The diffusive Lotka–Volterra competition model in fragmented patches I: Coexistence

It is an ecological imperative that we understand how changes in landscape heterogeneity affect population dynamics and coexistence among species residing in increasingly fragmented landscapes. Decades of research have shown the dispersal process to have major implications for individual fitness, species’ distributions, interactions with other species, population dynamics, and stability. Although theoretical models have played a crucial role in predicting population level effects of dispersal, these models have largely ignored the conditional dependency of dispersal (e.g., responses to patch boundaries, matrix hostility, competitors, and predators). This work is the first in a series where we explore dynamics of the diffusive Lotka–Volterra (L–V) competition model in such a fragmented landscape. This model has been extensively studied in isolated patches, and to a lesser extent, in patches surrounded by an immediately hostile matrix. However, little attention has been focused on studying the model in a more realistic setting considering organismal behavior at the patch/matrix interface. Here, we provide a mechanistic connection between the model and its biological underpinnings and study its dynamics via exploration of nonexistence, existence, and uniqueness of the model’s steady states. We employ several tools from nonlinear analysis, including sub-supersolutions, certain eigenvalue problems, and a numerical shooting method. In the case of weak, neutral, and strong competition, our results mostly match those of the isolated patch or immediately hostile matrix cases. However, in the case where competition is weak towards one species and strong towards the other, we find existence of a maximum patch size, and thus an intermediate range of patch sizes where coexistence is possible, in a patch surrounded by an intermediate hostile matrix when the weaker competitor has a dispersal advantage. These results support what ecologists have long theorized, i.e., a key mechanism promoting coexistence among competing species is a tradeoff between dispersal and competitive ability.     

具有生态位构建作用的种群进化动力学模型及其应用研究

依据进化动力学的理论与方法,系统探讨了生态位构建的机理与模式．通过建立生态位构建的空间模式及其适合度计算公式和具有生态位构建作用的单种群与两种群的进化动力学模型,并对其种群进化动态、种间竞争共存机制进行的理论与数值模拟分析,揭示了生物与环境资源的协同进化关系．结果表明:种群动态受其主要生态因子及资源含量的正反馈作用．生态位构建作用通过对种群适宜度的影响而产生进化响应．单种群动力系统存在种群大小的阈值效应;在两竞争种群动力系统中,生态位构建可以导致进化动力系统的多个竞争结果,从而为解释种间竞争与稳定共存提供了一种新的理论机制．     

具有时滞的周期非均匀竞争恒化器模型的空间动力学

本文提出了一个具有时滞的周期非均匀单种营养基——双种微生物的竞争恒化器模型,利用半群理论, 获得了该模型解的存在唯一性. 进一步, 建立了该模型的竞争排斥原理, 给出了两竞争物种共存的充分条件.     

A mutualism-competition model characterizing competitors with mutualism at low density

A macroscopic model of two species is considered, in which mutualism is the dominant interaction when the species are at low density and competition is the dominant interaction when they are at high density. Our aim is to show that species using the same or similar resources can coexist without niche differentiation and that mutualism at low population density can lead to high production. The specific model is a novel combination of the Lotka–Volterra cooperative (mutualism) model and Lotka–Volterra competitive model. By comparing the dynamics of the specific system with those of the Lotka–Volterra competitive model, we demonstrate the mechanism by which the mutualism at low density promotes competitive coexistence by creating regions of mutualism that maintain coexistence. We also show situations in which high production occurs by (i) displaying regions of net mutualism in which the species with higher competitive ability (the superior) approaches a density larger than its carrying capacity when in isolation from the inferior species, and (ii) displaying regions of net mutualism in which both of the species approach densities larger than their carrying capacities, respectively. By comparing the dynamics of the specific system with those of the Lotka–Volterra mutualism model, we show that competition at high density promotes stability of the system.     

Competitive exclusion in a discrete juvenile–adult model with continuous and seasonal reproduction

We develop a general discrete juvenile–adult population model that describes two competing species. We consider species in which the juveniles only compete with other juveniles, and the adults only compete with other adults, i.e. juveniles and adults of either species do not compete. This is typical of amphibians where juveniles (tadpoles) live in water and adults (frogs) live on land. Assuming competition efficiencies of the two species are similar, we analyse the cases where reproduction is either continuous or seasonal. In both cases, we develop conditions on the invasion net reproductive numbers of the two species that will lead to competitive exclusion. We show using numerical simulations that coexistence and bistability are possible outcomes when competition efficiencies of the two species are different.     

