especially those in North America who practice industrial- scale beekeep-
ing with tens of thousands of colonies of Apis mellifera, the species that is
the focus of this book—are experiencing colony mortality rates of 40
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Introduction 3
percent or more each year. To be sure, this is not due entirely to the colony
management practices of beekeepers. Changes in the crop production
practices of farmers, especially the use of systemic insecticides that are
absorbed by plants and contaminate their nectar and pollen, and the switch
in many places to growing corn and soybeans instead of clover and alfalfa,
also play roles in this sad story. But the heavy- handed manipulations of the
lives of the honey bee colonies housed in beekeepers’ hives certainly do
contribute to the sky- high rates of colony mortality. We will see that when
beekeepers force colonies to live crowded together in apiaries—where
the bees’ homes are less than 1 meter (ca. 3 feet) apart, rather than the
hundreds of meters (at least 1,000 feet) apart in nature—beekeepers
boost the efficiency of their work but they also foster the spread of the
bees’ diseases. Likewise, when beekeepers supersize their colonies by
housing them in huge hives that are nearly as tall as themselves, rather than
in smaller hives the size of the bees’ natural nesting cavities, they boost the
honey production of their colonies, but they also turn them into stupen-
dous hosts for the pathogens and parasites of Apis mellifera, such as the
deadly ectoparasitic mite Varroa destructor.
Given the harmful effects on the bees that arise from the standard prac-
tices of beekeeping, it is not surprising that many beekeepers are now
exploring alternative approaches to this craft. These folks are keen to use
nature as a model, and this requires a solid understanding of how honey
bees live on their own in nature. To help readers who want to adjust their
beekeeping practices to make them more bee- friendly, I have included a
final chapter on what I like to call “Darwinian beekeeping,” which is an
approach to beekeeping that aims to give bees the opportunity to live the
way they do in the wild.
FOCUS ON WILD COLONIES IN THE
NORTHEASTERN UNITED STATES
This book does not attempt to provide a comprehensive account of how
Apis mellifera lives in nature across its vast geographic range, which now
includes Europe, some of Asia, all of Africa except the great desert regions,
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4
Chapter 1
Fig. 1.1. Left: Bee- tree home of a wild colony
of honey bees living in the Arnot Forest, of
Cornell University, in the United States. Red
arrow indicates the small knothole entrance
of this colony’s nest. Right: Nest entrance of a
wild colony of honey bees living in Munich,
Germany.
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Introduction 5
most of North and South America, and parts of Australia and New Zea-
land. Instead, it focuses on how colonies of our most important pollinator
are living in the wild in the deciduous forests of the northeastern United
States, a place where they have thrived as an introduced species for nearly
400 years. This is also the place where, for more than 40 years, my collabo-
rators and I have studied the behavior, social life, and ecology of honey bees
living in the wild (Fig. 1.1). Although our studies are based on honey bees
living outside their native range, I believe that what we have learned about
how honey bees live in the woods in the northeastern corner of the United
States can help us understand how these bees originally lived in nature in
Europe, especially in its northern and western regions.
Until the mid- 1800s, all the honey bees living in the northeastern
United States were descendants of the colonies of honey bees that were
brought to North America from northern Europe starting in the early
1600s. Insect taxonomists recognize some 30 subspecies (geographic vari-
ants) of Apis mellifera, and they refer to the honey bees native to northern
Europe as members of the subspecies Apis mellifera mellifera Linnaeus. This
subspecies of Apis mellifera—also called the dark European honey bee—has
the distinction of being the first kind of honey bee to be described taxo-
nomically. This was done 360 years ago, in 1758, when Carl Linnaeus, a
professor of botany and zoology at Uppsala University in Sweden, pub-
lished his work Systema Naturae, in which he presented the system of taxo-
nomic classification that biologists have used ever since.
The dark European honey bee is so named because its body color ranges
from dark brown to jet black and historically it lived throughout northern
Europe, from the British Isles in the west to the Ural Mountains in the east,
and from the Pyrenees and Alps in the south to the coasts of the Baltic Sea
in the north (Fig. 1.2). We know from archaeological studies, which have
found traces of beeswax in fragments of pottery vessels dating from 7,200
to 7,500 years before present, that this bee was living in Germany and
Austria some 8,000 years ago. We also know from genetic studies of the
bees themselves that as the climate of northern Europe underwent post-
glacial warming, starting about 10,000 years ago, this bee expanded its
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6
Chapter 1
Arctic Circle
6
Ural Mt.
0˚
Oslo
Moscow
London
Berlin mel ifera
Paris
Vienna
Alps
Caucasus Mt.
