Behavioral adaptations of sandy beach macrofauna in face of climate change impacts: A conceptual framework

Sandy beaches are severely under-represented in the literature on climate-change ecology, yet different lines of evidence suggest that the macrofauna inhabiting these narrow and dynamic environments located at the land-sea interface is being reorganized under the influence of this large scale and long-lasting stressor. This is reflected in macrofaunal sensitivity to increasing sea surface temperature, sea-level rise, extreme events and erosion of the narrow physical habitat. However, evidence of behavioral responses by sandy beach macrofauna that are consistent with expectations under climate change is scarce and fragmentary. In this paper, specific hypotheses are formulated for how behavioral adaptations in sandy beach macrofauna are predicted to respond to climate change impacts. Firstly, a conceptual framework and an overview of macrofauna behavioral adaptation features are provided. Secondly, the effects of main climate change drivers on sandy beaches are summarized. Thirdly, a conceptual framework is developed giving behavioral adaptations of sandy beach macrofauna under climate change pressure. The degree to which observations on behavioral adaptations of beach animals conform to expectations under specific climate change drivers (sea level rise, sea surface temperature, winds and storminess, rainfall, acidification and eutrophication) is explored. Taking into account the empirical evidence and the theoretical framework detailed in the paper, emergent hypotheses/predictions are proposed. Climate change drivers are expected to impact habitat features and consequently the behavioral expression of macrofauna as active responses to habitat changes. Behavioral adaptations are expected to be impaired, more variable or disrupted, thus decreasing fitness, causing local population extirpations and potentially triggering a range of cascading effects of ecological change in the beach ecosystem. Biodiversity loss will be the outcome of the negative pressures driven by climate change. The specificity of sandy beaches as narrow ecotones between sea and land may be lost under climate change pressure, adversely affecting fine-tuned macrofaunal adaptations and therefore ecosystem functioning. Strictly adapted endemic sandy beach fauna will be especially subjected to local extirpations, while species with a large reaction norm (i.e. phenotypic and behavioral plasticity) may face changes by dispersal and exploitation of new niches. Under climate change impacts, biodiversity loss is predicted, which would hamper beach ecosystem resilience. The limits to which sandy beach macrofauna responds and can behaviorally adapt to environmental change are worthy of exploration, in view of the increasing influence of the long-lasting climate driven stressors threatening these ecosystems at risk.