Continental diatom biodiversity in stream benthos declines as more nutrients become limiting.

Biodiversity of both terrestrial ecosystems and lacustrine phytoplankton increases with niche dimensionality, which can be determined by the number of limiting resources (NLR) in the environment. In the present continental study, I tested whether niche dimensionality and, with this species, richness scale positively with NLR in running waters. Diatom richness in 2,426 benthic and 383 planktonic communities from 760 and 127 distinct localities, respectively, was examined as a function of NLR, including basic cations, silica, iron, ammonia, nitrate, and dissolved phosphorus. The patterns found in the two communities were opposite: as more resources became limiting, diatom richness declined in the benthos but increased in the phytoplankton. The divergence of benthic from both planktonic and terrestrial communities is attributed to the complex spatial organization of the benthos, generating strong internal resource gradients. Differential stress tolerance among benthic diatoms allows substantial overgrowth, which greatly reduces nutrient transport to the biofilm base and can be supported only by high ambient resource levels. Therefore, niche dimensionality in the benthos increases with the number of resources at high supply. These findings provide a mechanistic explanation of the well documented phenomenon of increased species richness after fertilization in freshwater as opposed to terrestrial ecosystems. Clearly, however, new theoretical approaches, retaining resource availability as an environmental constraint but incorporating a trade-off between tolerance and spatial positioning, are necessary to address coexistence in one of the major producer communities in streams, the algae.