Pyren.s
carnica
l
caucasica
i
40
g
˚
ustic
Istanbul
a
Madrid
macedonica
iberica
Fig. 1.2. Distribution map of the dark European honey bee, Apis mellifera mellifera.
Green line: original distribution limits to the west, north, and east. Vertically
hatched line: transition zone to the honey bee races of southern and eastern
Europe ( A. m. ligustica, carnica, macedonica, and caucasica). Red dashed line: northern limit of beekeeping.
range north and east from Ice Age refugia—pockets of woodlands in the
mountains of southern France and Spain—as it tracked the expansion of
forests populated with thermophilic trees such as willow, hazel, oak, and
beech. Evidently, Apis mellifera mellifera flourished as it spread and eventu-
ally extended its range farther north within Europe than any other subspe-
cies, drawing on the adaptations for winter survival that it had evolved
during the glacial period. It is estimated that within the eastern, heavily
forested two- thirds of its range (from eastern Germany to the Urals) there
once lived millions of colonies of the dark European honey bee. And there
is no doubt that tree beekeeping—cutting cavities i
n trees to create nest
sites and then harvesting honey without killing the colonies living in the
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Introduction 7
artificial hollows—provided most of the beeswax and honey traded in
Europe in the Middle Ages. A vestige of this centuries- old tradition of tree
beekeeping is found in the South Ural region of the republic of Bashkor-
tostan, part of the Russian Federation. Colonies of pure A. m. mellifera still
inhabit the forests of this region, and Bashkir tree- hive beekeepers still
harvest basswood ( Tilia cordata) honey from colonies residing in man-
made nest cavities high in the trees.
The dark European honey bee is superbly adapted to living in forested
regions with relatively cool summers and long, cold winters. It is not sur-
prising, therefore, that when bees of this subspecies were brought to Mas-
sachusetts, Delaware, and Virginia in North America by English and Swed-
ish immigrants starting in the early 1600s, they escaped (swarmed) from
the beekeepers’ hives and soon became an important part of the local
fauna. Already in 1720, a Mr. Paul Dudley published, in Philosophical
Transactions, a journal of the Royal Society of London, a letter titled “An
account of a method lately found out in New- England for discovering
where the bees hive in the woods, in order to get their honey.” Analysis of
letters, diaries, and accounts of travels in North America written in the
1600s and 1700s has revealed that these honey bees spread speedily across
the heavily forested eastern half of North America below the Great Lakes
(Fig. 1.3). Also, the journals of the Lewis and Clark Expedition document
the dark European honey bee’s rapid colonization of North America east
of the Mississippi River. For example, on Sunday, 25 March 1804, shortly
after the expedition party had left St. Louis and was camped along the
Kansas River, William Clark wrote in his journal: “River rose 14 Inch last
night, the men find numbers of Bee Trees, & take great quantities of honey.”
These days, the honey bees living wild in the forests of the northeastern
United States are no longer a genetically pure population of Apis mellifera
mellifera. This is because, in 1859, following the advent of steamship ser-
vice between Europe and the United States, American beekeepers began
importing queen bees of several other subspecies of Apis mellifera, ones that
are native to southern Europe or northern Africa. These imports contin-
ued for more than 60 years, during which time many thousands of mated
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8
Chapter 1
45˚ N
1794
Boston 1640
1790
1644
Chicago
40˚ N
1826
New York
1796
1698
1655
1622
1792
1780
Norfolk
1788
35˚ N
1750
1796
1796
30˚ N
1736
1770
New Orleans
1765
1773
Miami
Fig. 1.3. The dispersal of the dark European honey bee across eastern North
America following introductions (indicated by solid arrows) in Virginia, Massa-
chusetts, Connecticut, and Maryland in the 1600s, and in Alabama in 1773.
Dashed arrows indicate the bee’s subsequent spread.
queen bees were shipped to North America, but they abruptly ceased in
1922. This was the year the U.S. Congress passed the Honey Bee Act,
which prohibited further imports to protect honey bees in the United
States from the Isle of Wight disease, an unspecific but supposedly highly
infectious and lethal disease named for the location of its first reputed
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Introduction 9
outbreak, in southern England. As we shall see, the genetic composition
of the wild honey bees living in the northeastern United States is now a
blend of the genes of A. m. mellifera and several other subspecies of Apis
mellifera. Of those introduced in the late 1800s and early 1900s, the three
most important all came from south- central Europe: A. m. ligustica (from
Italy), A. m. carnica (from Slovenia), and A. m. caucasica (from the Caucasus
Mountains). Several other subspecies were introduced from the Middle
East and Africa— A. m. lamarckii (from Egypt), A. m. cypria (from Cyprus),
A. m. syriaca (from Syria and the eastern Mediterranean region), and
A. m. intermissa (from northern Africa)—but they did not prove popular,
and it seems that they are not well represented genetically anywhere in the
United States.
