Prey group defense and hunting cooperation among generalist-predators induce complex dynamics: a mathematical study.

Group defense in prey and hunting cooperation in predators are two important ecological phenomena and can occur concurrently. In this article, we consider cooperative hunting in generalist predators and group defense in prey under a mathematical framework to comprehend the enormous diversity the model could capture. To do so, we consider a modified Holling-Tanner model where we implement Holling type IV functional response to characterize grazing pattern of predators where prey species exhibit group defense. Additionally, we allow a modification in the attack rate of predators to quantify the hunting cooperation among them. The model admits three boundary equilibria and up to three coexistence equilibrium points. The geometry of the nontrivial prey and predator nullclines and thus the number of coexistence equilibria primarily depends on a specific threshold of the availability of alternative food for predators. We use linear stability analysis to determine the types of hyperbolic equilibrium points and characterize the non-hyperbolic equilibrium points through normal form and center manifold theory. Change in the model parameters leading to the occurrences of a series of local bifurcations from non-hyperbolic equilibrium points, namely, transcritical, saddle-node, Hopf, cusp and Bogdanov-Takens bifurcation; there are also occurrences of global bifurcations such as homoclinic bifurcation and saddle-node bifurcation of limit cycles. We observe two interesting closed 'bubble' form induced by global bifurcations due to change in the strength of hunting cooperation and the availability of alternative food for predators. A three dimensional bifurcation diagram, concerning the original system parameters, captures how the alternation in model formulation induces gradual changes in the bifurcation scenarios. Our model highlights the stabilizing effects of group or gregarious behaviour in both prey and predator, hence supporting the predator-herbivore regulation hypothesis. Additionally, our model highlights the occurrence of "saltatory equilibria" in ecological systems and capture the dynamics observed for lion-herbivore interactions.

Bridging Theories for Ecosystem Stability Through Structural Sensitivity Analysis of Ecological Models in Equilibrium.

Ecologists are challenged by the need to bridge and synthesize different approaches and theories to obtain a coherent understanding of ecosystems in a changing world. Both food web theory and regime shift theory shine light on mechanisms that confer stability to ecosystems, but from different angles. Empirical food web models are developed to analyze how equilibria in real multi-trophic ecosystems are shaped by species interactions, and often include linear functional response terms for simple estimation of interaction strengths from observations. Models of regime shifts focus on qualitative changes of equilibrium points in a slowly changing environment, and typically include non-linear functional response terms. Currently, it is unclear how the stability of an empirical food web model, expressed as the rate of system recovery after a small perturbation, relates to the vulnerability of the ecosystem to collapse. Here, we conduct structural sensitivity analyses of classical consumer-resource models in equilibrium along an environmental gradient. Specifically, we change non-proportional interaction terms into proportional ones, while maintaining the equilibrium biomass densities and material flux rates, to analyze how alternative model formulations shape the stability properties of the equilibria. The results reveal no consistent relationship between the stability of the original models and the proportionalized versions, even though they describe the same biomass values and material flows. We use these findings to critically discuss whether stability analysis of observed equilibria by empirical food web models can provide insight into regime shift dynamics, and highlight the challenge of bridging alternative modelling approaches in ecology and beyond.

Mathematical models for dengue fever epidemiology: A 10-year systematic review.

Mathematical models have a long history in epidemiological research, and as the COVID-19 pandemic progressed, research on mathematical modeling became imperative and very influential to understand the epidemiological dynamics of disease spreading. Mathematical models describing dengue fever epidemiological dynamics are found back from 1970. Dengue fever is a viral mosquito-borne infection caused by four antigenically related but distinct serotypes (DENV-1 to DENV-4). With 2.5 billion people at risk of acquiring the infection, it is a major international public health concern. Although most of the cases are asymptomatic or mild, the disease immunological response is complex, with severe disease linked to the antibody-dependent enhancement (ADE) - a disease augmentation phenomenon where pre-existing antibodies to previous dengue infection do not neutralize but rather enhance the new infection. Here, we present a 10-year systematic review on mathematical models for dengue fever epidemiology. Specifically, we review multi-strain frameworks describing host-to-host and vector-host transmission models and within-host models describing viral replication and the respective immune response. Following a detailed literature search in standard scientific databases, different mathematical models in terms of their scope, analytical approach and structural form, including model validation and parameter estimation using empirical data, are described and analyzed. Aiming to identify a consensus on infectious diseases modeling aspects that can contribute to public health authorities for disease control, we revise the current understanding of epidemiological and immunological factors influencing the transmission dynamics of dengue. This review provide insights on general features to be considered to model aspects of real-world public health problems, such as the current epidemiological scenario we are living in.

