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.

Species diversity and coexistence have long been central foci of ecology, but field studies are often limited to describing diversity patterns, while theory frequently ignores environmental variation. Scale transition theory is an ideal framework in which to study species diversity, as it explicitly accounts for this environmental variability and allows for the quantification of coexistence mechanisms. Each coexistence mechanism arises from specific types of biotic and abiotic interactions. Moreover, mechanism magnitudes provide information about how these interactions contribute to coexistence. By studying how the natural history of a community determines these biotic and abiotic interactions, insight can be gained into how that natural history influences coexistence. Environmental variation is a central hypothesis for the maintenance of diversity in both desert annual plants and streams. This dissertation is broadly interested in the way differences in the environmental responses of species interact with the structure of the environmental conditions to affect coexistence. I use scale transition theory to develop theoretical understanding of how life history and environmental structure in these communities influence coexistence mechanisms and diversity. In desert annual plants, the focus is on the environmental response itself: how germination depends on environmental conditions. I analyze how this life history interacts with variation in the environment to affect coexistence. The germination responses of desert annual plants to an unstudied type of environmental variation, duration of soil moisture after rainfall, generate species-specific but highly structured patterns of germination variation. Although this germination variation is one-dimensional, the nonlinearities that arise due to germination biology generate sufficient germination variation to promote coexistence by the temporal storage effect. In stream communities, I examine how the physical structure of stream environments affects coexistence given that species’ performance is environmentally dependent. This dissertation demonstrates that patterns of diversity along the stream are related to the strength of coexistence. The downstream drift of organisms has relatively minor effects on coexistence despite asymmetric shifts in the distribution of organism in the stream. This study identifies conditions that eliminate the effects of the branched structure of stream networks on coexistence. Branching has no effect on community dynamics if (a) tributaries have identical environmental conditions, (b) habitat size increases additively at confluences, and (c) demographic stochasticity is unimportant. Any effects of branching on coexistence caused by violating the environmental condition are asymptotically eliminated as streams increase in size. These studies provide a theoretical, mechanistic foundation for the study of stream communities that addresses environmental and life history factors long recognized as important by empirical stream ecologists.