Scale-Dependent Community Theory for Streams and Other Linear Habitats

The maintenance of species diversity occurs at the regional scale but depends on interacting processes at the full range of lower scales. Although there is a long history of study of regional diversity as an emergent property, analyses of fully multiscale dynamics are rare. Here, we use scale transition theory for a quantitative analysis of multiscale diversity maintenance with continuous scales of dispersal and environmental variation in space and time. We develop our analysis with a model of a linear habitat, applicable to streams or coastlines, to provide a theoretical foundation for the long-standing interest in environmental variation and dispersal, including downstream drift. We find that the strength of regional coexistence is strongest when local densities and local environmental conditions are strongly correlated. Increasing dispersal and shortening environmental correlations weaken the strength of coexistence regionally and shift the dominant coexistence mechanism from fitness-density covariance to the spatial storage effect, while increasing local diversity. Analysis of the physical and biological determinants of these mechanisms improves understanding of traditional concepts of environmental filters, mass effects, and species sorting. Our results highlight the limitations of the binary distinction between local communities and a species pool and emphasize species coexistence as a problem of multiple scales in space and time.