Effects of dispersal rates in a two-stage reaction-diffusion system.

It is well known that in reaction-diffusion models for a single unstructured population in a bounded, static, heterogeneous environment, slower diffusion is advantageous. That is not necessarily the case for stage structured populations. In (Cantrell et al. 2020), it was shown that in a stage structured model introduced by Brown and Lin (1980), there can be situations where faster diffusion is advantageous. In this paper we extend and refine the results of (Cantrell et al. 2020) on persistence to more general combinations of diffusion rates and to cases where either adults or juveniles do not move. We also obtain results on the asymptotic behavior of solutions as diffusion rates go to zero, and on competition between species that differ in their diffusion rates but are otherwise ecologically identical. We find that when the spatial distributions of favorable habitats for adults and juveniles are similar, slow diffusion is still generally advantageous, but if those distributions are different that may no longer be the case.

A model for the coupling of the Greater Bairam and local environmental factors in promoting Rift-Valley Fever epizootics in Egypt.

OBJECTIVES: Rift-Valley Fever (RVF) is a zoonotic mosquito-borne disease in Africa and the Arabian Peninsula. Drivers for this disease vary by region and are not well understood for North African countries such as Egypt. A deeper understanding of RVF risk factors would inform disease management policies. STUDY DESIGN: The present study employs mathematical and computational modeling techniques to ascertain the extent to which the severity of RVF epizootics in Egypt differs depending on the interaction between imported ruminant and environmentally-constrained mosquito populations. METHODS: An ordinary differential system of equations, a numerical model, and an individual-based model (IBM) were constructed to represent RVF disease dynamics between localized mosquitoes and ruminants being imported into Egypt for the Greater Bairam. Four cases, corresponding to the Greater Bairam's occurrence during distinct quarters of the solar year, were set up in both models to assess whether the different season-associated mosquito populations present during the Greater Bairam resulted in RVF epizootics of variable magnitudes. RESULTS: The numerical model and the IBM produced nearly identical results: ruminant and mosquito population plots for both models were similar in shape and magnitude for all four cases. In both models, all four cases differed in the severity of their corresponding simulated RVF epizootics. The four cases, ranked by the severity of the simulated RVF epizootics in descending order, correspond with the occurrence of the Greater Bairam on the following months: July, October, April, and January. The numerical model was assessed for sensitivity with respect to parameter values and exhibited a high degree of robustness. CONCLUSIONS: Limiting the importation of infected ruminants beginning one month prior to the Greater Bairam festival (on years in which the festival falls between the months of July and October: 2014-2022) might be a feasible way of mitigating future RVF epizootics in Egypt.

The effects of human movement on the persistence of vector-borne diseases.

With the recent resurgence of vector-borne diseases due to urbanization and development there is an urgent need to understand the dynamics of vector-borne diseases in rapidly changing urban environments. For example, many empirical studies have produced the disturbing finding that diseases continue to persist in modern city centers with zero or low rates of transmission. We develop spatial models of vector-borne disease dynamics on a network of patches to examine how the movement of humans in heterogeneous environments affects transmission. We show that the movement of humans between patches is sufficient to maintain disease persistence in patches with zero transmission. We construct two classes of models using different approaches: (i) Lagrangian models that mimic human commuting behavior and (ii) Eulerian models that mimic human migration. We determine the basic reproduction number R(0) for both modeling approaches. We show that for both approaches that if the disease-free equilibrium is stable (R(0)<1) then it is globally stable and if the disease-free equilibrium is unstable (R(0)>1) then there exists a unique positive (endemic) equilibrium that is globally stable among positive solutions. Finally, we prove in general that Lagrangian and Eulerian modeling approaches are not equivalent. The modeling approaches presented provide a framework to explore spatial vector-borne disease dynamics and control in heterogeneous environments. As an example, we consider two patches in which the disease dies out in both patches when there is no movement between them. Numerical simulations demonstrate that the disease becomes endemic in both patches when humans move between the two patches.

Deriving reaction-diffusion models in ecology from interacting particle systems.