Complexity of host-vector dynamics in a two-strain dengue model.

We introduce a compartmental host-vector model for dengue with two viral strains, temporary cross-immunity for the hosts, and possible secondary infections. We study the conditions on existence of endemic equilibria where one strain displaces the other or the two virus strains co-exist. Since the host and vector epidemiology follow different time scales, the model is described as a slow-fast system. We use the geometric singular perturbation technique to reduce the model dimension. We compare the behaviour of the full model with that of the model with a quasi-steady approximation for the vector dynamics. We also perform numerical bifurcation analysis with parameter values from the literature and compare the bifurcation structure to that of previous two-strain host-only models.

Analysis of a predator-prey model with specific time scales: a geometrical approach proving the occurrence of canard solutions.

We study a predator-prey model with different characteristic time scales for the prey and predator populations, assuming that the predator dynamics is much slower than the prey one. Geometrical Singular Perturbation theory provides the mathematical framework for analyzing the dynamical properties of the model. This model exhibits a Hopf bifurcation and we prove that when this bifurcation occurs, a canard phenomenon arises. We provide an analytic expression to get an approximation of the bifurcation parameter value for which a maximal canard solution occurs. The model is the well-known Rosenzweig-MacArthur predator-prey differential system. An invariant manifold with a stable and an unstable branches occurs and a geometrical approach is used to explicitly determine a solution at the intersection of these branches. The method used to perform this analysis is based on Blow-up techniques. The analysis of the vector field on the blown-up object at an equilibrium point where a Hopf bifurcation occurs with zero perturbation parameter representing the time scales ratio, allows to prove the result. Numerical simulations illustrate the result and allow to see the canard explosion phenomenon.

Is structural sensitivity a problem of oversimplified biological models? Insights from nested Dynamic Energy Budget models.

Many current issues in ecology require predictions made by mathematical models, which are built on somewhat arbitrary choices. Their consequences are quantified by sensitivity analysis to quantify how changes in model parameters propagate into an uncertainty in model predictions. An extension called structural sensitivity analysis deals with changes in the mathematical description of complex processes like predation. Such processes are described at the population scale by a specific mathematical function taken among similar ones, a choice that can strongly drive model predictions. However, it has only been studied in simple theoretical models. Here, we ask whether structural sensitivity is a problem of oversimplified models. We found in predator-prey models describing chemostat experiments that these models are less structurally sensitive to the choice of a specific functional response if they include mass balance resource dynamics and individual maintenance. Neglecting these processes in an ecological model (for instance by using the well-known logistic growth equation) is not only an inappropriate description of the ecological system, but also a source of more uncertain predictions.

The impact of a predator on the outcome of competition in the three-trophic food web.

We study the effects of predation on the competition of prey populations for two resources in a chemostat. We investigate a variety of small food web compositions: the bi-trophic food web (two resources-two competing prey) and the three-trophic food web (two resources-two prey-generalist predator) comparing different model formulations: substitutable resources and essential resources, namely Liebig's minimum law model (perfect essential resources) and complementary resources formulations. The prediction of the outcome of competition is solely based on bifurcation analysis in which the inflow of resources into the chemostat is used as the bifurcation parameter. We show that the results for different bi-trophic food webs are very similar, as only equilibria are involved in the long-term dynamics. In the three-trophic food web, the outcome of competition is manifested largely by non-equilibrium dynamics, i.e., in oscillatory behavior. The emergence of predator-prey cycles leads to strong deviations between the predictions of the outcome of competition based on Liebig's minimum law and the complementary resources. We show that the complementary resources formulation yields a stabilization of the three-trophic food web by decreasing the existence interval of oscillations. Furthermore, we find an exchange of a region of oscillatory co-existence of all three species in Liebig's formulation by a region of bistability of two limit cycles containing only one prey and the predator in the complementary formulation.

Ecoepidemic predator-prey model with feeding satiation, prey herd behavior and abandoned infected prey.

