Vicious circle in the intertidal: Facilitation between barnacle epibionts, a shell boring polychaete and trematode parasites in the periwinkle Littorina littorea

We studied interactions between three organisms associated with a common gastropod of northern Atlantic shores, the periwinkle Littorina littorea: barnacle epibionts Balanus crenatus, a shell boring polychaete Polydora ciliata, and tissue invading trematodes which use the periwinkles as first intermediate host. Snails collected shortly after barnacle settlement with > 50% cover of barnacles had significantly higher infestation of shell boring worms compared to unfouled snails, while trematode infestation was similar. The result was the same at two sites, and we conclude that the worm P. ciliata facilitates barnacle fouling on snails. The reverse was also the case. In an experiment with 14 weeks of exposure, snail treatments with barnacle epibionts had a significantly higher P. ciliata load than unfouled and cleaned snails. Again, trematode infestations were similar. The reciprocal positive interactions between barnacle epibionts and shell boring worms on snail houses is regarded as a case of facultative mutualism. On the other hand, for the snail basibiont, both barnacles and shell boring worms exert strongly negative effects by reducing fecundity, growth, and survival, resulting in a vicious circle for the snails. The combined effects of these associated organisms may rival in importance any competitive or predacious effects on the host L. littorea.