Large scale patterns and trophic structure of southern African rocky shores: The roles of geographic variation and wave exposure

In this study we revise the biogeographic delimitation, and large-scale patterns of community structure of the intertidal rocky shores of southern Africa. We use binary (presence/absence) and per-species biomass data collected at fifteen localities and thirty-seven different rocky sites, encompassing the shores of southern Namibia, South Africa and southern Mozambique. Multivariate analyses revealed that the shores of southern Africa (south of 25 degrees) can be divided into three main biogeographic provinces: the west coast or Namaqua province, the south coast or Agulhas province and the east coast or Natal province. The biomass structure of the intertidal rocky shores communities of southern Africa varied at a large scale, corresponding to biogeographic differences, while local-scale variation accorded with the intensity of local wave action. The average biomass of west coast communities was on average significantly greater than that of the south and east provinces. At a local scale, the community biomass on exposed shores was an order of magnitude greater than on sheltered shores, within all biogeographic provinces. Semi-exposed shores exhibited intermediate average biomass. The trophic structure of these communities varied significantly with wave action: autotrophs, filter-feeders and invertebrate predators were more prevalent on wave exposed than sheltered shores, whereas grazers were more abundant on sheltered and semi-exposed shores. Exposed shores were consistently dominated by far fewer species than semi-exposed and sheltered shores, independently of biogeographic differences. Within all biogeographic provinces semi-exposed and sheltered shores were more diverse than exposed shores. West coast intertidal communities therefore had high levels of biomass, but were consistently species-poor. Several working hypotheses that could explain these large and small-scale patterns are presented.