In this paper we analyse a predator-prey model where the prey population shows group defense and the prey individuals are affected by a transmissible disease. The resulting model is of the Rosenzweig-MacArthur predator-prey type with an SI (susceptible-infected) disease in the prey. Modeling prey group defense leads to a square root dependence in the Holling type II functional for the predator-prey interaction term. The system dynamics is investigated using simulations, classical existence and asymptotic stability analysis and numerical bifurcation analysis. A number of bifurcations, such as transcritical and Hopf bifurcations which occur commonly in predator-prey systems will be found. Because of the square root interaction term there is non-uniqueness of the solution and a singularity where the prey population goes extinct in a finite time. This results in a collapse initiated by extinction of the healthy or susceptible prey and thereafter the other population(s). When also a positive attractor exists this leads to bistability similar to what is found in predator-prey models with a strong Allee effect. For the two-dimensional disease-free (i.e. the purely demographic) system the region in the parameter space where bistability occurs is marked by a global bifurcation. At this bifurcation a heteroclinic connection exists between saddle prey-only equilibrium points where a stable limit cycle together with its basin of attraction, are destructed. In a companion paper (Gimmelli et al., 2015) the same model was formulated and analysed in which the disease was not in the prey but in the predator. There we also observed this phenomenon. Here we extend its analysis using a phase portrait analysis. For the three-dimensional ecoepidemic predator-prey system where the prey is affected by the disease, also tangent bifurcations including a cusp bifurcation and a torus bifurcation of limit cycles occur. This leads to new complex dynamics. Continuation by varying one parameter of the emerging quasi-periodic dynamics from a torus bifurcation can lead to its destruction by a collision with a saddle-cycle. Under other conditions the quasi-periodic dynamics changes gradually in a trajectory that lands on a boundary point where the prey go extinct in finite time after which a total collapse of the three-dimensional system occurs.

Competition for light and nutrients in layered communities of aquatic plants.

Dominance of free-floating plants poses a threat to biodiversity in many freshwater ecosystems. Here we propose a theoretical framework to understand this dominance, by modeling the competition for light and nutrients in a layered community of floating and submerged plants. The model shows that at high supply of light and nutrients, floating plants always dominate due to their primacy for light, even when submerged plants have lower minimal resource requirements. The model also shows that floating-plant dominance cannot be an alternative stable state in light-limited environments but only in nutrient-limited environments, depending on the plants' resource consumption traits. Compared to unlayered communities, the asymmetry in competition for light-coincident with symmetry in competition for nutrients-leads to fundamentally different results: competition outcomes can no longer be predicted from species traits such as minimal resource requirements ([Formula: see text] rule) and resource consumption. Also, the same two species can, depending on the environment, coexist or be alternative stable states. When applied to two common plant species in temperate regions, both the model and field data suggest that floating-plant dominance is unlikely to be an alternative stable state.

Analysis of an asymmetric two-strain dengue model.

In this paper we analyse a two-strain compartmental dengue fever model that allows us to study the behaviour of a Dengue fever epidemic. Dengue fever is the most common mosquito-borne viral disease of humans that in recent years has become a major international public health concern. The model is an extension of the classical compartmental susceptible-infected-recovered (SIR) model where the exchange between the compartments is described by ordinary differential equations (ode). Two-strains of the virus exist so that a primary infection with one strain and secondary infection by the other strain can occur. There is life-long immunity to the primary infection strain, temporary cross-immunity and after the secondary infection followed by life-long immunity, to the secondary infection strains. Newborns are assumed susceptible. Antibody Dependent Enhancement (ade) is a mechanism where the pre-existing antibodies to the previous dengue infection do not neutralize but rather enhance replication of the secondary strain. In the previously studied models the two strains are identical with respect to their epidemiological functioning: that is the epidemiological process parameters of the two strains were assumed equal. As a result the mathematical model possesses a mathematical symmetry property. In this manuscript we study a variant with epidemiological asymmetry between the strains: the force of infection rates differ while all other epidemiological parameters are equal. Comparison with the results for the epidemiologically symmetric model gives insight into its robustness. Numerical bifurcation analysis and simulation techniques including Lyapunov exponent calculation will be used to study the long-term dynamical behaviour of the model. For the single strain system stable endemic equilibria exist and for the two-strain system endemic equilibria, periodic solutions and also chaotic behaviour.

Describing dengue epidemics: Insights from simple mechanistic models.

We present a set of nested models to be applied to dengue fever epidemiology. We perform a qualitative study in order to show how much complexity we really need to add into epidemiological models to be able to describe the fluctuations observed in empirical dengue hemorrhagic fever incidence data offering a promising perspective on inference of parameter values from dengue case notifications.

