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    Local-scale nutrient regeneration facilitates seaweed growth on wave-exposed rocky shores in an upwelling system

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    Abstract This study shows that, even on exposed, wave-swept, rocky shores in a nutrient-replete upwelling ecosystem, mussels (Mytilus californianus) facilitate the growth of the seaweed Porphyra perforata by enhancing nutrient concentrations in the nearby water column. In field surveys on emergent substrate in the mid-intertidal zone, we found ten times greater abundance of P. perforata on mussels than on adjacent rock. In field experiments, P. perforata accumulated and grew more quickly on mussels than on bare rock or on mussel mimics, suggesting that nutrients excreted by mussels might be responsible for greater P. perforata cover. At high tide, water column ammonium concentrations over mussel beds were nearly double those found over bare rock. Correspondingly, tissue nitrogen concentrations were higher, and carbon-to-nitrogen ratios were lower in P. perforata growing on mussels compared to bare rock. Given the dominance of mussels in mid-intertidal regions of temperate coasts worldwide, ammonium regeneration could be a general contributor to local-scale nutrient availability, even in high-flow systems characterized by high nutrient concentrations
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