Barnacles, limpets and periwinkles: the effects of direct and indirect interactions on cyprid settlement and success

Conventionally, direct interactions between species are considered to be the most important biological factors determining community composition, structure and stability. However, it has been suggested that the indirect interactions occurring between species may be as important. One area of ecology where the direct effects of one species on another have been well studied is in the rocky intertidal. Examination of the effect of the presence of P. vulgata (limpets) and L. littorea (periwinkles) on the settlement and development of S. balanoides (cyprids/barnacles), over a cyprid settlement season and some six months later, in four different treatments (limpets only, limpets and periwinkles combined, periwinkles only and control (no animals)) revealed the following: (1) that the presence of limpets increased cyprid settlement and recruitment success above treatments containing no limpets; (2) that cyprid settlement and success were greatest on the limpets-only treatment, followed by the limpets-and-periwinkles treatment, then by the control treatment and then by the periwinkles-only treatment; (3) that the initial effects observed in the treatments were reflected in the long-term community structure; (4) that the effects of the treatments were independent of variations in algal biomass between treatments, i.e. the effects were not indirectly mediated through a second species (host); (5) that cyprid mortality was greatest on the periwinkles-only treatment; (6) that the source of the effect of limpets on cyprid settlement appeared to originate indirectly through the action of their residual pedal mucus trails. concluded that periwinkles can affect the settlement and success of barnacles directly through biological disturbance (i.e. surface ablation). However, although limpets may have a direct negative effect on barnacle settlement and success, at low to medium densities, limpets can positively indirectly influence the cyprid settlement and success. This effect operates at a factor greater than that afforded by the direct negative effects of periwinkles in a mixed-species treatment. These results illustrate how the indirect effects of one species on another can have a more important structuring effect than those derived from direct effects alone.