Interactions between sea urchins and macroalgae in south-western Australia: testing general predictions in a local context

Generalist herbivores profoundly influence the biomass and species composition of macroalgae assemblages. In subtidal ecosystems of temperate latitudes, large invertebrates are usually the most influential herbivores. I tested the prediction that exclusion of invertebrate herbivores would lead to changes in the biomass and species composition of the macroalgae assemblages that are a prominent feature of the reefs in south-western Australia. The most abundant invertebrate herbivores were sea urchins (Heliocidaris erythrogramma, Phyllacanthus irregularis and Centrostephanus tenuispinus), and these occupied different trophic positions. Heliocidaris was present at virtually all reefs surveyed, and was particularly abundant in the Fremantle region. Analyses of stable isotopes and direct observations of gut contents revealed that it was almost exclusively herbivorous, and that it mainly ate foliose brown algae. In contrast, Phyllacanthus and Centrostephanus were omnivorous; while they consumed large proportions of algae, a substantial proportion of the diet of both species was animal tissue. Because Heliocidaris is a generalist herbivore that occurs at high densities, it could exert a large influence on the macroalgae assemblage. This prediction was tested by a series of press experiments. Contrary to the prediction, Heliocidaris exerted a very minor influence on the biomass, and no detectable influence on the species composition, of attached macroalgae. However, it exerted a major influence on the retention of drift macroalgae and seagrass by trapping and feeding on drift. It exerted a particularly strong influence on retention of the kelp Ecklonia radiata. This kelp was not abundant in the attached algae assemblage (when all plots were pooled it ranked 35th in biomass), but was abundant as drift (ranking 1st). Most of the drift Ecklonia was retained by sea urchins, rather than freely drifting.