SUMMARYVisual landmarks guide humans and animals including insects to a goal location. Insects, with their miniature brains, have evolved a simple strategy to find their nests or profitable food sources; they approach a goal by finding a close match between the current view and a memorised retinotopic representation of the landmark constellation around the goal. Recent implementations of such a matching scheme use raw panoramic images ('image matching') and show that it is well suited to work on robots and even in natural environments. However, this matching scheme works only if relevant landmarks can be detected by their contrast and texture. Therefore, we tested how honeybees perform in localising a goal if the landmarks can hardly be distinguished from the background by such cues. We recorded the honeybees' flight behaviour with high-speed cameras and compared the search behaviour with computer simulations. We show that honeybees are able to use landmarks that have the same contrast and texture as the background and suggest that the bees use relative motion cues between the landmark and the background. These cues are generated on the eyes when the bee moves in a characteristic way in the vicinity of the landmarks. This extraordinary navigation performance can be explained by a matching scheme that includes snapshots based on optic flow amplitudes ('optic flow matching'). This new matching scheme provides a robust strategy for navigation, as it depends primarily on the depth structure of the environment.Supplementary material available online at http://jeb.biologists.org/cgi/content/full/213/17/2913/DC1 Key words: honeybee, landmark navigation, snapshot matching, vision.
THE JOURNAL OF EXPERIMENTAL BIOLOGY
2914be unnecessary (Zeil et al., 2003;Stürzl and Zeil, 2007). Zeil et al. show that the similarities between panoramic images of natural environments decrease smoothly with spatial distance between an observer and the goal location (Zeil et al., 2003). An animal that is sensitive to the similarity of views relative to the memorised view of the goal location could return to this location by maximising the similarities between images [modelled by simple image similarity gradient methods (Zeil et al., 2003)]. Thus, panoramic image similarities can be used for view-based homing in natural environments. Recently, the behaviour of ants and crickets in goal-finding tasks could be explained by 'image matching ' (Wystrach and Beugnon, 2009;Mangan and Webb, 2009).In our combined behavioural and modelling approach, we tested the content of the spatial memory in honeybees during complex navigational tasks. Honeybees were trained to locate an inconspicuous feeder surrounded by three cylinders, which we refer to as landmarks. By altering the spatial configuration and landmark texture and monitoring the approach flights to the feeder, we addressed the following questions: what role does the spatial configuration of the landmarks play? Does landmark texture play a role in navigational tasks? In particular, can landmarks b...