More recently, in 1987, a subspecies of Apis mellifera that is native to
eastern and southern Africa, A. m. scutellata, entered the southern United
States by way of Florida, perhaps when a beekeeper, seeking bees that
would thrive in subtropical Florida, smuggled in some queens of this trop-
ically adapted subspecies. Since then, colonies of A. m. scutellata have in-
deed thrived in Florida and have greatly influenced the genetics of the
honey bees living in the southeastern United States but not of the honey
bees living in the northeastern United States, probably because colonies
of A. m. scutellata cannot survive northern winters . Bees of the African
subspecies A. m. scutellata entered the United States a second time and in a
second place in 1990, when swarms flew across the U.S.–Mexico border
into Texas. Here again they mixed with the European honey bees already
in residence. Since then, populations of colonies that are hybrids of African
and European honey bees (so- called Africanized honey bees) have devel-
oped in the humid, subtropical parts of southern Texas and in the south-
ernmost parts of New Mexico, Arizona, and California. As of 2013, the
gene pool of the Africanized bees in southern Texas still had a small genetic
contribution (ca. 10%, for both mitochondrial and nuclear genes) from
European honey bees.
The complex history of countless introductions to North America of
honey bees from various regions of Europe, the Middle East, and Africa
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10 Chapter 1
raises an important question: What is the mix of subspecies of Apis mellifera
in the wild colonies living in the northeastern United States, the main
subjects of this book? Fortunately, we now have a clear answer to this ques-
tion for the colonies living in the vast woodlands of southern New York
State. In 1977 and then again in 2011, I collected worker bees from 32
wild colonies living in this heavily forested region. The 32 sets of bees from
1977 were stored as pinned (voucher) specimens in the Cornell University
Insect Collection, and the 32 sets of bees from 2011 were stored in vials
filled with ethanol, which preserves DNA quite nicely. In 2012, specimens
from both groups of bees were shipped to one of my former students,
Professor Alexa
nder S. Mikheyev, who heads the Ecology and Evolution
Unit at the Okinawa Institute of Science and Technology in Japan. There
the DNA was extracted from one bee from each of the 64 colonies that I
had sampled and an analysis based on whole- genome sequencing was per-
formed to determine the subspecies composition of both the 1977 (“old”)
and the 2011 (“modern”) populations of bees (Fig. 1.4).
This genetic detective work found that the bees in both the old and
modern samples are primarily descendants of two of the subspecies of Apis
mellifera that were imported from southern Europe, specifically from Italy
and from Slovenia: A. m. ligustica and A. m. carnica, respectively. This finding was not surprising, because these are the two subspecies that have proven
the most popular among beekeepers in North America since the 1800s.
Colonies of these two subspecies tend to be good- natured (not prone to
stinging) and good producers of honey. What was surprising, however, was
the discovery that the bees in both the old and modern samples also pos-
sessed many genes from the dark honey bees imported from north of the
Alps ( A. m. mellifera) starting in the 1600s and from the gray mountain
honey bees ( A. m. caucasica) imported from the Caucasus Mountains start-
ing in the late 1800s (see Fig. 1.4). This genetic sleuthing also revealed that
the bees in the modern (2011) sample, but not those in the old (1977)
sample, have a small percentage (less than 1%) of genes from two African
subspecies: A. m. scutellata, native to Africa south of the Sahara, and A. m.
yemenetica, native to the hot arid zones of Arabia (e.g., Saudi Arabia, Yemen,
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Introduction 11
Old
Modern
Proportion of ancestral population
ligustica,
mellifera
caucasica
scutellata
yemenetica
carnica
Fig. 1.4. Ancestries of the honey bees ( Apis mellifera) living in the forests south
of Ithaca, New York. Both the old (1970s) and the modern (2010s) populations
are largely descendants of bees from southern and southeastern Europe: the
subspecies A. m. ligustica, carnica, and caucasica, which have been popular with beekeepers in North America since the 1880s. Both old and modern populations
also have clear ancestry from the dark European honey bees ( A. m. mellifera) of
northern Europe that were introduced to North America starting in the 1600s.
The modern population also shows a small ancestry of bees from Africa ( A. m.
The Lives of Bees Page 2