We use a scaling procedure based on averaging Poisson distributed random variables to derive population level models from local models of interactions between individuals. The procedure is suggested by using the idea of hydrodynamic limits to derive reaction-diffusion models for population interactions from interacting particle systems. The scaling procedure is formal in the sense that we do not address the issue of proving that it converges; instead we focus on methods for computing the results of the scaling or deriving properties of rescaled systems. To that end we treat the scaling procedure as a transform, in analogy with the Laplace or Fourier transform, and derive operational formulas to aid in the computation of rescaled systems or the derivation of their properties. Since the limiting procedure is adapted from work by Durrett and Levin, we refer to the transform as the Durrett-Levin transform. We examine the effects of rescaling in various standard models, including Lotka-Volterra models, Holling type predator-prey models, and ratio-dependent models. The effects of scaling are mostly quantitative in models with smooth interaction terms, but ratio-dependent models are profoundly affected by the scaling. The scaling transforms ratio-dependent terms that are singular at the origin into smooth terms. Removing the singularity at the origin eliminates some of the unique dynamics that can arise in ratio-dependent models.

Spatial heterogeneity and critical patch size: Area effects via diffusion in closed environments.

We describe a class of mathematical models for critical patch size in which the mechanisms inducing area effects are based on source-sink population dynamics arising from dispersal throughout a closed, finite, but spatially heterogeneous environment. Our models are reaction-diffusion equations, but unlike classical KISS models for area effects they do not assume that there is dispersal across the boundary of the environment into a hostile exterior. We observe that simple rescaling has the same effects in our models as in KISS models and hence predicts the same sort of area effects, but that other sorts of rescaling may not predict area effects. The models considered here provide an alternative to the KISS models used in our previous work on species-area relationships in island biogeography.

Brucellosis, botflies, and brainworms: the impact of edge habitats on pathogen transmission and species extinction.

Ecological interactions between species that prefer different habitat types but come into contact in edge regions at the interfaces between habitat types are modeled via reaction-diffusion systems. The primary sort of interaction described by the models is competition mediated by pathogen transmission. The models are somewhat novel because the spatial domains for the variables describing the population densities of the interacting species overlap but do not coincide. Conditions implying coexistence of the two species or the extinction of one species are derived. The conditions involve the principal eigenvalues of elliptic operators arising from linearizations of the model system around equilibria with only one species present. The conditions for persistence or extinction are made explicit in terms of the parameters of the system and the geometry of the underlying spatial domains via estimates of the principal eigenvalues. The implications of the models with respect to conservation and refuge design are discussed.

How predator incursions affect critical patch size: the role of the functional response.

Understanding the impact of habitat edges provides a key to deciphering how community dynamics change as functions of habitat structure and spatial scale. Motivated by studies of predation on bird nests in forest fragments and other cases of "cross-boundary subsidies," we present results from a partial differential equation model in which a patch-resident prey species suffers incidental mortality from a generalist predator species residing in the surrounding matrix habitat. We demonstrate that predator intrusions have the potential to induce critical patch size effects for the prey species, even when the prey's dynamics would otherwise preclude such effects. We also demonstrate that the existence of critical patch size effects depends on the functional response of the predator, with Lotka-Volterra and Type II functional responses generating the effect (but not Type III). We conclude by discussing how predator-induced critical patch size effects can influence opportunities for regionwide persistence of the prey by altering the fraction and spatial distribution of meaningful patches within a metapopulation.

A comparison of foraging strategies in a patchy environment.

In this paper we compare foraging strategies that might be used by predators seeking prey in a patchy environment. The strategies differ in the extent to which predators aggregate in response to prey density. The approach to the comparison is suggested by the idea of evolutionarily stable strategies. A strategy is said to be evolutionarily stable if it cannot be invaded by another strategy. Thus we examine scenarios where a small number of individuals using one strategy are introduced into a situation where a large number of individuals using the other strategy are already present. However, our foraging models do not explicitly incorporate predator population dynamics, so we use net energy uptake as a surrogate for reproductive fitness. In cases where all of the patches visited by predators sustain prey populations, we find that for any pair of strategies one of them will have a higher net energy uptake than the other whether it is the resident or the introduced strain. However, which one is higher will typically depend on the total predator population, which is determined by the resident strain. If the predators leave prey densities high, the more aggregative strain will have the advantage. If the predators reduce prey densities to low levels the less aggregative strain will have the advantage. In cases where one strain of predators aggregates in response to prey density and the other does not, then there might be patches which do not contain prey but do contain (non-aggregating) predators. In those cases, there is the possibility that whichever strategy is used by the introduced strain will yield a higher energy uptake than that used by the resident strain. This suggests that if some patches are empty of prey then aggregative and non-aggregative strategies may be able to coexist.

Effects of spatial grouping on the functional response of predators.

