The Lives of Bees
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that occurred in 1800–1860 was followed by reversion to post- agricultural,
second- growth forests starting in the 1870s.
Today, most of the abandoned fields and pastures in the hills south of
Ithaca have grown back to closed- canopy woods that are dotted with the
stone foundations of long- gone houses and barns, stone- lined wells, and
wagon- size stone piles marking the boundaries of what were once plowed
fields. These forestlands also shade many abandoned cemeteries, such as
the small one along Irish Hill Road in the Arnot Forest. Of the 20 or so
graves here, most are detectable only as coffin- shaped depressions in the
soil; a few others are marked with unlettered rocks standing on edge, and
just six have a costly granite gravestone bearing names and dates. The years
of death chiseled into them are 1860, 1862, 1864, 1871, 1881, and 1884.
It seems that the population boom on Irish Hill was over by the 1880s,
whereupon the forests began their return.
The growth rings of the trees covering the hills south of Ithaca also
provide us with a clear record of the rewilding of this region over the last
130 or so years. When I moved back to Ithaca in 1986, my wife, Robin,
and I began buying up 40 hectares (100 acres) of forested land on a hillside
southeast of Ithaca, in a corner of Ellis Hollow, along Hurd Road. It is only
a few miles from where I grew up. Much of what is now our forest was
once the farm of the Hurd family. They lived on it for two generations,
starting in the early 1800s, when Asa Hurd came with Peleg Ellis to settle
the hollow, and ending in 1883, when Asa Hurd’s son, Wesley Hurd, sold
the land shortly before he died at age 82. For the next several decades, a
series of owners cut hay and raised stock on the farm’s fields, but none
lived on the farm, and bit by bit the house, the barn, and the fields were
abandoned. By the 1930s, all the land was growing back to a species- rich,
mostly hardwood forest with some widely spaced eastern white pines
( Pinus strobus) towering over all. There are also dark stands of eastern hem-
lock trees ( Tsuga canadensis) shading the steep, north- facing slopes, for
here the soil stays cool and damp.
Like most of the hilly woodlands south of Ithaca, ours contains diverse
native hardwood tree species. These include red, white, and chestnut oak
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Bees in the Forest, Still 21
( Quercus spp.); sugar, red, striped, and boxelder maple ( Acer spp.); white
and green ash ( Fraxinus spp.); shagbark, bitternut, and pignut hickory
( Carya spp.); yellow, black, and white birch ( Betula spp.); black and pin
cherry ( Prunus spp.); butternut and black walnut ( Juglans spp.); American
beech ( Fagus grandifolia); American basswood ( Tilia americana); tulip tree
( Liriodendron tulipifera); sassafras ( Sassafras albidum); cucumber magnolia
( Magnolia acuminata); American hornbeam ( Carpinus caroliniana); eastern
hophornbeam ( Ostrya virginiana); and even a few American chestnut ( Cas-
tanea dentata) trees. Not surprisingly, there are also old apple ( Malus pum-
ila) and pear ( Pyrus communis) trees around the site of the Hurd family’s
house, which remains clearly marked by its stone- walled cellar hole.
When I cleared the site for our house on this land in the winter of 1988,
I felled several white oaks ( Quercus alba) some 80 centimeters (32 inches)
in diameter at breast height and studied their growth rings. I learned that
these oaks were 100 to 110 years old. I also learned that for the first 50
years of their lives these trees had grown rapidly, quickly reaching diam-
eters of about 56 centimeters (22 inches), but that for the rest of their lives
they had grown more slowly, so that over the last 50 to 60 years their di-
ameters had enlarged by only another 25 centimeters (10 inches). I in-
ferred from the growth rings of these oaks, and from the stubs of rusty
barbed wire poking out of large eastern hemlock and sugar maple ( Acer
saccharum) trees marking a nearby property boundary, that these white
oaks had sprung up in an abandoned pasture in the 1880s and, with little
competition for light, had continued growing rapidly until the 1930s, and
then had grown more slowly as the forest canopy closed in. Today our for-
est, which is typical for the area, is dominated by large trees ranging in age
up to 140 or so years. These trees are large enough to provide homes for
the various animals that need good- size nesting cavities, including rac-
coons, pileated woodpeckers, barred owls, and honey bees. Indeed, in
August 2016, my wife, Robin Hadlock Seeley, heard the roar of a swarm
of bees flying over the treetops and succeeded in following it to a red
maple ( Acer rubrum) tree, where she found the bees flying in and out a dark
knothole, the entrance to their new home (Fig. 2.2).
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Fig. 2.2. Robin’s bee tree. The
entrance to the nest cavity in this
red maple ( Acer rubrum) tree is the
dark knothole near top of photo.
The entrance is 5.9 meters (19.4
feet) up from the base of the tree.
