Golden opportunities: A horizon scan to expand sandy beach ecology

Robust ecological paradigms and theories should, ideally, hold across several ecosystems. Yet, limited testing of generalities has occurred in some habitats despite these habitats offering unique features to make them good model systems for experiments. We contend this is the case for the ocean-exposed sandy beaches. Beaches have several distinctive traits, including extreme malleability of habitats, strong environmental control of biota, intense cross-boundary exchanges, and food webs highly reliant on imported subsidies. Here we sketch broad topical themes and theoretical concepts of general ecology that are particularly well-suited for ecological studies on sandy shores. These span a broad range: the historical legacies and species traits that determine community assemblages; food-web architectures; novel ecosystems; landscape and spatial ecology and animal movements; invasive species dynamics; ecology of disturbances; ecological thresholds and ecosystem resilience; and habitat restoration and recovery. Collectively, these concepts have the potential to shape the outlook for beach ecology and they should also encourage marine ecologists to embrace, via cross-disciplinary ecological research, exposed sandy beach systems that link the oceans with the land.