Facilitation cascades occur when multiple foundation species in a community are involved in a hierarchy of positive interactions, and consist of a primary facilitator which positively affects secondary facilitators, each supporting a suit of dependent species. There is no theoretical limit to the number of levels in a facilitation cascade, yet the existence of more than two has rarely been examined. We manipulated biogenic substrate produced by a primary facilitator (cockle shells) and a secondary facilitator (barnacles and their empty tests) in a space-limited subtidal community to test the hypothesis that solitary ascidians would be the third-level facilitator. In the field, most ascidians were found on barnacles, and most barnacles occupied cockle shells. To produce this pattern, barnacles could nurse ascidians (a longer ‘facilitation chain’) or outcompete them from cockle shells (a shorter chain). Experimental results clearly supported the nursing hypothesis providing evidence for a facilitation cascade with three hierarchical levels of foundation species. Our findings confirm that like predation and competition, positive interspecific interactions nest into multi-tier hierarchies with numerous levels. While the number of foundation species should increase community stability and resilience as it increases diversity and reduces environmental stress, facilitation chain length may have the opposite effect.

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.

Foundation species (FS) are strong facilitators providing habitat for numerous dependent organisms. The communities shaped by FS are commonly structured by interplay of facilitation and consumer control. Predators or grazers often indirectly determine community structure eliminating either FS or their principal competitors. Alternatively, they can prey on the dependent taxa directly, which is generally buffered by FS via forming complex habitats with numerous refuges. The latter case has been never investigated at high latitudes, where consumer control is widely considered weak. We manipulated the presence of common epibenthic crustacean predators to assess their effect on mobile macrofauna of the clusters developed by a FS (barnacle Balanus crenatus and its empty tests) in the White Sea shallow subtidal (65° N). While predation pressure on the FS itself here is low, the direct effects of a spider crab Hyas araneus and a shrimp Spirontocaris phippsii on the associated assemblages were unexpectedly strong. Removing the predators did not change species diversity, but tripled total abundance and altered multivariate community structure specifically increasing the numbers of amphipods, isopods (only affected by shrimp), and bivalves. Consumer control in the communities shaped by FS may not strictly follow the latitudinal predation gradient rule.