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Bees in the Forest, Still 23
What plants provide the forage for the honey bees living in these wood-
lands? Among the trees growing in the forests, the maple, cherry, bass-
wood, tulip, cucumber magnolia, and chestnut trees are all rich sources of
nectar or pollen or both. Likewise, various shrubs and herbaceous plants,
found in the forest understory or in sunny places along streams and marshy
areas, provide the bees with excellent food. The shrubs include common
alder ( Alnus incana), pussy willow ( Salix discolor), staghorn sumac ( Rhus
typhina), spicebush ( Lindera benzoin), serviceberry ( Amelanchier spp.), haw-thorns ( Crataegus spp.), and northern shrub honeysuckle ( Lonicera canaden-
sis). Among the herbaceous plants growing in sunny spots in these woods,
the most important as food sources for bees are the brambles ( Rubus spp.),
goldenrods ( Solidago spp.), and asters ( Aster spp.). And because honey bee
foragers can fly to food sources 10 kilometers (6 miles) or more from their
homes (discussed further in chapter 8), the colonies living up in the for-
ested hills can also exploit the food- rich flowers growing in the farms,
gardens, roadways, and waste areas down in the valleys. These sites include
apple orchards, stands of black locust trees ( Robinia pseudoacacia), fields of
buckwheat ( Fagopyrum esculentum), hayfields seeded with white and sweet
clovers ( Trifolium repens and Melilotus alba) and alfalfa ( Medicago sativa), and boggy places that contain many excellent sources of pollen and nectar,
including cattails ( Typha latifolia), jewelweed ( Impatiens capensis), and pur-
ple loosestrife ( Lythrum salicaria). In the flower gardens and roadsides, the
bees find many native and introduced plants that provide rich forage. Those
found most commonly are crocuses ( Crocus v
ernus), dandelions ( Taraxacum
officinale), chicory ( Cichorium intybus), milkweeds (e.g., Asclepias syriaca), Japanese knotweed ( Fallopia japonica), and various herbs, including catnip
( Nepeta cataria), borage ( Borago officinalis), and mints ( Mentha spp.).
HOW ABUNDANT ARE WILD HONEY BEE COLONIES
IN THE FORESTS AROUND ITHACA AND BEYOND?
It has long been known that honey bees were introduced to North America
in the mid- 1600s, and that wild colonies of honey bees were thriving
throughout the forests east of the Mississippi River by the late 1700s. It
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was not until the 1970s, however, that we began to get reliable information
about the abundance of honey bee colonies living in the wild anywhere
across this whole continent. Since then, the question of the density (and
thus the spacing) of honey bee colonies living in natural areas has been
investigated in multiple sites in North America, Europe, and Australia.
These explorations have been undertaken to enable us to better under-
stand this important element of the natural history of Apis mellifera.
I began wondering about the abundance of wild colonies back in the
1970s, when I started searching for whatever information was available
about how honey bees live when they inhabit natural cavities rather than
beekeepers’ hives. My richest find was a little book with the intriguing
title Survey of a Thousand Years of Beekeeping in Russia (1971), written by
Dorothy Galton, a scholar in the School of Slavonic and East European
Studies of the University College London. In it, Galton describes the ac-
tivities of tree beekeepers ( bortniki) in Russian forests in the 1100s to the
1600s. This was when Russian princes owned vast bee forests in which the
largest trees were protected by law to provide homes for honey bees. The
people living in these forests served their lords (to whom they belonged)
by gouging out nesting cavities for the bees high in the biggest trees, fitting
them with sturdy doors, and periodically inspecting them to see which
ones were occupied. In late summer, they would return to the trees with
bees, climb each one while carrying wooden pails, open the access door
to expose the bees’ nest, and remove some of the honeycombs, leaving
enough to sustain the colony over winter (Fig. 2.3). The honey and wax
were separated by crushing the honeycombs in water. The honey water was
turned into mead, and the wax bits floating in the honey water were
skimmed off and purified to supply the churches and monasteries in Russia
and the Byzantine Empire with beeswax candles.
Galton explains that the several hundred tons of beeswax exports each
year were hugely important to the economy of medieval Russia, so Russian
tree beekeeping was a well- organized activity. She also provides informa-
tion on the abundance of colonies in the Russian bee forests by citing re-
cords from the late 1600s for the Morozov estate near the city of Nizhny
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Fig. 2.3. Bashkir tree beekeeper extracting honey from a colony nesting in a
manmade tree hollow. Photo taken in the South Ural region of the republic of
Bashkortostan, Russian Federation.
Novgorod, which is east of Moscow. This grand estate possessed four bee
forests that ranged in size from 10 to 88 square kilometers (4 to 28 square
miles) and that contained from 3 to 50 trees with nesting cavities occupied
by honey bees. The average density of known colonies in these four forests
was 0.5 colonies per square kilometer (1.3 colonies per square mile). I
wondered, however: Might the total density of colonies residing in these
forests have been higher than these records indicate? Might there have
been many cryptic colonies living in natural dens high in the trees that the
bortniki had failed to find?
