ALLEE EFFECTS: POPULATION GROWTH,CRITICAL DENSITY,AND THE CHANCE OF EXTINCTION

This paper develops mathematical models to describe the growth, critical density, and extinction probability in sparse populations experiencing Allee effects. An Allee effect (or depensation) is a situation at low population densities where the per-individual growth rate is an increasing function of population density. A potentially important mechanism causing Allee effects is a shortage of mating encounters in sparse populations. Stochastic models are proposed for predicting the probability of encounter or the frequency of encounter as a function of population density. A negative exponential function is derived as such an encounter function under very general biological assumptions, including random, regular, or aggregated spatial patterns. A rectangular hyperbola function, heretofore used in ecology as the functional response of predator feeding rate to prey density, arises from the negative exponential function when encounter probabilities are assumed heterogeneous among individuals. These encounter functions produce Allee effects when incorporated into population growth models as birth rates. Three types of population models with encounter-limited birth rates are compared: (1) deterministic differential equations, (2) stochastic discrete birth-death processes, and (3) stochastic continuous diffusion processes. The phenomenon of a critical density, a major consequence of Allee effects, manifests itself differently in the different types of models. The critical density is a lower unstable equilibrium in the deterministic differential equation models. For the stochastic discrete birth-death processes considered here, the critical density is an inflection point in the probability of extinction plotted as a function of initial population density. In the continuous diffusion processes, the critical density becomes a local minimum (antimode) in the stationary probability distribution for population density. For both types of stochastic models, a critical density appears as an inflection point in the probability of attaining a small population density (extinction) before attaining a large one. Multiplicative (“environmental”) stochastic noise amplifies Allee effects. Harvesting also amplifies those effects. Though Allee effects are difficult to detect or measure in natural populations, their presence would seriously impact exploitation, management, and preservation of biological resources.