tree, which already I had prized as a good climbing tree and a
rich source of black walnuts, was super special. It was now a bee tree! I
visited it often that summer and gradually overcame my fear of the bees,
eventually learning that I could watch them close up (while perched atop
a stepladder) without being stung. It was a time of wonder.
My mother noted my curiosity about the bees, and for Christmas 1963
my parents gave me a beautifully illustrated children’s book on honey bees,
The Makers of Honey (1956), written by Mary Geisler Phillips. I read it
closely and liked how it introduced me to the biology of honey bees. It sits
on my writing desk as I type these words. I feel an especially deep connec-
tion to this little book because its author was also a Cornell University
professor, in the College of Home Economics (now the College of Human
Ecology), where she served as editor of the college’s radio scripts and
outreach publications. Moreover, the first professor of apiculture at Cor-
nell, Everett F. Phillips, was her husband.
Given these lovely introductions to honey bees as a boy, especially the
firsthand observations of a bee colony living wild in a tree, it is not surpris-
ing that, when I started graduate school in 1974 to earn a PhD in biology,
and I had to choose a topic for my thesis research, I decided to investigate
what honey bees seek when they (not a beekeeper) choose their living
quarters. In doing so, I figured that I could apply to honey bees the “know-
thy- animal- in- its- world” rule that I was learning from my thesis adviser at
Harvard, the German ethologist Bert Hölldobler. I also hoped that I could
foster a new approach to studying honey bees, one in which we view them
as amazing wild creatures that live in hollow trees in forests, not just as the
“angels of agriculture” that live in white boxes in apiaries. Furthermore, I
hoped that through this thesis research I would solve the mystery I had
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Preface xiii
sensed back in 1963 as I cautiously watched a swarm moving into its new
home: What was it about the dark cavity in the black walnut tree near my
parent’s house that attracted the bees to make it their dwelling place?
Watching that swarm take up residence in that tree on that day is the spark that ignited my long- standing passion to understand how honey bees live
in the wild.
Thomas D. Seeley
Ithaca, New York
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the
Lives
of
Bees
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1
INTRODUCTION
We have never known what we were doing
because we have never known what we were undoing.
We cannot know what we are doing until we know
what nature would be doing if we were doing nothing.
—Wendell Berry, “Preserving Wildness,” 1987
This book is about how colonies of the honey bee ( Apis mellifera) live in the
wild. Its purpose is to provide a synthesis of what is known about how
honey bee colonies function when they are not being managed by bee-
keepers for human purposes and instead are living on their own and in
ways that favor their survival, their reproduction, and thus their success
in contributing to the next generation of colonies. Our goal is to under-
stand the natural lives of honey bees—how they build and warm their
nests, rear their young, collect their food, thwart their enemies, achieve
their reproduction, and stay in tune with the seasons. Besides looking at
how honey bee colonies live in nature, we will examine why they live as
they do when they manage their affairs themselves. In other words, we
will also explore how natural selection has shaped the biology of this im-
portant species during its long journey through the labyrinth of evolu
tion.
Doing so will reveal how Apis mellifera achieved a native range that includes
Europe, western Asia, and most of Africa, and so became a world- class
species even before beekeepers introduced it to the Americas, Australia,
and eastern Asia.
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2
Chapter 1
Knowing how the honey bee lives in its natural world is important for
a broad range of scientific studies. This is because Apis mellifera has become
one of the model systems for investigating basic questions in biology, es-
pecially those related to behavior. Whether one is studying these bees to
solve some mystery in animal cognition, behavioral genetics, or social be-
havior, it is critically important to become familiar with their natural biol-
ogy before designing one’s experimental investigations. For example,
when sleep researchers used honey bees to explore the functions of sleep,
they benefited greatly from knowing that it is only the elderly bees within
a colony, the foragers, that get most of their sleep at night and in compara-
tively long bouts. If these researchers had not known which bees are a
colony’s soundest sleepers come nightfall, then they might have failed to
design truly meaningful sleep- deprivation experiments. A good experi-
ment with honey bees, as with all organisms, taps into their natural way
of life.
Knowing how honey bee colonies function when they live in the wild
is also important for improving the craft of beekeeping. Once we under-
stand the natural lives of honey bees, we can see more clearly how we
create stressful living conditions for these bees when we manage them
intensively for honey production and crop pollination. We can then start
to devise beekeeping practices that are better—for both the bees and our-
selves. The importance of using nature as a guide for developing sustain-
able methods of agriculture was expressed beautifully by the author, envi-
ronmentalist, and farmer Wendell Berry, when he wrote: “We cannot
know what we are doing until we know what nature would be doing if we
were doing nothing.”
The current state of beekeeping shows us all too clearly how problems
in the lives of animals under our management can arise when we fail to
consider how they would be living if we were not forcing them to live in
artificial ways that serve mainly our own interests. Many beekeepers—
The Lives of Bees Page 1
