Prey dispersal and predator persistence

To understand how patchiness influences population dynamics of a tri-trophic interaction, a tractable model is formulated in terms of differential equations. Motivated by the structure of systems such as plants, phytophagous mites and predatory mites, the model takes dispersal into account at the middle trophic level. The effect of dispersal for the middle level in a tri-trophic system could be either stabilising or destabilising since the middle level acts both as prey and as predator. First a simple model with logistic growth for the lowest level is formulated. A model with logistic growth for the lowest level and instantaneous dispersal has a globally stable three-species equilibrium, if this equilibrium exists. Addition of a middle level dispersal phase of non-negligible duration influences equilibrium stability. In the absence of the top trophic level a limit cycle can occur, caused by the delay that exists in the reaction of the middle level to the changes in the lowest level. With low predator efficiency, it is also possible to have an unstable three-species equilibrium. So addition of a middle level dispersal phase of non-negligible duration can work destabilising. Next the persistence of the third trophic level is studied. Even when the three-species equilibrium exists, the third trophic level need not be persistent. A two-species limit cycle can keep its stability when a three-species equilibrium exists; the system is then bistable. It is argued that such a bistability can offer an alternative explanation for pesticide-induced outbreaks of spider mites and failure of predator introduction.     

Spatial heterogeneity in three species,plant-parasite-hyperparasite,systems

This paper addresses the question of how heterogeneity may evolve due to interactions between the dynamics and movement of three-species systems involving hosts, parasites and hyperparasites in homogeneous environments. The models are motivated by the spread of soil-borne parasites within plant populations, where the hyperparasite is used as a biological control agent but where patchiness in the distribution of the parasite occurs, even when environmental conditions are apparently homogeneous. However, the models are introduced in generic form as three-species reaction-diffusion systems so that they have broad applicability to a range of ecological systems. We establish necessary criteria for the occurrence of population-driven patterning via diffusion-driven instability. Sufficient conditions are obtained for restricted cases with no host movement. The criteria are similar to those for the well-documented two-species reaction-diffusion system, although more possibilities arise for spatial patterning with three species. In particular, temporally varying patterns, that may be responsible for the apparent drifting of hot-spots of disease and periodic occurrence of disease at a given location, are possible when three species interact. We propose that the criteria can be used to screen population interactions, to distinguish those that cannot cause patterning from those that may give rise to population-driven patterning. This establishes a basic dynamical ''landscape'' against which other perturbations, including environmentally driven variations, can be analysed and distinguished from population-driven patterns. By applying the theory to a specific model example for host-parasite-hyperparasite interactions both with and without host movement, we show directly how the evolution of spatial pattern is related to biologically meaningful parameters. In particular, we demonstrate that when there is strong density dependence limiting host growth, the pattern is stable over time, whereas with less stable underlying host growth, the pattern varies with time.     

Biomass and water storage dynamics of epiphytes in old-growth and secondary montane cloud forest stands in Costa Rica

The way competition structures plant communities has been investigated intensely over many decades. Dominance structures due to competitive hierarchies, with consequences for species richness, have not received as much experimental attention, since their manipulation is a large logistic undertaking. Here the data from a model system based on bryophytes are presented to investigate competition structure in a three-species system. Grown in monocultures, pairwise and three-species mixtures under no and high nitrogen supply, the three moss species responded strongly to treatment conditions. One of them suffered from nitrogen fertilisation and hence performed better in mixtures, where the dominant species provided physical shelter from apparently toxic nitrogen spray. Accordingly, no linear competitive hierarchy emerged and qualitative transitivity remains restricted to the unfertilised treatments. Faciliation also affected other properties of the competition structure. The reciprocity of competition effects could not be observed. Moreover, the performances in three-species mixtures were not well predictable from the knowledge of monocultures and pairwise mixtures because competitive effects were not additive. This had implications for community stability at equilibrium: all two-species systems were stable, both fertilised and unfertilised, while the three-species system was only stable when fertilised. This stability under fertilisation has probably to do with the facilitative effect of the two dominant species on the third. In this experiment, little support for commonly held ideas was found about the way competition in plant communities is structured. On the other hand, this study shows that moss communities are ideal model systems to test predictions of theoretical models concerning properties and consequences of competition in plant communities.     