Galton’s estimate of the density of honey bee colonies in the medieval
Russian forests— at least one per square mile—made sense to me, because
over the summers of 1975, 1976, and 1977, I had studied the nest- site
preferences of honey bees and from this work had acquired evidence that
the density of wild colonies in the woods near Ithaca was at least one per
square mile. My study of the bees’ housing preferences involved setting up
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pairs of experimental nest boxes that differed in just one property, such as
cavity size or entrance size, and then seeing which box in a pair was oc-
cupied first by a wild swarm of bees (see the section on nest- site selection
in chapter 5). I nailed the boxes in each pair onto trees standing at least 10
meters (33 feet) apart along roads that ran through heavily forested areas
south of Ithaca in the towns of Dryden and Caroline. Because I spaced the
pairs of nest boxes about a mile apart, the density of my nest- box pairs was
approximately one pair per square mile. On average, I caught 0.5 swarms
per pair of boxes each summer, which told me that there were enough
wild colonies living in the area to generate 0.5 swarms per square mile
(0.2 swarms per square kilometer). (To the best of my knowledge, there
were few managed colonies in the areas where I conducted this study.) It
seemed likely, however, that more than 0.5 swarms had been produced per
square mile, because it was unlikely that my nest boxes had attracted all
the swarms in the places where I had put them. So, I wondered, might
there actually be several wild colonies per square mile in the woods around
Ithaca? To answer this question, I knew that I must find a large forest and
then try to locate all the wild honey bee colonies living within it.
Fortunately, Cornell University owns a sprawling research forest, the
17- square- kilometer (6.6- square- mile) Arnot Forest, which lies only
about 25 kilometers (15 miles) southwest of Ithaca (see Fig. 2.1). This for-
est extends over parts of the towns of Newfield (in Tompkins County) and
Cayuta (in Schuyler County) (Fig. 2.4). Moreover, the woodland habitat
found in the Arnot Forest doesn’t end at its boundary lines. Instead, it
continues beyond the Arnot Forest’s northern, western, and southern bor-
Fig. 2.4. Top: Aerial photo of the Arnot Forest with forest boundaries indicated
by yellow lines. Red bars: upper, 1 kilometer (upper); lower, 1 mile. North is up.
The complex of buildings in the valley to the west is a sawmill that processes
hardwood logs harvested from the vast woodlands in southern New York and
northern Pennsylvania. Bottom: View of the Arnot Forest, looking southeast from
Irish Hill Road. Photo taken in late September, as trees began to show their au-
tumn colors.
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ders into the steeply sloped Cliffside and Newfield State Forests and
beyond its eastern border into several private forests. Furthermore, on
the far sid
e of the narrow valley with good farmland (Pony Hollow) that
skirts the northwestern corner of the Arnot Forest lies the rugged,
47- square- kilometer (18- square- mile) Connecticut Hill Wildlife Manage-
ment Area, a state- owned preserve that is also mostly woodland. Agricul-
ture was abandoned in all these places more than 100 years ago, and today
they comprise mostly hardwood forests but also some wetlands and old
field habitat. The whole area provides a superb setting for the study of
wildlife, including wild honey bees.
In July 1978, a friend and fellow student of the bees, Kirk Visscher, and
I began to explore for wild colonies of honey bees living in the Arnot For-
est. We did so using the tools and methods of bee hunting (also called “bee
lining”), an outdoor pursuit that has been practiced for centuries in Europe
and North America. Most bee hunters search for wild colonies to get some
honey and have some fun. Our aim was purely scientific—to discover how
many wild colonies of honey bees were living in the Arnot Forest—but we
had some fun, too.
When Kirk and I began our bee hunting, we were rank novices and had
nobody to teach us the craft, but we had come across a first- rate guidebook
on the subject that was published in 1949, The Bee Hunter. It was written
by George H. Edgell, a bee hunter (and professor of architectural history
at Harvard University) with decades of experience finding wild colonies
of honey bees living in the mountains around his summer home in New-
port, New Hampshire. Edgell explains how you start a hunt for a wild
colony by going to a good- size clearing (the bigger the better) that is well
stocked with flowers attractive to honey bees. Here, you use a small, two-
chambered apparatus called a bee box to capture some bees that are forag-
ing on the flowers. Once you have imprisoned a half dozen or so bees in
your bee box, you tuck inside it a small square of beeswax comb filled with
sugar syrup. The bees in the box will find the syrup- filled comb, tank up
on its delicious contents, and make ready to fly home. After giving the bees
about five minutes to stumble upon the bait in your bee box, you let them
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