Ecophysiological responses to heat waves in the marine intertidal zone

One notable consequence of climate change is an increase in the frequency, scale and severity of heat waves. Heat waves in terrestrial habitats (atmospheric heat waves, AHW) and marine habitats (marine heat waves, MHW) have received considerable attention as environmental forces that impact organisms, populations and whole ecosystems. Only one ecosystem, the intertidal zone, experiences both MHWs and AHWs. In this Review, we outline the range of responses that intertidal zone organisms exhibit in response to heat waves. We begin by examining the drivers of thermal maxima in intertidal zone ecosystems. We develop a simple model of intertidal zone daily maximum temperatures based on publicly available tide and solar radiation models, and compare it with logged, under-rock temperature data at an intertidal site. We then summarize experimental and ecological studies of how intertidal zone ecosystems and organisms respond to heat waves across dimensions of biotic response. Additional attention is paid to the impacts of extreme heat on cellular physiology, including oxidative stress responses to thermally induced mitochondrial overdrive and dysfunction. We examine the energetic consequences of these mechanisms and how they shift organismal traits, including growth, reproduction and immune function. We conclude by considering important future directions for improving studies of the impacts of heat waves on intertidal zone organisms.