Extreme heatwave drives topography-dependent patterns of mortality in a bed-forming intertidal barnacle, with implications for associated community structure.

Heatwave frequency and intensity will increase as climate change progresses. Intertidal sessile invertebrates, which often form thermally benign microhabitats for associated species, are vulnerable to thermal stress because they have minimal ability to behaviourally thermoregulate. Understanding what factors influence the mortality of biogenic species and how heatwaves might impact their ability to provide habitat is critical. Here, we characterize the community associated with the thatched barnacle, Semibalanus cariosus (Pallass, 1788), in British Columbia (BC), Canada. Then, we investigate what site-level and plot-level environmental factors explained variations in barnacle mortality resulting from an unprecedented regional heatwave in BC, Canada. Furthermore, we used a manipulative shading experiment deployed prior to the heatwave to examine the effect of thermal stress on barnacle survival and recruitment and the barnacle-associated community. We identified 50 taxa inhabiting S. cariosus beds, with variations in community composition between sites. Site-scale variables and algal canopy cover did not predict S. cariosus mortality, but patch-scale variation in substratum orientation did, with more direct solar irradiance corresponding with higher barnacle mortality. The shading experiment demonstrated that S. cariosus survival, barnacle recruitment, and invertebrate community diversity were higher under shades where substratum temperatures were lower. Associated community composition also differed between shaded and non-shaded plots, suggesting S. cariosus was not able to fully buffer acute thermal stress for its associated community. While habitat provisioning by intertidal foundation species is an important source of biodiversity, these species alone may not be enough to prevent substantial community shifts following extreme heatwaves. As heatwaves become more frequent and severe, they may further reduce diversity via the loss of biogenic habitat, and spatial variation in these impacts may be substantial.