Climate change is forecasted to increase temperature variability and stochasticity. Most of our understanding of thermal physiology of intertidal organisms has come from laboratory experiments that acclimate organisms to submerged conditions and steady-state increases in temperatures. For organisms experiencing the ebb and flow of tides with unpredictable low tide aerial temperatures, the reliability of reported tolerances and thus predicted responses to climate change requires incorporation of environmental complexity into empirical studies. Using the mussel Mytilus californianus, our study examined how stochasticity of the thermal regime influences physiological performance. Mussels were acclimated to either submerged conditions or a tidal cycle that included either predictable, unpredictable or no thermal stress during daytime low tide. Physiological performance was measured through anaerobic metabolism, energy stores and cellular stress mechanisms just before low tide, and cardiac responses during a thermal ramp. Both air exposure and stochasticity of temperature change were important in determining thermal performance. Glycogen content was highest in the mussels from the unpredictable treatment, but there was no difference in the expression of heat shock proteins between thermal treatments, suggesting that mussels prioritise energy reserves to deal with unpredictable low tide conditions. Mussels exposed to fluctuating thermal regimes had lower gill anaerobic metabolism, which could reflect increased metabolic capacity. Our results suggest that while thermal magnitude plays an important role in shaping physiological performance, other key elements of the intertidal environment complexity such as stochasticity, thermal variability, and thermal history are also important considerations for determining how species will respond to climate warming.

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