Persistence in models of three interacting predator-prey populations

This paper considers a class of deterministic models of three interacting populations with a view towards determining when all of the populations persist. In analytical terms persistence means that liminf_t→∞x(t)> 0 for each population x(t); in geometric terms, that each trajectory of the modeling system of differential equations is eventually bounded away from the coordinate planes. The class of systems considered allows three level food webs, two competing predators feeding on a single prey, or a single predator feeding on two competing prey populations. As a corollary to the last case it is shown that the addition of a predator can lead to persistence of a three population system where, without a predator, the two competing populations on the lower trophic level would have only one survivor. The basic models are of Kolmogorov type, and the results improve several previous theorems on persistence.     

Cannibalism in discrete-time predator-prey systems

In this study, we propose and investigate a two-stage population model with cannibalism. It is shown that cannibalism can destabilize and lower the magnitude of the interior steady state. However, it is proved that cannibalism has no effect on the persistence of the population. Based on this model, we study two systems of predator-prey interactions where the prey population is cannibalistic. A sufficient condition based on the nontrivial boundary steady state for which both populations can coexist is derived. It is found via numerical simulations that introduction of the predator population may either stabilize or destabilize the prey dynamics, depending on cannibalism coefficients and other vital parameters.     

Remarks on "persistence in models of three interacting predator-prey populations".

For models of three interacting predator-prey populations, a result on the boundedness of solution orbits and a result on ultimate boundedness are presented. A counterexample to another result is also given for comparison.     

Persistence in predator-prey systems with ratio-dependent predator influence

Predator-prey models where one or more terms involve ratios of the predator and prey populations may not be valid mathematically unless it can be shown that solutions with positive initial conditions never get arbitrarily close to the axis in question, i.e. that persistence holds. By means of a transformation of variables, criteria for persistence are derived for two classes of such models, thereby leading to their validity. Although local extinction certainly is a common occurrence in nature, it cannot be modeled by systems which are ratio-dependent near the axes. Research partially supported by the Natural Sciences and Engineering Research Council of Canada, Grant No. NSERC A4823. Research carried out while visiting the University of Alberta.     

Analysis of a predator-prey system with predator switching

In this paper, we consider an interaction of prey and predator species where prey species have the ability of group defence. Thresholds, equilibria and stabilities are determined for the system of ordinary differential equations. Taking carrying capacity as a bifurcation parameter, it is shown that a Hopf bifurcation can occur implying that if the carrying capacity is made sufficiently large by enrichment of the environment, the model predicts the eventual extinction of the predator providing strong support for the so-called ‘paradox of enrichment’.     

Absolute stability in predator-prey models

An equilibrium of a time-lagged population model is said to be absolutely stable if it remains locally stable regardless of the length of the time delay, and it is argued that the criteria for absolute stability provide a valuable guide to the behavior of population models. For example, it is sometimes assumed that time delays have a limited impact until they exceed the natural time scale of a system; here it is stressed that under some conditions very short time delays can have a marked (and often maximal) destabilizing effect. Consequently it is important that our understanding of population dynamics is robust to the inclusion of the short time delays present in all biological systems. The absolute stability criteria are ideally suited for this role. Another important reason for using the criteria for absolute stability rather than using criteria which depend upon the details of a time delay is that biological time delays are unlikely to be constant. For example, a time delay due to maturation inevitably varies between individuals and the mean may itself vary over time. Here it is shown that the criteria for absolute stability are generally robust in the presence of distributed delays and of varying delays. The analysis presented is based upon a general predator-prey model and it is shown that absolute stability can be expected under a broad range of parameter values whenever the time delay is due to the maturation time of either the predator or the prey or of both. This stability occurs because of the interaction between delayed and undelayed dynamic features of the model. A time-delayed process, when viewed across all possible delays, always reduces stability and this effect occurs regardless of whether the process would act to stabilize or destabilize an undelayed system. Opposing the destabilization due to a time delay and making absolute stability a possibility are a number of processes which act without delay. Some of these processes can be identified as stabilizing from the analysis of undelayed models (for example, the type 3 functional response) but other cannot (for example, the nonreproductive numerical response of predators).     

Limit cycles in predator-prey models

The general model of interaction between one predator and one prey is studied. A unimodal function of rate of growth of the prey and concave down functional response of the predator is assumed. In this work it is shown that for a given natural number n there exist models possessing at least 2n + 1 limit cycles. It is also proved, applying the Hopf bifurcation theorem, that a model exists with a logistic growth rate of the prey and concave down functional response that has at least two limit cycles.     

Multiple limit cycles in predator-prey models

A series of one-predator one-prey models are studied using two parameter Hopf bifurcation techniques which allow the determination of two periodic orbits. The biological implications of the results, in terms of domains of attraction and multiple stable states, are discussed.     

