Resonance in physiologically structured population models

Ecologists have long sought to understand how the dynamics of natural populations are affected by the environmental variation those populations experience. A transfer function is a useful tool for this purpose, as it uses linearization theory to show the frequency spectrum of the fluctuations in a population’s abundance relates to the frequency spectrum of environmental variation. Here, we show how to derive and to compute the transfer function for a continuous-time model of populations that are structured by a continuous individual-level state variable such as size. To illustrate, we derive, compute, and analyze the transfer function for a size-structured population model of stony corals with open recruitment, parameterized for a common Indo-Pacific coral species complex. This analysis identifies a sharp, multi-decade resonance driven by space competition between existing coral colonies and incoming recruits. The resonant frequency is most strongly determined by the rate at which colonies grow, and the potential for resonant oscillations is greatest when colony growth is only weakly density-dependent. While these resonant oscillations are unlikely to be predominant dynamical features of degraded reefs, they suggest the dynamical possibilities for marine invertebrates in more pristine waters. The size-structured model that we analyze is a leading example of a broader class of physiologically structured population models, and the methods we present should apply to a wide variety of models in this class.