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Understanding the effects of climate-mediated environmental variation on the distribution of organisms is critically important in an era of global change. We used wavelet analysis to quantify the spatiotemporal (co)variation in daily water temperature for predicting the distribution of cryptic refugia across 16 intertidal sites that were characterized as ‘no’, ‘weak’ or ‘strong’ upwelling and spanned 2000 km of the European Atlantic Coast. Sites experiencing weak upwelling exhibited high synchrony in temperature but low levels of co-variability at monthly to weekly timescales, whereas the opposite was true for sites experiencing strong upwelling. This suggests upwelling generates temporal thermal refugia that can promote organismal performance by both supplying colder water that mitigates thermal stress during hot Summer months and ensuring high levels of fine-scale variation in temperature that reduce the duration of thermal extremes. Additionally, pairwise correlograms based on the Pearson-product moment correlation coefficient and wavelet coherence revealed scale dependent trends in temperature fluctuations across space, with a rapid decay in strong upwelling sites at monthly and weekly timescales. This suggests upwelling also generates spatial thermal refugia that can ‘rescue’ populations from unfavorable conditions at local and regional scales. Overall, this study highlights the importance of identifying cryptic spatiotemporal refugia thatmore »emerge from fine-scale environmental variation to map potential patterns of organismal performance in a rapidly changing world.

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Abstract The rocky intertidal zone is a highly dynamic and thermally variable ecosystem, where the combined influences of solar radiation, air temperature and topography can lead to differences greater than 15°C over the scale of centimetres during aerial exposure at low tide. For most intertidal organisms this small-scale heterogeneity in microclimates can have enormous influences on survival and physiological performance. However, the potential ecological importance of environmental heterogeneity in determining ecological responses to climate change remains poorly understood. We present a novel framework for generating spatially explicit models of microclimate heterogeneity and patterns of thermal physiology among interacting organisms. We used drone photogrammetry to create a topographic map (digital elevation model) at a resolution of 2 × 2 cm from an intertidal site in Massachusetts, which was then fed into to a model of incident solar radiation based on sky view factor and solar position. These data were in turn used to drive a heat budget model that estimated hourly surface temperatures over the course of a year (2017). Body temperature layers were then converted to thermal performance layers for organisms, using thermal performance curves, creating ‘physiological landscapes’ that display spatially and temporally explicit patterns of ‘microrefugia’. Our framework shows how non-linear interactionsmore »between these layers lead to predictions about organismal performance and survivorship that are distinct from those made using any individual layer (e.g. topography, temperature) alone. We propose a new metric for quantifying the ‘thermal roughness’ of a site (RqT, the root mean square of spatial deviations in temperature), which can be used to quantify spatial and temporal variability in temperature and performance at the site level. These methods facilitate an exploration of the role of micro-topographic variability in driving organismal vulnerability to environmental change using both spatially explicit and frequency-based approaches.« less