Effects of diffusion on the stability of the equilibrium in multi-species ecological systems

Continuous population distributions that undergo self-diffusion, migrational cross-diffusion and interaction in a region of (1-, 2- or 3-dimensional) space are described dynamically by a governing system of nonlinear reaction-diffusion equations. It is shown that the constants associated with migrational cross-diffusion are ordinarily nonnegative or nonpositive, contingent on the type of species interaction. A simple sign relationship obtains between the latter diffusivity constants and the rate constants for species interaction in the neighborhood of a spatially uniform equilibrium state, and this relationship of signs serves to simplify the general stability theory for the growth or decay of arbitrary perturbations on a spatially uniform equilibrium state. The stability of the equilibrium state is analyzed and discussed in detail for the case of a generic two-species model, where the self-diffusion and migrational cross-diffusion of species act to either stabilize or destabilize the equilibrium, depending essentially on the character of the species interaction and also on the magnitude of the Helmholtz eigenvalues associated with the region and boundary conditions. In particular, for a prey-predator or host-parasite model, self-diffusion usually helps to stabilize the equilibrium state and migrational cross-diffusion can only act as an additional stabilizing influence, as evidenced generally by the experiments on such two-species systems. Sufficient conditions are derived for stability of the equilibrium state in the general case for an arbitrarily large number of interacting species. It is shown that the equilibrium state is always stable if all species undergo significant self-diffusion and the Helmholtz eigenvalues are suitably large.     

Existence and bifurcation of stable equilibrium in two-prey,one-predator communities

In this paper, stability of two-prey, one-predator communities is investigated by Lyapunov's direct method and Hopf's bifurcation theory. Three patterns of three-species coexistence are possible. A globally stable non-negative equilibrium exists for the system even if two competing prey species without a predator cannot coexist. The stable equilibrium bifurcates to a periodic motion with a small amplitude when the predation rate increases. It is also shown that a chaotic motion emerges from the periodic motion when one of two prey has greater competitive abilities than the other. This predator-mediated coexistence can be realized by the intimate relationship between preferences of a predator and competitive abilities of two prey.     

Density dependent selection incorporating intraspecific competition 1. A haploid model

A haploid model is introduced and analyzed in which intraspecific competition is incorporated within a density dependent framework. It is assumed that each genotype has a unique carrying capacity corresponding to the equilibrium population size when fixed for that type. Each genotypic fitness at a single multi-allelic locus is a function of a distinctive effective population size formed by adding the numbers of each genotype present, weighted by an intraspecific competition coefficient. As a result, the fitnesses depend upon the relative frequencies of the various genotypes as well as the total population size. Intergenotypic interactions can have a profound effect upon the outcome of the population. In particular, when the density effect of one individual upon another depends upon their respective genotypes, a unique stable interior equilibrium is possible in which all alleles are present. This stands in contrast to the purely density dependent haploid system in which the only possible stable state corresponds to fixation for the type with the highest carrying capacity. In the present model selective advantage is determined by a balance between carrying capacity and sensitivity to density pressures from other genotypes. Fixation for the genotype with the highest carrying capacity, for instance, will not be stable if it exerts a sufficiently weak competitive effect upon the other genotypes. In the diallelic case, maintenance of both alleles at a stable equilibrium requires that the net intragenotypic competition between individuals of like genotype be stronger than that between unlike types. As for purely density regulated systems, there may be no stable equilibria and/or regular and chaotic cycling may occur. The results may also be interpreted in terms of a discrete time model of interspecific competition with each haplotype representing a different species.     

Global Stability of a System of Two Interacting Species with Convective and Dispersive Migration

Using Liapunov's direct method, effects of convective and dispersive migration on the global stability of the equilibrium state of a system of two interacting species are investigated. It is shown that the stable equilibrium state without dispersal remains so with dispersal. Further, it is pointed out that stability or instability of the equilibrium state of the system is not affected by convective migration. These results are justified in cases of a system of mutualistic interactions of species and a prey-predator system with functional response.     

Effects of convective and dispersive interactions on the stability of two species

The evolution and local stability of a system of two interacting species in a finite two-dimensional habitat is investigated by taking into account the effects of self- and cross-dispersion and convection of the species. In absence of cross-dispersion, an equilibrium state which is stable without dispersion is always stable with dispersion provided that the dispersion coefficients of the two species are equal. However, when the dispersion coefficients of the two species are different, the possibility of self-dispersive instability arises. It is also pointed out that the cross-dispersion of species may lead to stability or instability depending upon the nature and the magnitude of the cross-dispersive interactions in comparison to the self-dispersive interactions. The self-convective movement of species increases the stability of the equilibrium state and can stabilize an otherwise unstable equilibrium state. The effect of cross-convection (in absence of self-dispersion and self-convection) is to stabilize the equilibrium state in a prey-predator model with positive cross-dispersion coefficients for the prey species. Finally, it is shown that if the system is stable under homogeneous boundary conditions it remains so under non-homogeneous boundary conditions.     

