First published online October 21, 2005
Journal of Experimental Biology 208, 4049-4061 (2005)
Published by The Company of Biologists 2005
doi: 10.1242/jeb.01869
Does swarming cause honey bees to update their solar ephemerides?
William F. Towne*,
Christopher M. Baer,
Sarah J. Fabiny and
Lisa M. Shinn
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). White arrows are 200 m long and indicate the bees' outward
flights to the feeders. North is indicated by black arrowheads. The
agricultural fields adjacent to the treelines sloped downward away from the
treelines; adjacent contour lines are separated by 6.1 m of elevation.
Different sites served as donor and recipient sites in different experiments.
The north-facing treeline (A) is at 75°47'12.1'W,
40°37'6.4'N, and the south-facing treeline (B) is at
75°47'12.6'W, 40°36'56.7'N.
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Fig. 2. Dance indications of native and transplanted bees under overcast conditions
on 28 July 2002. 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 south-facing treeline
(Fig. 1B), where the direction
to the feeder, and thus the correct dance indication, was 87.5° clockwise
(CW) of N (Current site, horizontal black line in each panel). The predicted
direction for dances oriented by memory of the sun's course at the
north-facing treeline (Fig.
1A), to which the transplanted bees were native, was 253.5°
(Natal site, horizontal gray line in each panel). (A) Bees native to the
current site; (B) newly transplanted bees released on the day of the
observations; (C) long-term transplantees, released at the current site 2-15
days earlier. Bimodal dances are indicated by broken vertical lines connecting
the two dance directions. The sky bar above each panel indicates sky
conditions: black indicates complete overcast; white indicates that the sun
was visible; shading indicates the sun peeking frequently in and out. There
were no periods of blue sky without sun. The sun first appeared at 8:22 h LST,
indicated on each panel by the thin vertical line running the entire height of
the graph. Statistical analyses here and below exclude dances occurring after
the sun first appeared.
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Fig. 3. Dance indications of native and transplanted bees under overcast on 27 July
2002. (A) Bees native to the current site; (B) newly transplanted bees that
had less than one full day's experience under mostly cloudy skies at the
current site; (C) long-term transplantees. All other symbols and conventions
as in Fig. 2.
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Fig. 4. Dance indications of native and transplanted bees under overcast on 14 July
2002. All symbols and conventions as in Figs
2 and
3.
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Fig. 5. Hovering (A) and circling (B) learning flights of bees transplanted by
three different techniques: caging small groups of foragers (Ai, Bi), moving
the entire hive intact (Aii, Bii), and putting the colonies through a swarming
process as they were transplanted (Aiii, Biii). Note that the scales of the
graphs for the hovering flights (A) and circling flights (B) are different.
Also shown in each panel is the mean flight duration ±
S.D. and the sample size, the latter in parentheses. The
flight durations for Swarms 1 and 3 are indistinguishable (Aiii, Biii; shaded
compared to black), so the results from the two swarms are pooled for
statistical comparisons with the other groups (see Materials and methods).
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Fig. 6. Dance indications of non-swarmed control bees (A) and Swarm 1 bees (B,C)
under cloudy skies. Unlike the previous figures, the three panels are not from
the same day: A and B are from 15 July 2000, while C is from 17 July 2000. Sky
bars and other conventions as in Figs
2,
3,
4, except that shaded regions
on the sky bar indicate periods when blue sky was visible but not the sun.
Note also that the current (recipient) site is now the north-facing treeline,
making the `current site' prediction 253.5°.
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Fig. 7. Dance indications of non-swarmed control bees (A), Swarm 2 bees (B), and
Swarm 3 bees (C), under cloudy skies on 14 August 2000. Sky bar and other
conventions as in Fig. 6. In B,
consecutive dances of the two bees that danced the most are connected by thin
black lines. In C, the dances of the four individual bees that indicated only
the natal site prediction before the sun became visible are labeled with each
bee's identifying tag.
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© The Company of Biologists Ltd 2005