Modeling the dispersal-reproduction trade-off in an expanding population.

Trade-offs between dispersal and reproduction are known to be important drivers of population dynamics, but their direct influence on the spreading speed of a population is not well understood. Using integrodifference equations, we develop a model that incorporates a dispersal-reproduction trade-off which allows for a variety of different shaped trade-off curves. We show there is a unique reproductive-dispersal allocation that gives the largest value for the spreading speed and calculate the sensitivities of the reproduction, dispersal, and trade-off shape parameters. Uncertainty in the model parameters affects the expected spread of the population and we calculate the optimal allocation of resources to dispersal that maximizes the expected spreading speed. Higher allocation to dispersal arises from uncertainty in the reproduction parameter or the shape of the reproduction trade-off curve. Lower allocation to dispersal arises from uncertainty in the shape of the dispersal trade-off curve, but does not come from uncertainty in the dispersal parameter. Our findings give insight into how parameter sensitivity and uncertainty influence the spreading speed of a population with a dispersal-reproduction trade-off.

Inside Dynamics of Integrodifference Equations with Mutations.

The method of inside dynamics provides a theory that can track the dynamics of neutral gene fractions in spreading populations. However, the role of mutations has so far been absent in the study of the gene flow of neutral fractions via inside dynamics. Using integrodifference equations, we develop a neutral genetic mutation model by extending a previously established scalar inside dynamics model. To classify the mutation dynamics, we define a mutation class as the set of neutral fractions that can mutate into one another. We show that the spread of neutral genetic fractions is dependent on the leading edge of population as well as the structure of the mutation matrix. Specifically, we show that the neutral fractions that contribute to the spread of the population must belong to the same mutation class as the neutral fraction found in the leading edge of the population. We prove that the asymptotic proportion of individuals at the leading edge of the population spread is given by the dominant right eigenvector of the associated mutation matrix, independent of growth and dispersal parameters. In addition, we provide numerical simulations to demonstrate our mathematical results, to extend their generality and to develop new conjectures about our model.

Inside dynamics for stage-structured integrodifference equations.

A stage-structured model of integrodifference equations is used to study the asymptotic neutral genetic structure of populations undergoing range expansion. That is, we study the inside dynamics of solutions to stage-structured integrodifference equations. To analyze the genetic consequences for long term population spread, we decompose the solution into neutral genetic components called neutral fractions. The inside dynamics are then given by the spatiotemporal evolution of these neutral fractions. We show that, under some mild assumptions on the dispersal kernels and population projection matrix, the spread is dominated by individuals at the leading edge of the expansion. This result is consistent with the founder effect. In the case where there are multiple neutral fractions at the leading edge we are able to explicitly calculate the asymptotic proportion of these fractions found in the long-term population spread. This formula is simple and depends only on the right and left eigenvectors of the population projection matrix evaluated at zero and the initial proportion of each neutral fraction at the leading edge of the range expansion. In the absence of a strong Allee effect, multiple neutral fractions can drive the long-term population spread, a situation not possible with the scalar model.

Integrodifference equations in the presence of climate change: persistence criterion, travelling waves and inside dynamics.

To understand the effects that the climate change has on the evolution of species as well as the genetic consequences, we analyze an integrodifference equation (IDE) models for a reproducing and dispersing population in a spatio-temporal heterogeneous environment described by a shifting climate envelope. Our analysis on the IDE focuses on the persistence criterion, travelling wave solutions, and the inside dynamics. First, the persistence criterion, characterizing the global dynamics of the IDE, is established in terms of the basic reproduction number. In the case of persistence, a unique travelling wave is found to govern the global dynamics. The effects of the size and the shifting speed of the climate envelope on the basic reproduction number, and hence, on the persistence criterion, are also investigated. In particular, the critical domain size and the critical shifting speed are found in certain cases. Numerical simulations are performed to complement the theoretical results. In the case of persistence, we separate the travelling wave and general solutions into spatially distinct neutral fractions to study the inside dynamics. It is shown that each neutral genetic fraction rearranges itself spatially so as to asymptotically achieve the profile of the travelling wave. To measure the genetic diversity of the population density we calculate the Shannon diversity index and related indices, and use these to illustrate how diversity changes with underlying parameters.

Neutral Genetic Patterns for Expanding Populations with Nonoverlapping Generations.

We investigate the inside dynamics of solutions to integrodifference equations to understand the genetic consequences of a population with nonoverlapping generations undergoing range expansion. To obtain the inside dynamics, we decompose the solution into neutral genetic components. The inside dynamics are given by the spatiotemporal evolution of the neutral genetic components. We consider thin-tailed dispersal kernels and a variety of per capita growth rate functions to classify the traveling wave solutions as either pushed or pulled fronts. We find that pulled fronts are synonymous with the founder effect in population genetics. Adding overcompensation to the dynamics of these fronts has no impact on genetic diversity in the expanding population. However, growth functions with a strong Allee effect cause the traveling wave solution to be a pushed front preserving the genetic variation in the population. In this case, the contribution of each neutral fraction can be computed by a simple formula dependent on the initial distribution of the neutral fractions, the traveling wave solution, and the asymptotic spreading speed.

Eco-evolutionary dynamics of range expansion.

Understanding the movement of species' ranges is a classic ecological problem that takes on urgency in this era of global change. Historically treated as a purely ecological process, range expansion is now understood to involve eco-evolutionary feedbacks due to spatial genetic structure that emerges as populations spread. We synthesize empirical and theoretical work on the eco-evolutionary dynamics of range expansion, with emphasis on bridging directional, deterministic processes that favor evolved increases in dispersal and demographic traits with stochastic processes that lead to the random fixation of alleles and traits. We develop a framework for understanding the joint influence of these processes in changing the mean and variance of expansion speed and its underlying traits. Our synthesis of recent laboratory experiments supports the consistent role of evolution in accelerating expansion speed on average, and highlights unexpected diversity in how evolution can influence variability in speed: results not well predicted by current theory. We discuss and evaluate support for three classes of modifiers of eco-evolutionary range dynamics (landscape context, trait genetics, and biotic interactions), identify emerging themes, and suggest new directions for future work in a field that stands to increase in relevance as populations move in response to global change.

Modelling the biological invasion of Carcinus maenas (the European green crab).

This paper proposes a system of integro-difference equations to model the spread of Carcinus maenas, commonly called the European green crab, that causes severe damage to coastal ecosystems. A model with juvenile and adult classes is first studied. Here, standard theory of monotone operators for integro-difference equations can be applied and yields explicit formulas for the asymptotic spreading speeds of the juvenile and adult crabs. A second model including an infected class is considered by introducing a castrating parasite Sacculina carcini as a biological control agent. The dynamics are complicated and simulations reveal the occurrence of periodic solutions and stacked fronts. In this case, only conjectures can be made for the asymptotic spreading speeds because of the lack of mathematical theory for non-monotone operators. This paper also emphasizes the need for mathematical studies of non-monotone operators in heterogeneous environments and the existence of stacked front solutions in biological invasion models.