Impact of fear effect on the growth of prey in a predator-prey interaction model

Several field data and experiments on a terrestrial vertebrates exhibited that the fear of predators would cause a substantial variability of prey demography. Fear for predator population enhances the survival probability of prey population, and it can greatly reduce the reproduction of prey population. Based on the experimental evidence, we proposed and analyzed a prey-predator system introducing the cost of fear into prey reproduction with Holling type-II functional response. We investigate all the biologically feasible equilibrium points, and their stability is analyzed in terms of the model parameters. Our mathematical analysis exhibits that for strong anti-predator responses can stabilize the prey-predator interactions by ignoring the existence of periodic behaviors. Our model system undergoes Hopf bifurcation by considering the birth rate r₀ as a bifurcation parameter. For larger prey birth rate, we investigate the transition to a stable coexisting equilibrium state, with oscillatory approach to this equilibrium state, indicating that the greatest characteristic eigenvalues are actually a pair of imaginary eigenvalues with real part negative, which is increasing for r₀. We obtained the conditions for the occurrence of Hopf bifurcation and conditions governing the direction of Hopf bifurcation, which imply that the prey birth rate will not only influence the occurrence of Hopf bifurcation but also alter the direction of Hopf bifurcation. We identify the parameter regions associated with the extinct equilibria, predator-free equilibria and coexisting equilibria with respect to prey birth rate, predator mortality rates. Fear can stabilize the predator-prey system at an interior steady state, where all the species can exists together, or it can create the oscillatory coexistence of all the populations. We performed some numerical simulations to investigate the relationship between the effects of fear and other biologically related parameters (including growth/decay rate of prey/predator), which exhibit the impact that fear can have in prey-predator system. Our numerical illustrations also demonstrate that the prey become less sensitive to perceive the risk of predation with increasing prey growth rate or increasing predators decay rate.     

Effects of dispersion on stability of multispecies prey-predator systems

The general multispecies prey-predator system with Gompertz's antisymmetric interactions is nonlinearly stable in the absence of dispersion and continues to remain stable with dispersion under both homogeneous reservoir and zero flux boundary conditions in a region containing the equilibrium state. It is proved that a general multispecies food-web model without antisymmetric interactions is stable in the absence of dispersion and remains stable with dispersion in the above-mentioned region.     

Dynamics of commensalistic systems with self- and cross-inhibition

Up to three stable steady states are possible in a simple commensalistic system, taking place in an open-loop mixed reactor when the growth rates of the two species are inhibited by the substrates they prey on (Self-inhibition). Two stable states are possible in a system with noncompetitive inhibition of the species by the substrate they are not preying on (cross-inhibition). A large number of steady states as well as oscillatory states are possible when both self- and cross-inhibition are strong. Multiplicity of steady states is also possible in a reactor with biomas recirculation for these kinetics. Yet, the latter is more stable than the open-loop reactor in the sense that the domain of steady-state multiplicity is narrower. The stability of steady states and the dynamics of the systems for each of the investigated kinetics are summarized in a qualitative phase plane. The importance of the analysis for improving the selectively and yield of the system and for predicting the response of the system to changes in the operating conditions, is discussed.     

Stability of commensalistic systems

The response of two-species commensalistic systems in a chemostat has been investigated after perturbations in steady state conditions and after step changes in dilution rate. The system is inherently stable with not more than three overshoots and undershoots possible. More complicated commensalistic systems are less stable, with limit cycle response occurring after dilution rate changes when feedback inhibition and feedforward activation occurs. In general variation of feedback parameters is more effective in changing the behavior of the systems than variation of feedforward parameters. Limited agreement with the experimental data of Chao and Reilly was obtained.     

Ideal free distributions, evolutionary games, and population dynamics in multiple-species environments

In this article, we develop population game theory, a theory that combines the dynamics of animal behavior with population dynamics. In particular, we study interaction and distribution of two species in a two-patch environment assuming that individuals behave adaptively (i.e., they maximize Darwinian fitness). Either the two species are competing for resources or they are in a predator-prey relationship. Using some recent advances in evolutionary game theory, we extend the classical ideal free distribution (IFD) concept for single species to two interacting species. We study population dynamical consequences of two-species IFD by comparing two systems: one where individuals cannot migrate between habitats and one where migration is possible. For single species, predator-prey interactions, and competing species, we show that these two types of behavior lead to the same population equilibria and corresponding species spatial distributions, provided interspecific competition is patch independent. However, if differences between patches are such that competition is patch dependent, then our predictions strongly depend on whether animals can migrate or not. In particular, we show that when species are settled at their equilibrium population densities in both habitats in the environment where migration between habitats is blocked, then the corresponding species spatial distribution need not be an IFD. Thus, when species are given the opportunity to migrate, they will redistribute to reach an IFD (e.g., under which the two species can completely segregate), and this redistribution will also influence species population equilibrial densities. Alternatively, we also show that when two species are distributed according to the IFD, the corresponding population equilibrium can be unstable.     

