Browsing by Subject "Adaptive dynamics"

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  • Fritsch, Coralie; Campillo, Fabien; Ovaskainen, Otso (2017)
    We propose a numerical approach to study the invasion fitness of a mutant and to determine evolutionary singular strategies in evolutionary structured models in which the competitive exclusion principle holds. Our approach is based on a dual representation, which consists of the modeling of the small size mutant population by a stochastic model and the computation of its corresponding deterministic model. The use of the deterministic model greatly facilitates the numerical determination of the feasibility of invasion as well as the convergence-stability of the evolutionary singular strategy. Our approach combines standard adaptive dynamics with the link between the mutant survival criterion in the stochastic model and the sign of the eigenvalue in the corresponding deterministic model. We present our method in the context of a mass-structured individual-based chemostat model. We exploit a previously derived mathematical relationship between stochastic and deterministic representations of the mutant population in the chemostat model to derive a general numerical method for analyzing the invasion fitness in the stochastic models. Our method can be applied to the broad class of evolutionary models for which a link between the stochastic and deterministic invasion fitnesses can be established. (C) 2017 Elsevier Inc. All rights reserved.
  • Kisdi, Eva; Geritz, Stefan A.H. (2016)
    We study the joint adaptive dynamics of n scalar-valued strategies in ecosystems where n is the maximum number of coexisting strategies permitted by the (generalized) competitive exclusion principle. The adaptive dynamics of such saturated systems exhibits special characteristics, which we first demonstrate in a simple example of a host-pathogen-predator model. The main part of the paper characterizes the adaptive dynamics of saturated polymorphisms in general. In order to investigate convergence stability, we give a new sufficient condition for absolute stability of an arbitrary (not necessarily saturated) polymorphic singularity and show that saturated evolutionarily stable polymorphisms satisfy it. For the case , we also introduce a method to construct different pairwise invasibility plots of the monomorphic population without changing the selection gradients of the saturated dimorphism.
  • Lehtinen, Sami O.; Geritz, Stefan A.H. (2019)
    We investigate the coevolution of cannibalistic predators and timid prey, which seek refuge upon detecting a predator. To understand how the species affect each other’s evolution, we derived the ecological model from individual-level processes using ordinary differential equations. The ecological dynamics exhibit bistability between equilibrium and periodic attractors, which may disappear through catastrophic bifurcations. Using the critical function analysis of adaptive dynamics, we classify general trade-offs between cannibalism and prey capture that produce different evolutionary outcomes. The evolutionary analysis reveals several ways in which cannibalism emerges as a response to timidity of the prey. The long-term coevolution either attains a singularity, or becomes cyclic through two mechanisms: genetical cycles through Hopf bifurcation of the singularity, or ecogenetical cycles involving abrupt switching between ecological attractors. Further diversification of cannibalism occurs through evolutionary branching, which is predicted to be delayed when simultaneous prey evolution is necessary for the singularity’s attainability. We conclude that predator-prey coevolution produces a variety of outcomes, in which evolutionary cycles are commonplace.
  • Berardo, Cecilia; Geritz, Stefanus (2021)
    The war of attrition in game theory is a model of a stand-off situation between two opponents where the winner is determined by its persistence. We model a stand-off between a predator and a prey when the prey is hiding and the predator is waiting for the prey to come out from its refuge, or when the two are locked in a situation of mutual threat of injury or even death. The stand-off is resolved when the predator gives up or when the prey tries to escape. Instead of using the asymmetric war of attrition, we embed the stand-off as an integral part of the predator-prey model of Rosenzweig and MacArthur derived from first principles. We apply this model to study the coevolution of the giving-up rates of the prey and the predator, using the adaptive dynamics approach. We find that the long term evolutionary process leads to three qualitatively different scenarios: the predator gives up immediately, while the prey never gives up; the predator never gives up, while the prey adopts any giving-up rate greater than or equal to a given positive threshold value; the predator goes extinct. We observe that some results are the same as for the asymmetric war of attrition, but others are quite different. (C) 2021 The Author(s). Published by Elsevier Ltd.
  • Lehtinen, Sami O.; Geritz, Stefanus A.H. (2019)
    We investigate the evolution of timidity in a prey species whose predator has cannibalistic tendencies. The ecological model is derived from individual-level processes, in which the prey seeks refuge after detecting a predator, and the predator cannibalises on the conspecific juveniles. Bifurcation analysis of the model reveals ecological bistability between equilibrium and periodic attractors. Using the framework of adaptive dynamics, we classify ten qualitatively different evolutionary scenarios induced by the ecological bistability. These scenarios include ecological attractor switching through catastrophic bifurcations, which can reverse the direction of evolution. We show that such reversals often result in evolutionary cycling of the level of timidity. In the absence of cannibalism, the model never exhibits ecological bistability nor evolutionary cycling. We conclude that cannibalistic predator behaviour can completely change both the ecological dynamics and the evolution of prey. (C) 2019 The Authors. Published by Elsevier Ltd.
