Convergence of generalized urn models to non-equilibrium attractors

Generalized Polya urn models have been used to model the establishment dynamics of a small founding population consisting of k different genotypes or strategies. As population sizes get large, these population processes are well-approximated by a mean limit ordinary differential equation whose state space is the k simplex. We prove that if this mean limit ODE has an attractor at which the temporal averages of the population growth rate is positive, then there is a positive probability of the population not going extinct (i.e. growing without bound) and its distribution converging to the attractor. Conversely, when the temporal averages of the population growth rate is negative along this attractor, the population distribution does not converge to the attractor. For the stochastic analog of the replicator equations which can exhibit non-equilibrium dynamics, we show that verifying the conditions for convergence and non-convergence reduces to a simple algebraic problem. We also apply these results to selection-mutation dynamics to illustrate convergence to periodic solutions of these population genetics models with positive probability.Comment: 29 pages, 2 figure

Pushed beyond the brink: Allee effects, environmental stochasticity, and extinction

A demographic Allee effect occurs when individual fitness, at low densities, increases with population density. Coupled with environmental fluctuations in demographic rates, Allee effects can have subtle effects on population persistence and extinction. To understand the interplay between these deterministic and stochastic forces, we analyze discrete-time single species models allowing for general forms of density-dependent feedbacks and stochastic fluctuations in demographic rates. Our analysis provide criteria for stochastic persistence, asymptotic extinction, and conditional persistence. Stochastic persistence requires that the geometric mean of fitness at low densities is greater than one. When this geometric mean is less than one, asymptotic extinction occurs with a high probability whenever the initial population density is low. If in addition the population only experiences positive density-dependent feedbacks, conditional persistence occurs provided the geometric mean of fitness at high population densities is greater than one. However, if the population experiences both positive and negative density-dependent feedbacks, conditional persistence is only possible if fluctuations in demographic rates are sufficiently small. Applying our results to stochastic models of mate-limitation, we illustrate counter-intuitively that the environmental fluctuations can increase the probability of persistence when populations are initially at low densities, and decrease the likelihood of persistence when populations are initially at high densities. Alternatively, for stochastic models accounting for predator saturation and negative density-dependence, environmental stochasticity can result in asymptotic extinction at intermediate predation rates despite conditional persistence occurring at higher predation rates.Comment: 19 pages, 3 figure

Robust permanence for ecological equations with internal and external feedbacks.

Species experience both internal feedbacks with endogenous factors such as trait evolution and external feedbacks with exogenous factors such as weather. These feedbacks can play an important role in determining whether populations persist or communities of species coexist. To provide a general mathematical framework for studying these effects, we develop a theorem for coexistence for ecological models accounting for internal and external feedbacks. Specifically, we use average Lyapunov functions and Morse decompositions to develop sufficient and necessary conditions for robust permanence, a form of coexistence robust to large perturbations of the population densities and small structural perturbations of the models. We illustrate how our results can be applied to verify permanence in non-autonomous models, structured population models, including those with frequency-dependent feedbacks, and models of eco-evolutionary dynamics. In these applications, we discuss how our results relate to previous results for models with particular types of feedbacks

Robust permanence for interacting structured populations

The dynamics of interacting structured populations can be modeled by $\frac{dx_i}{dt}= A_i (x)x_i$ where $x_i\in \R^{n_i}$, $x=(x_1,\dots,x_k)$, and $A_i(x)$ are matrices with non-negative off-diagonal entries. These models are permanent if there exists a positive global attractor and are robustly permanent if they remain permanent following perturbations of $A_i(x)$. Necessary and sufficient conditions for robust permanence are derived using dominant Lyapunov exponents $\lambda_i(\mu)$ of the $A_i(x)$ with respect to invariant measures $\mu$. The necessary condition requires $\max_i \lambda_i(\mu)>0$ for all ergodic measures with support in the boundary of the non-negative cone. The sufficient condition requires that the boundary admits a Morse decomposition such that $\max_i \lambda_i(\mu)>0$ for all invariant measures $\mu$ supported by a component of the Morse decomposition. When the Morse components are Axiom A, uniquely ergodic, or support all but one population, the necessary and sufficient conditions are equivalent. Applications to spatial ecology, epidemiology, and gene networks are given

Generalized Urn Models of Evolutionary Processes

Generalized Polya urn models can describe the dynamics of finite populations of interacting genotypes. Three basic questions these models can address are: Under what conditions does a population exhibit growth? On the event of growth, at what rate does the population increase? What is the long-term behavior of the distribution of genotypes? To address these questions, we associate a mean limit ordinary differential equation (ODE) with the urn model. Previously, it has been shown that on the event of population growth, the limiting distribution of genotypes is a connected internally chain recurrent set for the mean limit ODE. To determine when growth and convergence occurs with positive probability, we prove two results. First, if the mean limit ODE has an ``attainable'' attractor at which growth is expected, then growth and convergence toward this attractor occurs with positive probability. Second, the population distribution almost surely does not converge to sets where growth is not expecte

Protected polymorphisms and evolutionary stability of patch-selection strategies in stochastic environments

We consider a population living in a patchy environment that varies stochastically in space and time. The population is composed of two morphs (that is, individuals of the same species with different genotypes). In terms of survival and reproductive success, the associated phenotypes differ only in their habitat selection strategies. We compute invasion rates corresponding to the rates at which the abundance of an initially rare morph increases in the presence of the other morph established at equilibrium. If both morphs have positive invasion rates when rare, then there is an equilibrium distribution such that the two morphs coexist; that is, there is a protected polymorphism for habitat selection. Alternatively, if one morph has a negative invasion rate when rare, then it is asymptotically displaced by the other morph under all initial conditions where both morphs are present. We refine the characterization of an evolutionary stable strategy for habitat selection from [Schreiber, 2012] in a mathematically rigorous manner. We provide a necessary and sufficient condition for the existence of an ESS that uses all patches and determine when using a single patch is an ESS. We also provide an explicit formula for the ESS when there are two habitat types. We show that adding environmental stochasticity results in an ESS that, when compared to the ESS for the corresponding model without stochasticity, spends less time in patches with larger carrying capacities and possibly makes use of sink patches, thereby practicing a spatial form of bet hedging.Comment: Revised in light of referees' comments, Published on-line Journal of Mathematical Biology 2014 http://link.springer.com/article/10.1007/s00285-014-0824-