Analysis of cyclic fluctuations in larch bud moth populations by means of discrete-time dynamic models

Analysed are the data of larch bud moth (Zeiraphera diniana Gn.) fluctuations in Swiss Alps. The analysis applies simplest mathematical models of isolated population dynamics (in particular, Kostitzin model, Skellam model, the discrete logistic model, and some other ones), which include the minimal number of unknown parameters. The parameters have been estimated, for all the models in hand, by the least-squares method, to fit certain data from the Global Population Dynamics Database (N 1407 and N 6195), the sequences of the data deviations from the model trajectories being treated as well. The best approximations are shown to be achieved with Moran-Ricker model and the discrete logistic model. Statistical criteria (Kolmogorov-Smirnov and Shapiro-Wilk tests) reveal that the hypotheses of normal distribution of residuals must be rejected for one of the time series (N 1407); some models demonstrate serial correlations in the sequence of residuals (according to Durbin-Watson test). This leads to the conclusion that periodic fluctuations in the larch bud moth population (N 1407) can hardly be explained by self-regulation mechanisms alone. For another time series (N 6195), the modified discrete logistic model has appeared to be acceptable as a mode of fluctuations.     

Individual base model of predator-prey system shows predator dominant dynamics

Individual base model of predator-prey system is constructed. Both predator and prey species have age structure and cohorts of early reproductive age have competitive advantage. The model has linear functional response in predation behavior and includes the effect of interference among predators and delay of population growth from resource intake, not by functional response but by calculation procedure. Each foraging action is calculated successively and surplus or scarce of acquired resources is interpreted into population size through individual birth and death. This model shows that biomass of prey killed by predator is dependent on demand of predator and that heterogeneity in predator population is essential in persistency and stability of predator-prey system. Heterogeneity of predator makes predator individuals of less competing ability die rapidly. Rapid death of weak individuals causes rapid decrease of total demand of predator and that makes enough room for survived predators. Therefore, the biomass of killed prey is dependent on predator's demand. As young or infant population of predator are the more vulnerable to shortage of prey, and when many of them cannot survive to reproductive age, they can stabilize the system by wasting excessive prey with only temporal numerical increase of predator population.     

The importance of census times in discrete-time growth-dispersal models

Dispersal has been the focus of spatial ecology for a few decades. What should be a proper theoretical framework for understanding and modelling of dispersal processes remains a controversial issue though. Integrodifference equations (IDE) model the spatial dynamics of a population with distinct growth and dispersal stages in their life cycle. Depending on the stage observed, the equations take on different forms, only one of which is usually studied in the literature. Here we reveal that while these different forms are mathematically equivalent, the biological conclusions drawn from the different forms may differ considerably. We provide a summary of similarities and differences and point out the greatest potential caveats when applying IDE.     

The importance of census times in discrete-time growth-dispersal models

Dispersal has been the focus of spatial ecology for a few decades. What should be a proper theoretical framework for understanding and modelling of dispersal processes remains a controversial issue though. Integrodifference equations (IDE) model the spatial dynamics of a population with distinct growth and dispersal stages in their life cycle. Depending on the stage observed, the equations take on different forms, only one of which is usually studied in the literature. Here we reveal that while these different forms are mathematically equivalent, the biological conclusions drawn from the different forms may differ considerably. We provide a summary of similarities and differences and point out the greatest potential caveats when applying IDE.     

Testing for predator dependence in predator-prey dynamics: a non-parametric approach

The functional response is a key element in all predator-prey interactions. Although functional responses are traditionally modelled as being a function of prey density only, evidence is accumulating that predator density also has an important effect. However, much of the evidence comes from artificial experimental arenas under conditions not necessarily representative of the natural system, and neglecting the temporal dynamics of the organism (in particular the effects of prey depletion on the estimated functional response). Here we present a method that removes these limitations by reconstructing the functional response non-parametrically from predator-prey time-series data. This method is applied to data on a protozoan predator-prey interaction, and we obtain significant evidence of predator dependence in the functional response. A crucial element in this analysis is to include time-lags in the prey and predator reproduction rates, and we show that these delays improve the fit of the model significantly. Finally, we compare the non-parametrically reconstructed functional response to parametric forms, and suggest that a modified version of the Hassell-Varley predator interference model provides a simple and flexible function for theoretical investigation and applied modelling.     

