Dynamics of Reciprocal Pulsed Subsidies in Local and Meta-Ecosystems

Temporally variable and reciprocal subsidies between ecosystems are ubiquitous. These spatial flows can generate a suite of direct and indirect effects in local and meta-ecosystems. The focus of most subsidy research, however, has been on the response of consumers in recipient ecosystems to constant subsidies over very short or very long time scales. We derive a meta-ecosystem model to explicitly consider the dynamic feedbacks between local ecosystems coupled through reciprocal pulsed subsidies. We predict oscillating reinforcing and dampening effects of reciprocal pulsed herbivore flows. Maximum reinforcing effects between reciprocal pulsed herbivore flows occur when these flows are in phase with the dynamics of neighboring predators. This prediction is robust to a range of pulse quantities and frequencies. Reciprocal pulsed herbivore subsidies lead to spatial and temporal variability in the strength of trophic cascades in local and meta-ecosystems but these cascading effects are the strongest when reciprocal pulsed subsidies are temporally concentrated. When predators demonstrate a behavioral response to prey abundance, reciprocal pulsed subsidies dampen the strength of local trophic cascades but lead to strong trophic cascades across local ecosystems. The timing of reciprocal pulsed subsidies is a critical component that determines the cascading effects of spatial flows. We show that spatial and temporal variabilities in resources and consumers can have a significant influence on the strength of cascading trophic interactions; therefore, our ability to detect and understand trophic cascades may depend on the scale of inquiry of ecological studies.