First published online October 5, 2006
Journal of Experimental Biology 209, 3984-3989 (2006)
Published by The Company of Biologists 2006
doi: 10.1242/jeb.02472
Interspecific variation in beeswax as a biological construction material
Robert Buchwald1,*,
Michael D. Breed1,
Alan R. Greenberg2 and
Gard Otis3
1 Department of Ecology and Evolutionary Biology and Institute for
Behavioral Genetics, University of Colorado at Boulder, Campus Box 427,
Boulder, CO, 80309-0427, USA
2 Department of Mechanical Engineering, University of Colorado at Boulder,
Campus Box 427, Boulder, CO, 80309-0427, USA
3 Department of Environmental Biology, University of Guelph, Guelph, ON, N0B
2J0, Canada

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Fig. 1. Schematic representation of wax sample before (A) and after (B) a
compression test.
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Fig. 2. Sample output from a wax compression test. Strain is on the abscissa,
calculated as the change in length divided by the original length of the
sample, and has no units. Stress, in MPa, is on the ordinate and is calculated
as the force applied to the sample divided by its cross-sectional area. The
proportional limit is defined as the point where the curve leaves linearity
and the sample moves from elastic to plastic deformation. The yield point is
the point of local maximum stress that the sample can withstand. Resilience is
defined by the area under the curve at the proportional limit and is a measure
of the energy that the sample can absorb before deformation is permanent.
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Fig. 3. Comparisons among the waxes of five honeybee (Apis) subspecies for
six mechanical measures: yield stress (A), yield strain (B), stress at the
proportional limit (C), strain at the proportional limit (D), stiffness (E)
and resilience (F). Bars represent means ± s.e.m. Different letters
indicate statistically different groups.
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Fig. 4. Photograph of a dwarf honeybee nest (A) and giant honeybee nest (B) in
Southern China. Note how the dwarf honeybee nest completely surrounds the
branch to which it is attached instead of hanging from the bottom of the
branch like the giant honeybee nest and those of other honey bee species. Also
note the differences in size; scale bars 10 cm (A), 50 cm (B). Photo: Deng
Xiao-bao.
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© The Company of Biologists Ltd 2006