About deterministic extinction in ratio-dependent predator-prey models

Ratio-dependent predator-prey models set up a challenging issue regarding their dynamics near the origin. This is due to the fact that such models are undefined at (0, 0). We study the analytical behavior at (0, 0) for a common ratio-dependent model and demonstrate that this equilibrium can be either a saddle point or an attractor for certain trajectories. This fact has important implications concerning the global behavior of the model, for example regarding the existence of stable limit cycles. Then, we prove formally, for a general class of ratio-dependent models, that (0, 0) has its own basin of attraction in phase space, even when there exists a non-trivial stable or unstable equilibrium. Therefore, these models have no pathological dynamics on the axes and at the origin, contrary to what has been stated by some authors. Finally, we relate these findings to some published empirical results.     

Stabilizing dispersal delays in predator-prey metapopulation models

Time delays produced by dispersal are shown to stabilize Lotka-Volterra predator-prey models. The models are formulated as integrodifferential equations that describe local predator-prey dynamics and either intrapatch or interpatch dispersal. Dispersing individuals may (or may not) differ in the duration of their trips; these differences are captured via a distributed delay in the models. Our results include those of previous studies as special cases and show that the stabilizing effect continues to operate when the dispersal process is modeled more realistically.     

The effect of predator avoidance and travel time delay on the stability of predator-prey metacommunities

The stability conditions for an isolated specialist predator-prey community are fairly well understood. The spatial coupling of several such systems through dispersal of individuals can generate new dynamic behavior that is not yet completely understood. Many factors are known to be stabilizing or neutral, e.g., random dispersal or time delays, while others may induce instabilities in some cases but not others, e.g., density-dependent movement. We study the combination of two stabilizing mechanisms in a two-patch Rosenzweig-MacArthur model with a novel density-dependent movement term. Specifically, we assume that prey move between patches according to their perceived predation risk, and we include travel time between patches as a time delay. We show that the combination of mechanisms may be destabilizing even though each mechanism by itself is stabilizing. Our results show that a detailed knowledge of mechanisms and their temporal scales is necessary to correctly predict the stability of a metacommunity.     

Spatiotemporal dynamics of two generic predator-prey models

We present the analysis of two reaction-diffusion systems modelling predator-prey interactions, where the predator displays the Holling type II functional response, and in the absence of predators, the prey growth is logistic. The local analysis is based on the application of qualitative theory for ordinary differential equations and dynamical systems, while the global well-posedness depends on invariant sets and differential inequalities. The key result is an L (∞)-stability estimate, which depends on a polynomial growth condition for the kinetics. The existence of an a priori L ( p )-estimate, uniform in time, for all p≥1, implies L (∞)-uniform bounds, given any nonnegative L (∞)-initial data. The applicability of the L (∞)-estimate to general reaction-diffusion systems is discussed, and how the continuous results can be mimicked in the discrete case, leading to stability estimates for a Galerkin finite-element method with piecewise linear continuous basis functions. In order to verify the biological wave phenomena of solutions, numerical results are presented in two-space dimensions, which have interesting ecological implications as they demonstrate that solutions can be 'trapped' in an invariant region of phase space.     

Ongoing collapse of coral-reef shark populations

Marine ecosystems are suffering severe depletion of apex predators worldwide; shark declines are principally due to conservative life-histories and fisheries overexploitation. On coral reefs, sharks are strongly interacting apex predators and play a key role in maintaining healthy reef ecosystems. Despite increasing fishing pressure, reef shark catches are rarely subject to specific limits, with management approaches typically depending upon no-take marine reserves to maintain populations. Here, we reveal that this approach is failing by documenting an ongoing collapse in two of the most abundant reef shark species on the Great Barrier Reef (Australia). We find an order of magnitude fewer sharks on fished reefs compared to no-entry management zones that encompass only 1% of reefs. No-take zones, which are more difficult to enforce than no-entry zones, offer almost no protection for shark populations. Population viability models of whitetip and gray reef sharks project ongoing steep declines in abundance of 7% and 17% per annum, respectively. These findings indicate that current management of no-take areas is inadequate for protecting reef sharks, even in one of the world's most-well-managed reef ecosystems. Further steps are urgently required for protecting this critical functional group from ecological extinction.     

Directed movement of predators and the emergence of density-dependence in predator-prey models.

We consider a bitrophic spatially distributed community consisting of prey and actively moving predators. The model is based on the assumption that the spatial and temporal variations of the predators' velocity are determined by the prey gradient. Locally, the populations follow the simple Lotka-Volterra interaction. We also assume predator reproduction and mortality to be negligible in comparison with the time scale of migration. The model demonstrates heterogeneous oscillating distributions of both species, which occur because of the active movements of predators. One consequence of this heterogeneity is increased viability of the prey population, compared to the equivalent homogeneous model, and increased consumption. Further numerical analysis shows that, on the spatially aggregated scale, the average predator density adversely affects the individual consumption, leading to a nonlinear predator-dependent trophic function, completely different from the Lotka-Volterra rule assumed at the local scale.