First published online November 14, 2008
Journal of Experimental Biology 211, 3737-3743 (2008)
Published by The Company of Biologists 2008
doi: 10.1242/jeb.003640
Honeybees can learn the relationship between the solar ephemeris and a newly-experienced landscape
William F. Towne
Department of Biology, Kutztown University of Pennsylvania, Kutztown, PA
19530, USA

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Fig. 1. Aerial views of the field sites indicating locations of hives (H) and
feeders (F). The feeder at the valley site (A) was only 2 m from the hive
entrance and is not shown. The white arrows in B and C are 190 m long and
indicate the bees' outward flights to the feeder. North is indicated by the
black arrowhead in C. Adjacent contour lines in all three figures are
separated by 6.1 m of elevation. The hive at the valley site (A) was at the
bottom of a thinly wooded valley. The hives at the treelined sites B and C
were placed along sloping treelines adjacent to agricultural fields, and the
bees flew uphill to the feeder in each case. The valley site A is at 75
deg.47'43.4''W, 40 deg.36'41.4''N; the training treeline
B is at 75 deg.46'10.5''W, 40 deg.36'27.9''N; and the
test treeline C is at 75 deg.47'10.2''W, 40
deg.37'12.9''N. Photos by the United States Geological Survey,
courtesy of the USGS and TerraServer-USA.
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Fig. 2. Dance indications of control bees native to the training treeline (A) and
experimental bees native to the valley site (B) under overcast skies at the
test site on 7 August 2005. Times are given in local solar time (LST). The
valley natives had recently acquired several days' experience at the training
treeline. Each symbol represents the visual average of at least five wagging
runs during a single bout of dancing; each bee was scored only once after a
single trip to the feeder. The hive was at the test treeline
(Fig. 1C), where the direction
to the feeder, and thus the correct dance indication, was 353.5 deg.
(horizontal black line in each panel). The predicted direction for dances
oriented by memory of the sun's course at the training treeline was 165 deg.
(horizontal gray line in each panel). The dances of seven different valley
natives that started dancing before the sun first appeared (vertical line at
08:38 h) are shown with unique symbols in B (the bees' identities and symbols
are given in the legend), and the dances of six additional bees that started
dancing after the sun appeared are all shown with open squares (`others' in
the legend). The sky bar above each panel indicates sky conditions: black
indicates complete overcast; white indicates that the sun was visible. There
were no periods of blue sky without sun. Statistical analyses include only
those dances that occurred before the sun first appeared (thin vertical line
at 08:38 h). CW; clockwise.
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Fig. 3. Dance indications of control bees native to the training treeline (A) and
experimental bees native to the valley site (B) under overcast skies at the
test site on 8 August 2005. In A, the dances of three different control bees
are distinguished with unique symbols: these are the two control bees that
danced the most (open and shaded circles) and a third bee (triangles) that
showed an unusual dancing pattern. The other six bees are all shown with black
circles. In B, the two valley natives that danced the most are shown with
unique symbols (open and shaded circles); all other bees (26) are shown with
black squares. In B, bimodal dances, in which the bee indicated two different
directions on alternate wagging runs, are indicated with broken vertical lines
connecting the two dance directions. One of the bees that danced the most
(shaded circles) performed one bimodal dance; the two directions are indicated
by shaded circles with dots in the center connected by a broken vertical line
(further explanation in the text). Disoriented dances are indicated with
shaded triangles on the bottom axis. The control bees A performed no bimodal
or disoriented dances. Other symbols and conventions as in
Fig. 2.
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Fig. 4. Dance indications of control bees native to the training treeline (A) and
experimental bees native to the valley site (B) under overcast skies at the
test site on 16 August 2005. The two bees that danced the most are indicated
by open and shaded circles in each panel; these two bees happened to dance in
different directions in each case. All other symbols and conventions as in
Fig. 3.
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© The Company of Biologists Ltd 2008