竞争性生态系统中两物种的共存问题

利用Banach空间中锥与半序理论研究了竞争性生态系统中两物种的共存问题,得出依赖于某一共同生存环境的两物种在一定条件下必可达唯一共存状态．     

A mathematical model for chemical defense mechanism of two competing species

In this paper, a non-linear mathematical model is proposed and analyzed to study the phenomenon of a chemical defense mechanism involving two competing species, where each species produces a toxicant affecting the other. It is shown that if the emission rate coefficient of toxicant, produced by one species increases, the equilibrium density of the other species decreases and its magnitude is lower than its original carrying capacity. It is found that the usual principle of competitive exclusion (coexistence) in the absence of toxicant may change in the case under consideration. It is also observed that increases in the values of production rates of toxicants by the competing species and depletion rates of environmental toxicants due to its assimilation by the species has a destabilizing effect, and decrease in the washout rates of environmental toxicants has a destabilizing effect on the dynamics of the system. In the case of allelopathy, where only one species produces a toxicant affecting the other species, it is shown that the affected species is driven to extinction for large production rate of this toxicant.     

Dynamical behaviors of a classical Lotka–Volterra competition–diffusion–advection system

This paper deals with a two-species competition model in a homogeneous advective environment, where two species are subjected to a net loss of individuals at the downstream end. Under the assumption that the advection and diffusion rates of two species are proportional, we give a basic classification on the global dynamics by employing the theory of monotone dynamical system. It turns out that bistability does not happen, but coexistence and competitive exclusion may occur. Furthermore, we present a complete classification on the global dynamics in terms of the growth rates of two species. However, once the above assumption does not hold, bistability may occur. In detail, there exists a tradeoff between growth rates of two species such that competition outcomes can shift between three possible scenarios, including competitive exclusion, bistability and coexistence. These results show that growth competence is important to determine dynamical behaviors.     

Exclusionary population dynamics in size-structured,discrete competitive systems

A discrete multi-species size-structured competition model is considered. By using decreasing growth functions, we achieve the self-regulation of species. We develop various biologically significant conditions for global convergence to the extinction state of the dominated species in the competitive system. With an example we illustrate coexistence in a chaotic supr transient. The chaotic attractor has an unusual pulsating nature.     

On competitive Lotka-Volterra model in random environments

Focusing on competitive Lotka-Volterra model in random environments, this paper uses regime-switching diffusions to model the dynamics of the population sizes of n different species in an ecosystem subject to the random changes of the external environment. It is demonstrated that the growth rates of the population sizes of the species are bounded above. Moreover, certain long-run-average limits of the solution are examined from several angles. A partial stochastic principle of competitive exclusion is also derived. Finally, simple examples are used to demonstrate our findings.     

Advection-mediated competition in general environments

We consider a reaction–diffusion–advection system of two competing species with one of the species dispersing by random diffusion as well as a biased movement upward along resource gradient, while the other species by random diffusion only. It has been shown that, under some non-degeneracy conditions on the environment function, the two species always coexist when the advection is strong. In this paper, we show that for general smooth environment function, in contrast to what is known, there can be competitive exclusion when the advection is strong, and, we give a sharp criterion for coexistence that includes all previously considered cases. Moreover, when the domain is one-dimensional, we derive in the strong advection limit a system of two equations defined on different domains. Uniqueness of steady states of this non-standard system is obtained when one of the diffusion rates is large.     

Coexistence in a discrete competition model with dispersal

We propose a discrete-time competition model between two populations to study the effects of dispersal upon population interactions. It is assumed that dispersal occurs after reproduction and in synchrony. We first analyse a two-patch single species population model with no interspecific competition. Based on these results, we derive sufficient conditions for population coexistence. It is proved that the system is uniformly persistent and possesses a unique coexisting equilibrium.     

Crowding effects on coexistence solutions in the unstirred chemostat

This paper deals with the unstirred chemostat model with crowding effects. The introduction of crowding effects makes the conservation law invalid, and the equations cannot be combined to eliminate one of the variables. Consequently, the usual reduction of the system to a competitive system of one order lower is lost. Thus the system with predation and competition is non-monotone, and the single population model cannot be reduced to a scalar system. First, the uniqueness and asymptotic behaviors of the semi-trivial solutions are established. Second, the existence and structure of coexistence solutions are given by the degree theory and bifurcation theory. It turns out that the positive bifurcation branch connects one semi-trivial solution branch with another. Finally, the stability and asymptotic behaviors of coexistence solutions are discussed in some cases. It is shown that crowding effects are sufficiently effective in the occurrence of coexisting, and overcrowding of a species has an inhibiting effect on itself.     

