Even in nitrogen-replete ecosystems, microhabitats exist where local-scale nutrient limitation occurs. For example, coastal waters of the northeastern Pacific Ocean are characterized by high nitrate concentrations associated with upwelling. However, macroalgae living in high-zone tide pools on adjacent rocky shores are isolated from this upwelled nitrate for extended periods of time, leading to nutrient limitation. When high-intertidal pools are isolated during low tide, invertebrate-excreted ammonium accumulates, providing a potential nitrogen source for macroalgae. I quantified the influence of mussels (Mytilus californianus Conrad) on ammonium accumulation rates in tide pools. I then evaluated the effects of ammonium loading by mussels on nitrogen assimilation and growth rates of Odonthalia floccosa (Esp.) Falkenb., a common red algal inhabitant of pools on northeastern Pacific rocky shores. Odonthalia was grown in artificial tide pool mesocosms in the presence and absence of mussels. Mesocosms were subjected to a simulated tidal cycle mimicking emersion and immersion patterns of high-intertidal pools on the central Oregon coast. In the presence of mussels, ammonium accumulated more quickly in the mesocosms, resulting in increased rates of nitrogen assimilation into algal tissues. These increased nitrogen assimilation rates were primarily associated with higher growth rates. In mesocosms containing mussels, Odonthalia individuals added 41% more biomass than in mesocosms without mussels. This direct positive effect of mussels on macroalgal biomass represents an often overlooked interaction between macroalgae and invertebrates. In nutrient-limited microhabitats, such as high-intertidal pools, invertebrate-excreted ammonium is likely an important local-scale contributor to macroalgal productivity.

Many ecological phenomena are characterized by context dependency, and the relationship between diversity and productivity is no exception. We examined the relationship between macroalgal diversity,and nutrient availability by evaluating the effects of reduced nutrients and their subsequent replacement via local-scale nutrient loading in tide pools. Macroalgae in Oregon coast high-intertidal pools have evolved in a nitrate-rich upwelling ecosystem, but instead of settling on low-intertidal reefs (where algae are often immersed in nutrient-rich nearshore waters) these individuals have colonized high-zone pools, where they are isolated from the ocean for extended periods of time and are subjected to extended periods, of nitrate depletion. In some pools, this nutrient stress was ameliorated by a positive interaction: the excretion of ammonium by invertebrates. We conducted experimental manipulations to quantify invertebrate-mediated ammonium loading and macroalgal ammonium uptake in high-intertidal pools. Variation in tide pool volume and invertebrate biomass created a gradient of local-scale nutrient inputs, allowing us to address the relationship between nitrogen loading and algal diversity. Slow-growing species tolerant of low nitrogen availability were joined by fast-growing species with higher nitrogen requirements in pools with higher ammonium loading rates. A fourfold increase in the ammonium loading rate was associated with a doubling in the number of macroalgal species, and macroalgal assemblages in more species-rich pools were characterized by higher rates of biomass-specific ammonium uptake. These patterns contrast with productivity-diversity relationships in terrestrial systems, where local-scale nutrient enrichments generally result in reduced producer diversity due to displacement of subordinate species by aggressive competitors. Our data suggest that the effect of enrichment on diversity is context-dependent. Each ecosystem has a critical level of nutrient availability, determined by the level of nutrients typically available in that system. Below this critical level, local-scale nutrient additions increase diversity, but above it, diversity declines with enrichment.

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.