Abstract The task of nectar foraging in honey-bees is
partitioned between foragers and receivers. Foragers
typically transfer a nectar load in the nest as sub-loads to
several receivers rather than as a single transfer. Foragers
experience delays in finding receivers and use these
delays to balance the number of foragers and receivers.
A short delay results in the forager-recruiting waggle dance
whereas a long delay results in the receiver-recruiting
tremble dance. Several nectar transfers increase the cost
of this system by introducing additional delays in finding
extra receivers. We tested four hypotheses to explain the
occurrence of multiple transfer. We found no evidence
that multiple transfer is due to different crop capacities
of foragers and receivers or that it results from extensive
trophallactic interactions with nest-mates. Receiver bees
frequently evaporate nectar in their mouthparts to hasten
the production of honey. The suggestion has been made
that multiple transfer is driven by receivers who take
partial loads from foragers to enhance nectar evaporation.
An alternative suggestion is that foragers drive multiple
transfer to gain better information on the balance of
foragers and receivers. Multiple sampling of the delay in
finding a receiver reduces the standard deviation of the
delay mean and so provides foragers with better informa-
tion than is provided by a single delay. The enhanced-
evaporation hypothesis predicts that receivers break
foragers’ first transfer whereas the information improve-
ment hypothesis predicts foragers break their first trans-
fers. Furthermore, only the information improvement
hypothesis predicts a high level of multiple receptions.
Data on transfer break-off and receiver behaviour strongly
support the information improvement hypothesis and
reject the enhanced-evaporation hypothesis. We suggest
that multiple transfer is an adaptive sampling mechanism,
which improves foragers’ information on colony work
allocation, and that multiple sampling is a common
feature of social insect societies.
Keywords Honey-bee · Nectar transfer · Receiver ·
Foraging · Information improvement
Introduction
Adaptive organisation and integration depend upon
information and communication (Detrain et al. 1999).
Nowhere is this more pertinent than in the organisation
of the activities of the many individuals that comprise an
insect society (Seeley 1995). Foraging is one vital area
of colony life that uses information to co-ordinate indi-
vidual activities. Efficient organisation of foraging
normally involves two complementary processes. One is
directing foragers to profitable food patches, via mecha-
nisms such as tandem running, pheromone trails or the
waggle dance of honey-bees (Wilson 1971; Hölldobler
and Wilson 1990; Seeley 1995). The second is adjusting
the total foraging effort to the colony’s ability to use,
process or store food (Seeley 1995; Anderson and Ratnieks
1999). In the honey-bee, for example, nectar collected by
foragers is transferred to receiver bees, who then store it
in cells. Efficient nectar collection, therefore, requires
both the direction of unemployed foragers to good food
patches and a balance between the work capacities of
nectar foragers and receivers.
Waggle dances made by honey-bee foragers function
in both processes. The dance directs foragers to profitable
food patches and, because the bees recruited were
previously unemployed foragers (Seeley 1995), it also
functions in balancing the work capacities of foragers
and receivers (Anderson and Ratnieks 1999). Clearly, the
waggle dance, and more generally whether or not a
forager should dance, is at the heart of efficient foraging
organisation in the honey-bee (Seeley 1995). As a result,
there should be strong selection making the ability of
individual foragers to dance or not appropriate to the
Communicated by R. Crozier
A.G. Hart (
✉
) · F.L.W. Ratnieks
Department of Animal and Plant Sciences,
University of Sheffield, Sheffield, S10 2TN, UK
e-mail: bop98agh@sheffield.ac.uk
Tel.: +44-114-2220149, Fax: +44-114-2760159
Behav Ecol Sociobiol (2001) 49:244–250
DOI 10.1007/s002650000306
ORIGINAL ARTICLE
Adam G. Hart · Francis L.W. Ratnieks
Why do honey-bee (Apis mellifera) foragers transfer nectar
to several receivers? Information improvement through multiple
sampling in a biological system
Received: 7 February 2000 / Revised: 7 August 2000 / Accepted: 25 September 2000 / Published online: 20 December 2000
© Springer-Verlag 2000