Top-down versus bottom-up: Grazing and upwelling regime alter patterns of primary productivity in a warm-temperate system.

Community structure is driven by the interaction of physical processes and biological interactions that can vary across environmental gradients and the strength of top-down control is expected to vary along gradients of primary productivity. In coastal marine systems, upwelling drives regional resource availability through the bottom-up effect of nutrient subsidies. This alters rates of primary production and is expected to alter algae-herbivore interactions in rocky intertidal habitats. Despite the potential for upwelling to alter these interactions, the interaction of upwelling and grazing pressure is poorly understood, particularly for warm-temperate systems. Using in situ herbivore exclusion experiments replicated across multiple upwelling regimes, we investigated the effects of both grazing pressure, upwelling, and their interactions on the sessile invertebrate community and biomass of macroalgal communities in a warm-temperate system. The sessile invertebrate cover showed indirect effects of grazing, being consistently low where algal biomass was high at upwelling sites and at nonupwelling sites when grazers were excluded. The macroalgal cover was greater at upwelling sites when grazers were excluded and there was a strong effect of succession throughout the experimental period. Grazing effects were greater at upwelling sites, particularly during winter months. There was a nonsignificant trend toward greater grazing pressure on early than later successional stages. Our results show that the positive bottom-up effects of nutrient supply on algal production do not overwhelm top-down control in this warm-temperate system but do have knock-on consequences for invertebrates that compete with macroalgae for space. We speculate that global increases in air and sea surface temperatures in warm-temperate systems will promote top-down effects in upwelling regions by increasing herbivore metabolic and growth rates.