Abstract
The strength of top-down control by consumers is predicted to decrease with latitude, but most data confirming this assumption come from latitudes <60°, while empirical studies of predation in sub-arctic and arctic marine habitats are few. A barnacle Balanus crenatus is a native foundation species in the shallow subtidal of the White Sea (65° N), hosting a diverse (250+ species) assemblage of macrobenthic organisms. On mixed sediments live barnacles share primary substrates (shells and gravel) with numerous empty barnacle tests, 7% of which had drill holes of an unidentified origin. We manipulated the densities of (i) adult muricid whelks Boreotrophon clathratus (of previously unknown feeding habits), to check if they prey on barnacles, (ii) other predators to reveal their effect on juvenile Boreotrophon, and (iii) empty tests to assess the community-wide effect of predation on barnacles. The abundance of drilled empty tests in the field correlated with that of Boreotrophon. A year-long caging experiment clearly confirmed predation, showing the highest barnacle mortality and proportion of drilled tests in whelk enclosures, and the lowest--in predator exclosure treatments. Boreotrophon preferred the barnacles attached to conspecifics to those from primary substrates. Because of its scarcity Boreotrophon had a minor direct effect on barnacle abundance in the field. Yet, initially defaunated empty tests and live barnacles developed markedly different macrobenthic assemblages, suggesting a strong indirect effect of the predation. Juvenile Boreotrophon were 5-6 times less abundant in open and partial cages than in exclosures and enclosures, which indicates that the recruitment and, consequently, the abundance of Boreotrophon and its predation on Balanus are top-down controlled by apex predators. In contrast, in tropical and temperate intertidal the predation on barnacles is stronger and primarily limited by environmental stress and prey availability.