The depth of the honeybee's backup sun-compass systems.

Honeybees have at least three compass mechanisms: a magnetic compass; a celestial or sun compass, based on the daily rotation of the sun and sun-linked skylight patterns; and a backup celestial compass based on a memory of the sun's movements over time in relation to the landscape. The interactions of these compass systems have yet to be fully elucidated, but the celestial compass is primary in most contexts, the magnetic compass is a backup in certain contexts, and the bees' memory of the sun's course in relation to the landscape is a backup system for cloudy days. Here we ask whether bees have any further compass systems, for example a memory of the sun's movements over time in relation to the magnetic field. To test this, we challenged bees to locate the sun when their known celestial compass systems were unavailable, that is, under overcast skies in unfamiliar landscapes. We measured the bees' knowledge of the sun's location by observing their waggle dances, by which foragers indicate the directions toward food sources in relation to the sun's compass bearing. We found that bees have no celestial compass systems beyond those already known: under overcast skies in unfamiliar landscapes, bees attempt to use their landscape-based backup system to locate the sun, matching the landscapes or skylines at the test sites with those at their natal sites as best they can, even if the matches are poor and yield weak or inconsistent orientation.