Abstract. Increasing evidence emphasizes that the effects of human impacts on ecosystems must be investigated using designs that incorporate the responses across levels of biological organization as well as the effects of multiple stressors. Here we implemented a mesocosm experiment to investigate how the effects of CO2 enrichment and its interaction with eutrophication, scale-up from changes in primary producers at the individual- (biochemistry) or population-level (production, reproduction, and/or abundance) to higher levels of community (macroalgae abundance, herbivory, and global metabolism) and ecosystem organization (detritus release and carbon sink capacity). The responses of Zostera noltii seagrass meadows growing in low- and high- nutrient field conditions were compared. In both meadows, the effect of elevated CO2 levels was mediated by epiphyte proliferation (mostly the cyanobacterium Microcoleus spp.), but not through changes in plant biochemistry or population-level traits. In the low-nutrient meadow, epiphyte proliferation suppressed the CO2 benefits on Z. noltii leaf production and led to increased detritus and decreased organic matter in sediment. Faster and stronger responses to nutrients than to CO2 were observed. Nutrient addition enhanced the nutritional quality of Z. noltii (high N, low C : N and phenolics) and the loss of leaves and shoots, while promoted the proliferation of pennate diatoms and purple bacteria. These changes led to a reduced sediment organic matter, but had no significant effects on herbivory nor on community metabolism. Interestingly, the interaction with CO2 attenuated eutrophication effects. In the high-nutrient meadow, a striking shoot decline caused by amphipod overgrazing was observed, with no response to CO2 and nutrient additions. Our results reveal that under future scenarios of CO2, the responses of seagrass ecosystems will be complex, being mediated by epiphyte proliferation rather than by effects on plant biochemistry. The multi-level responses of the system to nutrients overwhelmed those to CO2 enrichment, but the interaction between stressors reduced the effects of eutrophication. Both, CO2 and nutrient enrichments can reduce the carbon sink capacity of seagrass meadows.
Toxic effects of increased sediment nutrient and organic matter loading on the seagrass Zostera noltii
