Coexistence and extinction in flow-kick systems: An invasion growth rate approach

Natural populations experience a complex interplay of continuous and discrete processes: continuous growth and interactions are punctuated by discrete reproduction events, dispersal, and external disturbances. These dynamics can be modeled by impulsive or flow-kick systems, where continuous flows alternate with instantaneous discrete changes. To study species persistence in these systems, an invasion growth rate theory is developed for flow-kick models with state-dependent timing of kicks and auxiliary variables that can represent stage structure, trait evolution, or environmental forcing. The invasion growth rates correspond to Lyapunov exponents that characterize the average per-capita growth of species when rare. Two theorems are proven that use invasion growth rates to characterize permanence, a form of robust coexistence where populations remain bounded away from extinction. The first theorem uses Morse decompositions of the extinction set and requires that there exists a species with a positive invasion growth rate for every invariant measure supported on a component of the Morse decomposition. The second theorem uses invasion growth rates to define invasion graphs whose vertices correspond to communities and directed edges to potential invasions. Provided the invasion graph is acyclic, permanence and extinction are fully characterized by the signs of the invasion growth rates. Invasion growth rates are also used to identify the existence of extinction-bound trajectories and attractors that lie on the extinction set. To demonstrate the framework's utility, these results are applied to three ecological systems: (i) a microbial serial transfer model where state-dependent timing enables coexistence through a storage effect, (ii) a spatially structured consumer-resource model showing intermediate reproductive delays can maximize persistence, and (iii) an empirically parameterized Lotka-Volterra model demonstrating how disturbance can lead to extinction by disrupting facilitation. Mathematical challenges, particularly for systems with cyclic invasion graphs, and promising biological applications are discussed. These results reveal how the interplay between continuous and discrete dynamics creates ecological outcomes not found in purely continuous or discrete systems, providing a foundation for predicting population persistence and species coexistence in natural communities subject to gradual and sudden changes.