Behaviour of mobile macrofauna is a key factor in beach ecology as response to rapid environmental changes

Sandy beach animals show behavioural adaptations that are expressed as contingencies during the life history of individuals to face periodic and episodic environmental changes. Such adaptations include activity rhythms, orientation, zonation, burrowing, escape responses and feeding strategies, the first two being common adaptations to all mobile animals. The complex conditions of a particular beach environment may be integrated in a learning process enhancing the adaptation and survival of individuals and eventually of populations. Evidence exists of genetic determination of some behavioural features that are adaptive in the long term (throughout generations) by increasing individual survival and reproductive potential. The environmental features integrated with the life history of beach animals shape the individual behaviour through ontogenetic processes, as well as population behaviour through evolutionary processes. Thus, behavioural differences among individuals may reflect environmental variation at the local and small/medium temporal scales of beach processes, whereas within-population behavioural coherence and differences among populations may reflect variation at the geographic scale. The different foci stressed by different authors and the variety of evidence dependent upon local geographical and ecological conditions have often resulted in compartmentalised explanations, making generalizations and the repeatability of behavioural studies of beach ecology challenging. There was a need to developing a more synthetic paradigm for beach animal behaviour. This paper gives a brief overview of the theoretical background and keystone studies, which have contributed to our understanding of animal behaviour in sandy beach ecology, and proposes testable hypotheses to be integrated in the beach ecology paradigm.