A unified mechanistic approach is given for the derivation of various forms of functional response in predator-prey models. The derivation is based on the principle of mass action but with the crucial refinement that the nature of the spatial distribution of predators and/or opportunities for predation are taken into account in an implicit way. If the predators are assumed to have a homogeneous spatial distribution, then the derived functional response is prey-dependent. If the predators are assumed to form a dense colony or school in a single (possibly moving) location, or if the region where predators can encounter prey is assumed to be of limited size, then the functional response depends on both predator and prey densities in a manner that reflects feeding interference between predators. Depending on the specific assumptions, the resulting functional response may be of Beddington-DeAngelis type, of Hassell-Varley type, or ratio-dependent.

Diffusion models for population dynamics incorporating individual behavior at boundaries: applications to refuge design.

We construct models for dispersal of a population which incorporate the response of individuals to interfaces between habitat types. The models are based on random walks where there may be a bias in the direction an individual moves when it encounters an interface. This sort of dispersal process is called skew Brownian motion. Our models take the form of diffusion equations with matching conditions across the interface between regions for population densities and fluxes. We combine the dispersal models with linear population growth models which assume that the population growth rate differs between regions of different habitat types. We use those models to study issues of refuge design. We specifically consider how the effectiveness of buffer zones depends on their size, quality, and the population's response to the interface between the buffer zone and the refuge.

Competitive reversals inside ecological reserves: the role of external habitat degradation.

Habitat degradation is the slow--and often subtle--deterioration in habitat quality that accompanies human activities through increases in road density, pesticide use, hunting pressure, etc. Such degradation is of particular concern in fragmented habitats where economic or jurisdictional boundaries rather than ecological ones determine the level of exploitation adjoining habitat patches endure. To examine the consequences habitat degradation might have on species interactions, we posited a patch of pristine habitat surrounded by "matrix" habitat whose degradation level was variable. Using a coupled pair of diffusive Lotka-Volterra competition equations with Robin (mixed) boundary conditions, we modeled the dynamics of two competing species inhabiting the pristine patch and incorporated matrix degradation through a tunable "hostility" parameter representing species' mortality rates in the matrix. We found that the numerical range of competition coefficients over which one species is the competitive dominant and the other inferior may grow or shrink as matrix quality deteriorates. In some cases, degradation of the exterior habitat would bring about a complete competitive reversal inside the preserve. This result, wherein a formerly inferior species supplants a formerly dominant one--even inside the "protected" remnant patch itself--has policy implications for both nature reserve design and management of human activities outside park boundaries.

Spatially explicit models for the population dynamics of a species colonizing an island.

We construct reaction-diffusion models for the population dynamics of a species colonizing an island from a source population on a continent. We view the source population as inducing a density or flux of immigrants onto the island and interpret colonization as succeeding if the population on the island is predicted to persist even when immigration from the continent is stopped. To capture the observation that a sufficiently large population or density must be attained for colonization to succeed, we assume Allee (i.e., bistable) dynamics rather than logistic dynamics for the colonizing population. We consider the cases of colonization in both the absence and presence of a competitor. We use reaction-diffusion theory, especially comparison methods and sub- and supersolutions, to determine how parameters such as the distance from the continent to the island and the dispersal, birth and mortality rates, carrying capacity, and minimum viable population density of the colonizing species affect the outcome of the attempted colonization. In the case of colonization in the presence of a competitor we consider a number of scenarios involving different types and strengths of competition. Our analysis permits us to draw conclusions about the characteristics of a species that make it a good colonizer.

Models for the effects of individual size and spatial scale on competition between species in heterogeneous environments.

A spatially explicit model for competition with dispersal in a heterogeneous environment is used to study the effects of individual size and the spatial scale of the environment on the competitive interactions between species. The model is a Lotka-Volterra competition system with diffusion and with spatial variation in some coefficients. The coefficients in the model are taken to reflect a situation where the larger competitor typically disperses farther in unit time than the smaller and reproduces less rapidly, but has an advantage in contests or other forms of interference competition. The environment is assumed to be closed, i.e., it is assumed that individuals do not leave through the boundary. The environment is generally assumed to consist of a patch of favorable habitat surrounded by less favorable regions. The effects of spatial scale are studied by examining how the predictions of the model change as the size of the favorable patch is varied. The predictions turn out to be in qualitative agreement with the results of some empirical studies.

Threshold conditions for a diffusive model of a myelinated axon.

This paper develops and uses comparison principles to study the time evolution of solutions to problems of the form (formula; see text) Such a system models an infinite myelinated axon with discrete, excitable nodes spaced unit distant apart.