“…Some of them, like celestial and astronomical cues, are of a general nature, whereas others are local. Among location-independent compass cues, many species of sandhoppers possess sun, moon and magnetic compasses (see Pardi and Ercolini, 1986;Ugolini, 2001a;Ugolini, 2001b;Ugolini et al, 2002;Ugolini, 2003). Local cues are also important: through learning, sandhoppers can modify the innate compass reference direction to improve their orientation along the sea-land axis direction or when they are displaced to a differently orientated beach (Ugolini and Macchi, 1988;.…”