Persistence and Space Occupancy by Subtidal Blue Mussel Patches

At a subtidal, soft-bottom site in the western Baltic Sea, mussel (Mytilus edulis) patches co-occur with high predator abundances. Sea star (Asterias rubens) biomasses, in particular, exceed reported values considered sufficient for restricting mussels to the intertidal zone. To determine how mussels can persist in the face of intense predation we decomposed patch space occupancy into the relative contributions of newly arriving individuals (recruitment) and of increases in body size of the individuals already present in the patch over 13 mo. Sea stars, as major predators, were only able to control 77% of the potential per capita recruitment rate of 91 individuals/yr in 2 m depth. The remaining recruitment rate of 21 individuals/yr was sufficient to allow patches to occupy 1.6 times more space per year. Transplantation of patches to 6 m depth, where recruitment is negligible, revealed that sea stars were also ineffective in controlling mussel coverage through consumption of larger mussels ($>$1 yr,$>$30 mm shell length). In deeper water, space occupancy of patches through increases in mussel body size was able to balance predation mortality, demonstrating that mussels attained a relative refuge in size at only 33 mm shell length. Based on the measured shell growth rates, mussels attain this size after ≈ 15 mo. In situ observations of Asterias feeding activity, the ratios between necessary predator sizes to attack prey of a given size, and predator size distributions suggest that sea stars were on average too small to feed effectively on adult ($>$1 yr) mussels. Probably, Asterias cannot respond to abundant prey and increase its maximal body size at the site because salinities are at its lower tolerable limit (12-18 g/kg). Thus, bottom-up factors such as high prey productivity in concert with subtle size-based ineffectiveness of the predator population allow otherwise unstable predator-prey populations of a generalist predator and its preferred prey to coexist. Although mussel predators were unable to decimate mussels to local extinction, a release of experimental mussel patches from predation with strong recruitment (2 m depth) resulted in an approximately sevenfold yearly areal increase in shallow treatments, which would lead to a 100% mussel cover at the site within 1 yr. Given that mussels can dominate both rocky substratum and soft sediment, we also studied the effect of substratum quality in factorial combination with presence/absence of predation and