A nonequilibrium framework for community responses to pulse perturbations

Understanding responses of ecological communities to shocks that displace species abundances is of paramount importance given the increasing frequency of extreme climatic events. However, current theory on responses to such pulse perturbations focuses on equilibrium points and we lack a unified framework that accommodates other common, but more complicated, types of population fluctuations such as transients and cycles. Here we introduce this frame-work by deriving metrics that quantify the minimum, typical, and maximum amplification of perturbed abundances under nonequilibrium population dynamics. By simulating models under several nonequilibrium scenarios, we demonstrate that these metrics accurately characterize the full range of amplification of perturbed abundances in the short and long terms. Notably, we show that perturbation amplification depends strongly on community state in the short term, but this state dependency vanishes in the long term. Overall, our framework enables stability analysis for model and natural communities that do not exhibit equilibrium dynamics.