Coexistence and stability of a competition—diffusion system in population dynamics

The coexistence and stability of the population densities of two competing species in a bounded habitat are investigated in the present paper, where the effect of dispersion (transportation) is taken into consideration. The mathematical problem involves a coupled system of Lotka-Volterra-type reaction-diffusion equations together with some initial and boundary conditions, including the Dirichlet, Neumann and third type. Necessary and sufficient conditions for the coexistence and competitive exclusion are established and the effect of diffusion is explicitly given. For the stability problem, general criteria for the stability and instability of a steady-state solution are established and then applied to various situations depending on the relative magnitude among the physical parameters. Also given are necessary and sufficient conditions for the existence of multiple steady-state solutions and the stability or instability of each of these solutions. Special attention is given to the Neumann boundary condition with respect to which some threshold results for the coexistence and stability or instability of the four uniform steady states are characterized. It is shown in this situation that only one of the four constant steady states is asymptotically stable while the remaining three are unstable. The stability or instability of these states depends solely on the relative magnitude among the various rate constants and is independent of the diffusion coefficients.     

Competitive exclusion and coexistence for competitive systems on ordered Banach spaces

The dynamics of competitive maps and semiflows defined on the product of two cones in respective Banach spaces is studied. It is shown that exactly one of three outcomes is possible for two viable competitors. Either one or the other population becomes extinct while the surviving population approaches a steady state, or there exists a positive steady state representing the coexistence of both populations.

     

Pathogen coexistence induced by saturating contact rates

In this paper, a two-strain epidemic model with saturating contact rates is considered. It is shown that if the social activity of infected individuals does not vary with strains, then the competitive exclusion principle holds; if the social activity of infected individuals varies with different strains, the coexistence of pathogens is possible under a certain condition which involves the invasion reproduction numbers. The stability of the dominance equilibria and coexistence equilibrium is also examined. Numerical simulations are presented to illustrate the results.     

Complex dynamics in the Leslie–Gower type of the food chain system with multiple delays

In this paper, we present a Leslie–Gower type of food chain system composed of three species, which are resource, consumer, and predator, respectively. The digestion time delays corresponding to consumer-eat-resource and predator-eat-consumer are introduced for more realistic consideration. It is called the resource digestion delay (RDD) and consumer digestion delay (CDD) for simplicity. Analyzing the corresponding characteristic equation, the stabilities of the boundary and interior equilibrium points are studied. The food chain system exhibits the species coexistence for the small values of digestion delays. Large RDD/CDD may destabilize the species coexistence and induce the system dynamic into recurrent bloom or system collapse. Further, the present of multiple delays can control species population into the stable coexistence. To investigate the effect of time delays on the recurrent bloom of species population, the Hopf bifurcation and periodic solution are investigated in detail in terms of the central manifold reduction and normal form method. Finally, numerical simulations are performed to display some complex dynamics, which include multiple periodic solution and chaos motion for the different values of system parameters. The system dynamic behavior evolves into the chaos motion by employing the period-doubling bifurcation.     

Coexistence of Two Species in a Strongly Coupled Schoener’s Competitive Model

This paper deals with the existence of positive solution to a strongly coupled system with homogeneous Dirichlet boundary conditions describing a Schoener’s competitive interaction of two species. Making use of the Schauder fixed point theorem, a sufficient condition is given for the system to have a coexistence. And true solutions are constructed based on monotone iterative method. Our results show that this model possesses at least one coexistence state if cross-diffusions and intra-specific competitions are weak.     

具有阶段结构的竞争系统中自食的稳定性作用

该文讨论了两种群竞争系统解的动力学行为，其中一种群分幼年，成年两阶段，当不考虑自食时，得到了阶段结构的竞争系统也可以出现三种典型的动力学行为，共存（coexietence)，双稳定（bistability),占优（dominance)，进一步，在没有自食时竞争系统是占优的情形下，考虑自食的影响，得到了所有种群永久持续生存的充分条件，这表明自食有稳定性的作用。     

Stable coexistence mediated by specialist harvesting in a two zooplankton–phytoplankton system

This paper deals with a predator–prey model with specialist harvesting, representing a two predators (Zooplankton) and one resource (Phytoplankton) system. First, the existence and stability of equilibria is analyzed both from local and global point of view. Our results indicate that a specialist harvesting which is discriminate may mediate the coexistence of the two zooplankton species which competitively exclude each other in absence harvesting. Although in most cases increasing harvesting reduces the two zooplankton species numbers, when harvesting leads to coexistence, it may also lead to increase the two zooplankton species numbers. Furthermore, to protect fish population from over exploitation a control instrument tax is imposed. The problem of optimal taxation policy is then solved by using Pontryagin’s maximal principle. It is established that the zero discounting leads to the maximization of the net economic revenue to the society and an infinite discount rate leads to complete dissipation of the net economic revenue to the society. Finally, the impact of harvesting is mentioned along with numerical results to provide some support to the analytical findings.