Direct and indirect effects of giant kelp determine benthic community structure and dynamics

Indirect facilitation can occur when a species positively affects another via the suppression of a shared competitor. In giant kelp forests, shade from the canopy of the giant kelp, Macrocystis pyrifera, negatively affects understory algae, which compete with sessile invertebrates for space. This raises the possibility that giant kelp indirectly facilitates sessile invertebrates, via suppression of understory algae. We evaluated the effect of giant kelp on the relative abundance of algae and invertebrates by experimentally manipulating kelp abundance on large artificial reefs located off San Clemente, California, USA. The experiments revealed a negative effect of giant kelp on both light availability and understory algal abundance and a positive effect on the abundance of sessile invertebrates, which was consistent with an indirect effect mediated by shade from the kelp canopy. The importance of these processes to temporal variability in benthic community structure was evaluated at 16 locations on natural reefs off Santa Barbara, California, over an eight-year period. Interannual variability in the abundance of understory algae and in the abundance of sessile invertebrates was significantly and positively related to interannual variability in the abundance of giant kelp. Analysis of these observational data using Structural Equation Modeling (SEM) indicated that the magnitude of the indirect effect of giant kelp on invertebrates was six times larger than the direct effect on invertebrates. Results suggest that the dynamics of this system are driven by variability in the abundance of a single structure-forming species that has indirect positive, as well as direct negative, effects on associated species.