Insect navigation en route to the goal: multiple strategies for the use of landmarks.

There are at least four distinct ways in which familiar landmarks aid an insect on its trips between nest and foraging site. Recognising scenes: when bees are displaced unexpectedly from their hive to one of several familiar locations, they are able to head in the direction of home as though they had previously linked an appropriate directional vector to a view of the scene at the release site. Biased detours: ants recognise familiar landmarks en route and will correct their path by steering consistently to the left or to the right around them. Aiming at beacons: bees and ants also guide their path by approaching familiar landmarks lying on or close to the direct line between start and finish. Simulations suggest that such mechanisms acting together may suffice to account for the routes taken by desert ants through a landmark-strewn environment: the stereotyped trajectories of individual ants can be modelled by a weighted combination of dead reckoning, biased detours and beacon-aiming. These mechanisms guide an insect sufficiently close to an inconspicuous goal for image matching to be successfully employed to locate it. Insects then move until their current retinal image matches a stored view of the surrounding panorama seen from a vantage point close to the goal. Bees and wasps perform learning flights on their first departure from a site to which they will return. These flights seem to be designed to pick up the information needed for several navigational strategies. Thus, a large portion of the learning flight of a bee leaving a feeder tends to be spent close to the feeder so aiding the acquisition of a view from that vantage point, as is needed for image matching. Bees and social wasps also tend to inspect their surroundings while facing along preferred directions and to adopt similar bearings before landing, thereby making it easy to employ retinotopically stored patterns in image matching. Aiming at beacons, in contrast, requires a landmark to be familiar to the frontal retina. Objects tend to be viewed frontally while the insect circles through arcs centred on the goal. This procedure may help insects to pick out those objects close to the goal that are best suited for guiding later returns.