Influence of consumer mutual interference on the stabilization of a three-level trophic system

In this paper, we present a three-level (food–prey–predator) trophic food chain which includes consumer mutual interference (MIF). In contrast with other analyses, we consider the effect of both prey and predator MIF on the dynamics of a three-level trophic system. MIF is generally considered to exert a stabilizing effect on population dynamics based on the predator–prey model. However, results from analytical and numerical simulations utilizing a simple three-species food chain model suggest that while the addition of prey MIF to the model provides a stabilizing influence, as the chaotic dynamics collapse to a stable steady state, adding only predator MIF to the model can only stabilize the system at intermediate MIF values. The three-species trophic food chain is also stabilized when combination of both prey and predator MIF is added to the model. Our work serves to provide insight into the effects of MIF in the real world.     

The Role of Space in the Exploitation of Resources

In order to understand the role of space in ecological communities where each species produces a certain type of resource and has varying abilities to exploit the resources produced by its own species and by the other species, we carry out a comparative study of an interacting particle system and its mean-field approximation. For a wide range of parameter values, we show both analytically and numerically that the spatial model results in predictions that significantly differ from its nonspatial counterpart, indicating that the use of the mean-field approach to describe the evolution of communities in which individuals only interact locally is invalid. In two-species communities, the disagreements between the models appear either when both species compete by producing resources that are more beneficial for their own species or when both species cooperate by producing resources that are more beneficial for the other species. In particular, while both species coexist if and only if they cooperate in the mean-field approximation, the inclusion of space in the form of local interactions may prevent coexistence even in cooperative communities. Introducing additional species, cooperation is no longer the only mechanism that promotes coexistence. We prove that, in three-species communities, coexistence results either from a global cooperative behavior, or from rock-paper-scissors type interactions, or from a mixture of these dynamics, which excludes in particular all cases in which two species compete. Finally, and more importantly, we show numerically that the inclusion of space has antagonistic effects on coexistence depending on the mechanism involved, preventing coexistence in the presence of cooperation but promoting coexistence in the presence of rock-paper-scissors interactions. Although these results are partly proved analytically for both models, we also provide somewhat more explicit heuristic arguments to explain the reason why the models result in different predictions.     

Three-step food chains in Gompertz and Lotka-Volterra models

Some new results for three-step prey-predator food chains, which appear in the exactly solvable Gompertz model, are shown to follow in the Lotka-Volterra model also. Possible implications of these results are discussed.     

Chaotic dynamics of a three species prey-predator competition model with bionomic harvesting due to delayed environmental noise as external driving force

We consider a biological economic model based on prey-predator interactions to study the dynamical behaviour of a fishery resource system consisting of one prey and two predators surviving on the same prey. The mathematical model is a set of first order non-linear differential equations in three variables with the population densities of one prey and the two predators. All the possible equilibrium points of the model are identified, where the local and global stabilities are investigated. Biological and bionomical equilibriums of the system are also derived. We have analysed the population intensities of fluctuations i.e., variances around the positive equilibrium due to noise with incorporation of a constant delay leading to chaos, and lastly have investigated the stability and chaotic phenomena with a computer simulation.     

A new model for interacting populations—II: Principle of competitive exclusion

The principle of competitive exclusion is investigated within the framework of the solvable model proposed earlier for two-species systems. The results elucidate the recent controversy over the interpretation of experimental data onDrosophila equilibrium. It is shown that the necessary and sufficient conditions for stable coexistence of competing species is that the product of intraspecific rate constants be greater than the product of interspecific rate constants. Inequalities between rate constants for the occurrence of stable equilibriumbelow andabove the line joining single species equilibria are derived. The availability of larger domain of coexistence suggests that the model presented here has the potential to accommodate a wider class of phenomena than the Gause—Volterra model according to which coexistence is possible only above the line of single species equilibrium.     

Conditions for periodic solutions of volterra differential systems

As a contribution to the discussion of oscillatory models for interacting species it is shown that two-species Volterra models can never have limit cycles, and a complete enumeration is given of conditions which the parameters of these models must satisfy in order that a part of the phase space be filled with a family of closed curves; sketches of phase portraits are also given. These results complement and correct older results by Bautin and by Coppel on quadratic differential systems. The paper opens with a brief discussion of some more practical aspects of the ecological application of oscillatory models.     

What makes ecological systems reactive?

Although perturbations from a stable equilibrium must ultimately vanish, they can grow initially, and the maximum initial growth rate is called reactivity. Reactivity thus identifies systems that may undergo transient population surges or drops in response to perturbations; however, we lack biological and mathematical intuition about what makes a system reactive. This paper presents upper and lower bounds on reactivity for an arbitrary linearized model, explores their strictness, and discusses their biological implications. I find that less stable systems (i.e. systems with long transients) have a smaller possible range of reactivities for which no perturbations grow. Systems with more species have a higher capacity to be reactive, assuming species interactions do not weaken too rapidly as the number of species increases. Finally, I find that in discrete time, reactivity is determined largely by mean interaction strength and neither discrete nor continuous time reactivity are sensitive to food web topology.