Complex interactions of climatic and ecological controls on macroalgal recruitment

Little is known about the cumulative effects of multiple (.2) environmental controls on species performance and interactions in aquatic ecosystems. We asked how changes in climatic (temperature, ultraviolet radiation) and ecological controls (nutrients, grazing) affect recruitment of the green macroalga Enteromorpha intestinalis, which forms destructive algal blooms in coastal ecosystems worldwide. We designed factorial laboratory experiments to analyze the recruitment response to (1) single and combined effects of nutrient enrichment, grazing pressure, and grazer species composition and (2) the cumulative effects of ultraviolet (UV) radiation, temperature, nutrients, and grazing. Recruitment of E. intestinalis increased exponentially with nutrient enrichment. Grazers could control algal recruitment until a nutrient threshold was reached depending on grazer species composition. Snails (Littorina lit-torea) had strong negative effects on recruit density, whereas amphipods (Gammarus oceanicus) had weak grazing effects and favored algal recruitment through excretion when nutrient supply was low. Temperature and nutrients both enhanced algal recruitment but also the effects of grazers, which led to a significant three-way interaction among these factors. Similarly, effects of UV radiation depended on grazer presence and temperature. When grazers were absent, UV radiation reduced recruitment at 11 and 178C but enhanced recruitment at 58C. No effects were seen in the presence of grazers. Our results indicate that multiple human influences, such as climate change