Does Imprecision in The Waggle Dance Fit Patterns Predicted by The Tuned-Error Hypothesis?
David A. Tanner
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P. Kirk Visscher
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P. K. Visscher Entomology Department, University of California
, Riverside, Riverside,
CA 92521, USA
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) Biology Department, Utah State University
, 5305 Old Main Hill, Logan,
UT 84322, USA
The waggle dance of the honey bee is used to recruit nest mates to a resource, though direction indicated for a resource may vary greatly within a single dance. Some authors suggest that this variation exits as an adaptation to distribute recruits across a patch of flowers, and that, due to the variation's inverse relationship with distance, the shape of the recruit distribution will remain constant for resources at different distances. In this study, we test this hypothesis by examining how variation in the indication of direction and distance changes with respect to distance. We find that imprecision in the communication of direction does not diminish rapidly enough to accommodate an adaptive-error hypothesis, and we also find that variation in the indication of distance has a positive relationship with the distance of a resource from the hive.
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Evolution is the biological process primarily responsible for the rich diversity of
Earths biota and is accomplished most prominently through natural selection
(Darwin 1859). The potential effect of selective pressures on any organism, and the
form and function that a trait may realize, can be limited, however, by the current
state and evolutionary history of that trait (Gould and Lewontin 1979). Occasionally,
students of evolution atomize an individual into discrete and independent traits
without considering how traits interrelate and influence the evolution of the entire
organism. This tendency may lead to ingenious or fanciful explanations of the
adaptive nature of a trait, and also to the misapprehension that all character states
in nature are products of natural selection and exist as optima (Gould and
Lewontin 1979).
Of recent interest is the utility of imprecision in the waggle dance of the honey
bee. The dance is an adaptation for social recruitment (Sherman and Visscher 2002;
Dornhaus and Chittka 2004; Dornhaus et al. 2006) and is used to communicate the
location of that resource relative to the position of the sun (von Frisch 1967).
The dance works as a ritualized reenactment of the foraging flight, in which the
direction flown to a resource, relative to the suns azimuth, is indicated by the
body orientation and movement of the dancing bee relative to up on a vertical
comb (Fig. 1). This phase of the waggle dance is called a waggle phase. After
completing a waggle phase, a dancer returns to its origin via a return phase to
perform another waggle phase. Distance is communicated through sound bursts
produced during the waggle phase of the dance, the duration of which correlates to
the distance of a resource (Wenner 1962; Esch and Kerr 1965; von Frisch 1967).
There is considerable variation, however, in the direction communicated within a
single dance (Haldane and Spurway 1954; Wilson 1962; von Frisch 1967; Edrich
1975; Gould 1975; Towne 1985; Towne and Gould 1988; Weidenmller and Seeley
1999). Though some of this error may be mitigated by the manner in which
information from the dance is processed (Tanner and Visscher 2008). The angular
difference in the direction between consecutive waggle phases is the divergence
angle. Interestingly, the magnitude of the divergence angle has a negative
relationship with the distance of a resource from the hive. Some authors contend
that the divergence angle has evolved as an adaptation in temperate honey bees to
distribute recruits across a resource patch (Wilson 1962; Towne and Gould 1988),
and that the relationship between divergence angle and the distance of a resource
from the hive represents a tuning to produce a recruit distribution of optimal size that
remains constant with distance (Towne 1985; Towne and Gould 1988; Weidenmller
Fig. 1 Waggle dance of the honey bee. In this figure, the waggle phase is oriented 45 to the left of up
in the hive, communicating that a resource is located 45 to the left of the suns azimuth. Upon completing
this waggle phase, the dancer bee will return to its origin via a return phase, and perform another waggle
phase that may indicate a different direction.
and Seeley 1999). A recent study of recruit distributions, however, showed that, at a
distance of 250 m from the hive, the distribution of bees was significantly smaller
than the imprecision in the dance would predict (Tanner and Visscher 2008). In this
study, we test if the imprecision in the indication of distance and direction are
sufficient to produce a distribution of recruits that will have the same spatial
dimensions as a resource becomes increasingly distant. For a distribution to remain
constant with distance, not only must the imprecision in direction decrease, but also
the imprecision in distance communication must remain constant. An increase or
decrease in variation will alter the size o (...truncated)