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Epilithic biofilms play a key role in marine ecosystems. They also provide a tractable system to investigate the relative roles of environmental stressors, bottom-up physicochemical factors and top-down biological control in regulating communities. Patterns of photosynthetic microbial biomass were recorded over a four-year period at several tidal levels on shores in the Isle of Man. Photosynthetic biomass and the abundance of diatoms were consistently greater during winter than summer. Biomass was negatively correlated with insolation stress and air temperature, but was not correlated with grazing intensity, dissolved nutrients, sea temperature, or planktonic chlorophyll. Field experiments confirmed that reducing insolation stress led to substantial increases in photosynthetic biomass, predominantly of diatoms and macroalgal germlings. Reducing grazing intensity also led to considerable increases in photosynthetic biomass, but reducing desiccation stress or increasing nutrient availability had no effect. Although grazing can regulate microalgal biomass, seasonal patterns of grazing activity were driven by temperature and were decoupled from photosynthetic biomass. Our study demonstrates the importance of physiological stresses for the direct and indirect regulation of the balance between primary producers and consumers. Based on these findings, we present a model that combines the roles of stressors together with bottom-up forcing and top-down regulation in controlling communities on wave-exposed shores.

The dominant components of the mid-shore community of 4 sheltered, rocky shores in the south of the Isle of Man, UK, were surveyed. A 2-way factorial experiment to investigate the community structuring roles of Ascophyllum nodosum (hereafter AscophyUum) canopies and Patella vulgata grazing was undertaken at 1 site and monitored over a period of 6 yr. Removal of the canopy had a marked impact on the understorey community, with both direct and indirect effects. In contrast, the effect of limpet removal was limited, owing to the restriction of this grazer to small patches of bare substrate within a turf of red algae. The Ascophyllum canopy directly facilitated the presence of the red algal turf in the mid-shore; canopy removal resulted in bleachmg and death of turf species with a consequent loss of entrapped silt. This degradation of the turf resulted in an increase in the area grazed by limpets and a subsequent increase in limpet recruitment, proportional to the increased area of bare substratum. Thus, the AscophyUwn canopy indirectly limits the population of P. vulgata by facilitating the growth of a red algal turf. Eighteen months after Ascophyllum removal, a mixed canopy of Fucus vesiculosus and Fucus serratus developed. This acted in a similar manner to the original canopy, providlng shade for turf species, which resulted in restoration of the balance between algal turf and limpet grazing. Despite this, the red algal turf had not fully recovered 5 yr after it was originally bleached, illustrating the long-term effects of Ascophyllum canopy loss on this community. Removal of the canopy also resulted in high levels of Ascophyllum recruitment, but the slow growth rate of these juvenile plants meant that nearly 6 yr after canopy removal, an Ascophyllum canopy had still not developed. Nonetheless, we predict that Ascophyllum will eventually outcompete the established Fucus spp. canopy. Detailed exammation of the distribution of Ascophyllum juveniles suggests that the low density of juveniles in the natural population is due to a lack of substratum free from both limpets and space-occupying turf, rather than a direct effect of the canopy.