Spatial flows and the regulation of ecosystems.

Spatial flows of materials and organisms across ecosystem boundaries are ubiquitous. Understanding the consequences of these flows should be a basic goal of ecosystem science, and yet it has received scant theoretical treatment to date. Here, using a simple, open, nutrient-limited ecosystem model with trophic interactions, we explore theoretically how spatial flows affect the functioning of local ecosystems, how physical mass-balance constraints interact with biological demographic constraints in the regulation of this functioning, and how failure to consider these constraints explicitly can lead to models that are ecologically inconsistent. In particular, we show that standard prey-dependent models for trophic interactions may lead to implausible outcomes when embedded in an ecosystem context with appropriate mass flows and mass-balance constraints. Our analysis emphasizes the need for integration of population, community, and ecosystem perspectives in ecology and the critical consequences of assuming closed versus open systems.