Distinguishing between direct and indirect effects of predators in complex ecosystems.
Mounting concern over the loss of marine biodiversity has increased the urgency of understanding its consequences. This urgency spurred the publication of many short-term studies, which often report weak effects of diversity (species richness) driven by the presence of key species (the sampling effect). Longer-term field experiments are slowly accumulating, and they more often report strong diversity effects driven by species complementarity, calling into question the generality of earlier findings. However, differences among study systems in which short- and long-term studies are conducted currently limit our ability to assess whether these differences are simply due to biological or environmental differences among systems. In this paper, we compared the effect of intertidal seaweed species richness on biomass accumulation in mesocosms and field experiments using the same pool of species. We found that seaweed species richness increased biomass accumulation in field experiments in both short (2-month) and long (3-year) experiments, although effects were stronger in the long-term experiment. In contrast, richness had no effect in mesocosm experiments, where biomass accumulation was completely a function of species identity. We argue that the short-term experiments, like many published experiments on the topic, detect only a subset of possible mechanisms that operate in the field over the longer term because they lack sufficient environmental heterogeneity to allow expression of niche differences, and they are of insufficient length to capture population-level responses, such as recruitment. Many published experiments, therefore, likely underestimate the strength of diversity on ecosystem processes in natural ecosystems.