The role of seasonality and import in a minimalistic multi-strain dengue model capturing differences between primary and secondary infections: complex dynamics and its implications for data analysis.

In many countries in Asia and South-America dengue fever (DF) and dengue hemorrhagic fever (DHF) has become a substantial public health concern leading to serious social-economic costs. Mathematical models describing the transmission of dengue viruses have focussed on the so-called antibody-dependent enhancement (ADE) effect and temporary cross-immunity trying to explain the irregular behavior of dengue epidemics by analyzing available data. However, no systematic investigation of the possible dynamical structures has been performed so far. Our study focuses on a seasonally forced (non-autonomous) model with temporary cross-immunity and possible secondary infection, motivated by dengue fever epidemiology. The notion of at least two different strains is needed in a minimalistic model to describe differences between primary infections, often asymptomatic, and secondary infection, associated with the severe form of the disease. We extend the previously studied non-seasonal (autonomous) model by adding seasonal forcing, mimicking the vectorial dynamics, and a low import of infected individuals, which is realistic in the dynamics of dengue fever epidemics. A comparative study between three different scenarios (non-seasonal, low seasonal and high seasonal with a low import of infected individuals) is performed. The extended models show complex dynamics and qualitatively a good agreement between empirical DHF monitoring data and the obtained model simulation. We discuss the role of seasonal forcing and the import of infected individuals in such systems, the biological relevance and its implications for the analysis of the available dengue data. At the moment only such minimalistic models have a chance to be qualitatively understood well and eventually tested against existing data. The simplicity of the model (low number of parameters and state variables) offer a promising perspective on parameter values inference from the DHF case notifications.

Parameter Estimation in Epidemiology: from Simple to Complex Dynamics.

We revisit the parameter estimation framework for population biological dynamical systems, and apply it to calibrate various models in epidemiology with empirical time series, namely influenza and dengue fever. When it comes to more complex models like multi-strain dynamics to describe the virus-host interaction in dengue fever, even most recently developed parameter estimation techniques, like maximum likelihood iterated filtering, come to their computational limits. However, the first results of parameter estimation with data on dengue fever from Thailand indicate a subtle interplay between stochasticity and deterministic skeleton. The deterministic system on its own already displays complex dynamics up to deterministic chaos and coexistence of multiple attractors.

Food quality in producer-grazer models: a generalized analysis.

Stoichiometric constraints play a role in the dynamics of natural populations but are not explicitly considered in most mathematical models. Recent theoretical works suggest that these constraints can have a significant impact and should not be neglected. However, it is not yet resolved how stoichiometry should be integrated in population dynamical models, as different modeling approaches are found to yield qualitatively different results. Here we investigate a unifying framework that reveals the differences and commonalities between previously proposed models for producer-grazer systems. Our analysis reveals that stoichiometric constraints affect the dynamics mainly by increasing the intraspecific competition between producers and by introducing a variable biomass conversion efficiency. The intraspecific competition has a strongly stabilizing effect on the system, whereas the variable conversion efficiency resulting from a variable food quality is the main determinant for the nature of the instability once destabilization occurs. Only if the food quality is high can an oscillatory instability, as in the classical paradox of enrichment, occur. While the generalized model reveals that the generic insights remain valid in a large class of models, we show that other details such as the specific sequence of bifurcations encountered in enrichment scenarios can depend sensitively on assumptions made in modeling stoichiometric constraints.

Ecological consequences of global bifurcations in some food chain models.

Food chain models of ordinary differential equations (ode's) are often used in ecology to gain insight in the dynamics of populations of species, and the interactions of these species with each other and their environment. One powerful analysis technique is bifurcation analysis, focusing on the changes in long-term (asymptotic) behaviour under parameter variation. For the detection of local bifurcations there exists standardised software, but until quite recently most software did not include any capabilities for the detection and continuation of global bifurcations. We focus here on the occurrence of global bifurcations in four food chain models, and discuss the implications of their occurrence. In two stoichiometric models (one piecewise continuous, one smooth) there exists a homoclinic bifurcation, that results in the disappearance of a limit cycle attractor. Instead, a stable positive equilibrium becomes the global attractor. The models are also capable of bistability. In two three-dimensional models a Shil'nikov homoclinic bifurcation functions as the organising centre of chaos, while tangencies of homoclinic cycle-to-cycle connections 'cut' the chaotic attractors, which is associated with boundary crises. In one model this leads to extinction of the top predator, while in the other model hysteresis occurs. The types of ecological events occurring because of a global bifurcation will be categorized. Global bifurcations are always catastrophic, leading to the disappearance or merging of attractors. However, there is no 1-on-1 coupling between global bifurcation type and the possible ecological consequences. This only emphasizes the importance of including global bifurcations in the analysis of food chain models.

