The consequences of declining biodiversity remain controversial, in part because many studies focus on a single metric of ecosystem functioning and fail to consider diversity's integrated effects on multiple ecosystem functions. We used tide pool microcosms as a model system to show that different conclusions about the potential effects of producer diversity on ecosystem functioning may result when ecosystem functions are measured separately vs. together. Specifically, we found that in diverse seaweed assemblages, uptake of either nitrate or ammonium alone was equal to the average of the component monocultures. However, when nitrate and ammonium were available simultaneously, uptake by diverse assemblages was 22% greater than the monoculture average because different species were complementary in their use of different nitrogen forms. Our results suggest that when individual species have dominant effects on particular ecosystem processes (i.e., the sampling effect), multivariate complementarity can arise if different species dominate different processes. Further, these results suggest that similar mechanisms ( complementary nutrient uptake) may underlie diversity - functioning relationships in both algal and vascular-plant-based systems.

Although nutrient availability and oceanographic context can influence top-down processes in intertidal communities, the reciprocal effects of consumers on bottom-up processes, such as nitrogen uptake by macroalgae, are less well-documented. We used a combination of field observations, mesocosm experiments, and laboratory nitrate-uptake measurements to evaluate the interaction between the kelp crab Pugettia producta and its preferred food source, the kelp Egregia menziesii, in northern California, USA. We found that R producta fed selectively on E. menziesii, removing tissues with a high surface area to volume ratio (SA:V), which resulted in a 66% decrease in overall SA:V. Because these high SA:V tissues are disproportionately more responsible for nitrate uptake, selective herbivory by P. producta decreased E. menziesii's biomass-specific uptake of nitrogen, the primary limiting nutrient in this ecosystem, by 65% per gram of remaining algal tissue. In field surveys, where P. producta abundances were high, E. menziesii thalli more frequently exhibited signs of intense grazing and thus had lower SA:V ratios, suggesting that grazing is sufficiently intense in the field to generate differences in algal morphology and nitrate uptake. P. producta therefore influences E. menziesii biomass and productivity from the top down, by removing substantial amounts of biomass, and from the bottom up, by reducing the kelp's subsequent ability to acquire nutrients. Because of selective grazing on specialized tissues, the effects of mesoherbivores, like P. producta, on the structure and dynamics of marine communities may often be greater than predicted simply by the biomass they consume.