Spatial scale mediates the effects of biodiversity on marine primary producers

Most studies evaluating the effects of biodiversity on ecosystem functioning are conducted at a single location, limiting our understanding of how diversity-function relationships may change when measured across different spatial scales. We conducted a species-removal experiment at three sites nested in each of three regions along the rocky intertidal coastline of the Gulf of Maine, USA, to evaluate the potential for scale-dependent effects of species loss on the biomass of intertidal seaweed assemblages. We randomly assigned 50 plots in the mid-intertidal zone at each site to one of five treatments (n = 10 plots each): an unmanipulated control, a polyculture plot that contained our three target seaweed species, and three monoculture plots. We manipulated diversity by removing all non-target species from monoculture and polyculture plots, then removing additional biomass from polyculture plots, proportionate to species' relative abundances, so that the average amount removed from monoculture and polyculture plots was equivalent at each site. At the largest spatial scale, all sites considered together, after accounting for region and site nested within region seaweed diversity had consistent, positive effects on seaweed cover. Diverse polyculture plots always had higher cover than was predicted by the average performance of the component seaweed species and usually had higher cover than even the best-performing component species. Diversity effects weakened and became less consistent at smaller spatial scales, so that at the scale of individual sites, diverse polycultures only performed better than the average of monocultures ~40% of the time. Hence, our results indicate that weak and/or inconsistent biodiversity effects at the level of individual sites may scale up to stronger, more consistent effects at larger spatial scales. Quantitative summaries of biodiversity experiments conducted at the scale of individual sites do not capture this spatial aspect of biodiversity effects and may therefore underestimate the functional consequences of biodiversity loss.