Keynote papers on sandhopper orientation and navigation

This article analyses the relevant studies that have made sandhoppers a model subject for the study of orientation, and traces the development of the paradigm through innovative hypotheses and empirical evidence. Sandhoppers are able to maintain their direction without sensorial contact with the goal, which is their burrowing zone extended along the beach, but very narrow across it. They actively determine the direction of their movements, according to their internal state and the environmental features encountered. Each population shows an 'innate directional tendency' adapted to the shoreline of origin, and the inexpert laboratory-born young behave in a similar way to the adults. Genetic differences have been demonstrated between, as well as within natural populations. The question of the calibration of the sun compass to orientation on a particular shoreline implies a redundancy of mechanisms of orientation. Orientation mechanisms may involve environmental cues perceived through diverse sensory modalities, and range from simple orientation reflexes to sun compass navigational systems. These include scototaxis and geotaxis, and the response to the silhouette of the dune, in addition to sun and moon orientation, which is dependent on the time of the day and orientates daily migrations on the beach. Different modalities of orientation may operate singly, or in conjunction with each other, and their ecological significance may vary according to the habitat and lifestyle of the animals. Taken collectively, the orientation behaviour of the group appears to be a most accommodating phenotype, with considerable adaptive potential. The evidence from comparative studies of different populations promotes consideration of behavioural plasticity as an adaptation to changing coastlines.