Direct and indirect effects of pollutants on algae and algivorous ciliates in an aquatic indoor microcosm.

An aquatic indoor microcosm was used to study effects of the pesticides parathion-methyl and prometryn on phototrophic flagellates (Cryptomonas sp.) and predatory ciliates (Urotricha furcata). Parathion-methyl caused effects to flagellates and ciliates at the range of low mg L(-1), regardless of whether the organisms were exposed separately or combined in the multi-species test system. Prometryn caused effects on the flagellates at low microg L(-1) concentrations, resulting in a NOEC of 6.9 microg L(-1) in the single-species test and a NOEC of 15.2 microg L(-1) in the multi-species microcosm. For ciliates the NOEC decreased by factor 145 in the multi-species test compared to the NOEC of 2.2 mg L(-1) in the single-species test when exposed to prometryn. The lower NOEC for ciliates exposed to prometryn in the microcosm was most likely caused by an indirect effect due to reduced availability of flagellates as food. The measurement of nutrient concentrations in the test media and organisms facilitated the modelling of effects. The presented aquatic indoor microcosm is considered as a tool which could be standardised and applied as an instrument to provide data for higher tier risk assessment.

Effects of density-dependent migrations on stability of a two-patch predator-prey model.

We consider a predator-prey model in a two-patch environment and assume that migration between patches is faster than prey growth, predator mortality and predator-prey interactions. Prey (resp. predator) migration rates are considered to be predator (resp. prey) density-dependent. Prey leave a patch at a migration rate proportional to the local predator density. Predators leave a patch at a migration rate inversely proportional to local prey population density. Taking advantage of the two different time scales, we use aggregation methods to obtain a reduced (aggregated) model governing the total prey and predator densities. First, we show that for a large class of density-dependent migration rules for predators and prey there exists a unique and stable equilibrium for migration. Second, a numerical bifurcation analysis is presented. We show that bifurcation diagrams obtained from the complete and aggregated models are consistent with each other for reasonable values of the ratio between the two time scales, fast for migration and slow for local demography. Our results show that, under some particular conditions, the density dependence of migrations can generate a limit cycle. Also a co-dim two Bautin bifurcation point is observed in some range of migration parameters and this implies that bistability of an equilibrium and limit cycle is possible.

Heteroclinic orbits indicate overexploitation in predator-prey systems with a strong Allee effect.

Species establishment in a model system in a homogeneous environment can be dependent not only on the parameter setting, but also on the initial conditions of the system. For instance, predator invasion into an established prey population can fail and lead to system collapse, an event referred to as overexploitation. This phenomenon occurs in models with bistability properties, such as strong Allee effects. The Allee effect then prevents easy re-establishment of the prey species. In this paper, we deal with the bifurcation analyses of two previously published predator-prey models with strong Allee effects. We expand the analyses to include not only local, but also global bifurcations. We show the existence of a point-to-point heteroclinic cycle in these models, and discuss numerical techniques for continuation in parameter space. The continuation of such a cycle in two-parameter space forms the boundary of a region in parameter space where the system collapses after predator invasion, i.e. where overexploitation occurs. We argue that the detection and continuation of global bifurcations in these models are of vital importance for the understanding of the model dynamics.

Bifurcation analysis of a predator-prey model with predators using hawk and dove tactics.

Most classical prey-predator models do not take into account the behavioural structure of the population. Usually, the predator and the prey populations are assumed to be homogeneous, i.e. all individuals behave in the same way. In this work, we shall take into account different tactics that predators can use for exploiting a common self-reproducing resource, the prey population. Predators fight together in order to keep or to have access to captured prey individuals. Individual predators can use two behavioural tactics when they encounter to dispute a prey, the classical hawk and dove tactics. We assume two different time scales. The fast time scale corresponds to the inter-specific searching and handling for the prey by the predators and the intra-specific fighting between the predators. The slow time scale corresponds to the (logistic) growth of the prey population and mortality of the predator. We take advantage of the two time scales to reduce the dimension of the model and to obtain an aggregated model that describes the dynamics of the total predator and prey densities at the slow time scale. We present the bifurcation analysis of the model and the effects of the different predator tactics on persistence and stability of the prey-predator community are discussed.