Interactions between substratum rugosity, colonization density and periwinkle grazing efficiency

Both surface texture and littorinid grazing are known to influence the establishment of many shallow-water benthic hard-bottom communities. However, the effects of these factors and their interactions have not yet been investigated in a quantifiable manner. This investigation aims to assess the interactive effect of both factors in a strictly standardized manner. Natural recruitment by diatoms, the barnacle Balanus improvisus and the tube-building polychaete Polydora sp. was monitored under a 2-factorial treatment: grazing by the periwinkle Littorina littorea (Factor 1; 2 levels: 1 or no snails per plate) on artificial recruitment plates of different initial surface rugosities (Factor 2; 5 levels: smooth, 0.1, 0.5, 1 and 5 mm rugosity elements). In the absence of grazers, barnacle recruitment decreased with increasing initial rugosity, polychaete recruitment peaked at intermediate rugosities, and diatoms exhibited contrasting recruitment patterns in an in vitro and an in situ experiment. When preferred recruitment sites coincided for Polydora and B. improvisus, a competition for space could be inferred from a negative correlation between the 2 species. However, when the overlap of requirements weakened on the 5 rugosities, the relationship was positive, but was not statistically significant. Grazing efficiency by L. littorea depended on initial rugosity, generally showing minimum values on intermediate rugosities which is attributable to a mismatch between radula dimensions and surface structures in these rugosity classes. Additionally, grazing effects tended to increase with higher prey densities. As all factors—initial rugosity, grazing, colonizer species—interact with each other, the outcome of recruitment under combined factors is difficult to predict from single factor effects.