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To test the relative importance of long-term and working spatial memories in short-range foraging in bumblebees, we compared the performance of two groups of bees. One group foraged in a stable array of six flowers for 40 foraging bouts, thereby enabling it to establish a long-term memory of the array, and adjust its spatial movements accordingly. The other group was faced with an array that changed between (but not within) foraging bouts, and thus had only access to a working memory of the flowers that had been visited. Bees in the stable array started out sampling a variety of routes, but their tendency to visit flowers in a repeatable, stable order ("traplining") increased drastically with experience. These bees used shorter routes and converged on four popular paths. However, these routes were mainly formed through linking pairs of flowers by near-neighbour movements, rather than attempting to minimize overall travel distance. Individuals had variations to a primary sequence, where some bees used a major sequence most often, followed by a minor less used route, and others used two different routes with equal frequency. Even though bees foraging in the spatially randomized array had access to both spatial working memory and scent marks, this manipulation greatly disrupted foraging efficiency, mainly via an increase in revisitation to previously emptied flowers and substantially longer search times. Hence, a stable reference frame greatly improves foraging even for bees in relatively small arrays of flowers.

Chemicals used in communication are divided into signals and cues. Signals are moulded by natural selection to carry specific meanings in specific contexts. Cues, on the other hand, have not been moulded by natural selection to carry specific information for intended receivers. Distinguishing between these two modes of information transfer is difficult when animals do not perform obvious secretion behaviours. Although a number of insects have been suspected of leaving cues at food sites and nest entrances, studies have not attempted to experimentally distinguish between cues and signals. Here, we examine the chemical composition of the scent marks left by the bumblebee Bombus terrestris at food sites and compare it to those found at a neutral location. If bees are depositing a cue, we expect the same chemicals to be found at both sites, but if they deposit a signal we only expect to find the scent marks at the food site. We were also interested in identifying the chemicals left at the nest entrance to determine if they differed from those used to mark food sites. We find that bees deposit the same chemicals at food, nest and neutral sites. Therefore, bumblebees leave behind general chemical footprints everywhere they walk and we propose that they learn to use these footprints in a manner that ultimately enhances their fitness, for example, to improve their foraging efficiency and locate their nest.Experimentally, distinguishing these two modes of information transfer is crucial for understanding how they interact to shape animal behaviour and what chemical bouquets are under natural selection.

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