  • Lehtinen, Sami (2021)
    In many terrestrial, marine, and freshwater predator-prey communities, young predators can be vulnerable to attacks by large prey. Frequent prey counter-attacks may hinder the persistence of predators. Despite the commonness of such role reversals in nature, they have rarely been addressed in evolutionary modelling. To understand how role reversals affect ecological and evolutionary dynamics of a predatorprey community, we derived an ecological model from individual-level processes using ordinary differential equations. The model reveals complex ecological dynamics, with possible bistability between alternative coexistence states and an Allee effect for the predators. We find that when prey counter-attacks are frequent, cannibalism is necessary for predator persistence. Using numerical analysis, we also find that a sudden ecological shift from coexistence to predator extinction can occur through several catastrophic bifurcations, including 'saddle-node', 'homoclinic', and 'subcritical Hopf'. The analysis of single-species evolution reveals that predator selection towards increasing or decreasing cannibalism triggers a catastrophic shift towards an extinction state of the predators. Such an evolutionary extinction of the predators may also be caused by prey selection towards increasing foraging activity because it facilitates encounters with vulnerable, young predators. The analysis of predator-prey coevolution further demonstrates that predator's catastrophic extinction becomes an even more likely outcome than in single-species evolution. Our results suggest that when young predators are vulnerable to prey attacks, a sudden extinction of the predators may be more common than currently understood. (C) 2020 The Author(s). Published by Elsevier Ltd.
  • Weigang, Helene C.; Kisdi, Eva (2015)
    Resources invested in dispersal structures as well as time and energy spent during transfer may often decrease fecundity. Here we analyse an extended version of the Hamilton-May model of dispersal evolution, where we include a fecundity-dispersal trade-off and also mortality between competition and reproduction. With adaptive dynamics and critical function analysis we investigate the evolution of dispersal strategies and ask whether adaptive diversification is possible. We exclude evolutionary branching for concave trade-offs and show that for convex trade-offs diversification is promoted in a narrow parameter range. We provide theoretical evidence that dispersal strategies can monotonically decrease with increasing survival during dispersal. Moreover, we illustrate the existence of two alternative attracting dispersal strategies. The model exhibits fold bifurcation points where slight changes in survival can lead to evolutionary catastrophes. (C) 2015 Elsevier Ltd. All rights reserved.
  • Karisto, Petteri; Kisdi, Eva (2017)
    The pattern of connectivity between local populations or between microsites supporting individuals within a population is a poorly understood factor affecting the evolution of dispersal. We modify the well-known Hamilton May model of dispersal evolution to allow for variable connectivity between microsites. For simplicity, we assume that the microsites are either solitary, i.e., weakly connected through costly dispersal, or part of a well-connected cluster of sites with low-cost dispersal within the cluster. We use adaptive dynamics to investigate the evolution of dispersal, obtaining analytic results for monomorphic evolution and numerical results for the co-evolution of two dispersal strategies. A monomorphic population always evolves to a unique singular dispersal strategy, which may be an evolutionarily stable strategy or an evolutionary branching point. Evolutionary branching happens if the contrast between connectivities is sufficiently high and the solitary microsites are common. The dimorphic evolutionary singularity, when it exists, is always evolutionarily and convergence stable. The model exhibits both protected and unprotected dimorphisms of dispersal strategies, but the dimorphic singularity is always protected. Contrasting connectivities can thus maintain dispersal polymorphisms in temporally stable environments.
  • Cai, Yuhua (2022)
    We study the adaptive dynamics of the colonization rate of species living in a patchy habitat when there is a trade-off with the competitive strength for individual patches. To that end, we formulate a continuous-time competition-colonization model that also includes ownership effects as well as random disturbance affecting the mortality rate. We find that intermediate disturbance (as measured by the fluctuation intensity of the mortality rate), a strong competition-colonization trade-off, and a weak ownership effect are necessary conditions for evolutionary branching and hence for the emergence of polymorphisms (i.e., coexistence) by small evolutionary steps. Specifically, concerning ownership we find that with low-intermediate disturbance, a weak ownership advantage favours evolutionary branching while ownership disadvantage does not. This asymmetry disappears at the higher-intermediate disturbance. Moreover, at a low-intermediate disturbance, the effect of the strength of the competition-colonization trade-off on evolutionary branching is non-monotonic disappears because the possibility of branching disappears again when the trade-off is too strong. We also find that there can be multiple evolutionary attractors for polymorphic populations, each with its own basin of attraction. With small but non-zero random evolutionary steps and depending on the initial polymorphic condition just after branching, a coevolutionary trajectory may come arbitrarily close to the shared boundary of two such basins and may even jump from one side to the other, which can lead to various kinds of long-term evolutionary dynamics, including evolutionary branching-extinction cycles. (C) 2021 The Author(s). Published by Elsevier Ltd.
  • Vitale, Caterina; Kisdi, Eva (2019)
    Under the threat of predation, a species of prey can evolve to its own extinction. Matsuda and Abrams (Theor Popul Biol 45:76-91, 1994a) found the earliest example of evolutionary suicide by demonstrating that the foraging effort of prey can evolve until its population dynamics cross a fold bifurcation, whereupon the prey crashes to extinction. We extend this model in three directions. First, we use critical function analysis to show that extinction cannot happen via increasing foraging effort. Second, we extend the model to non-equilibrium systems and demonstrate evolutionary suicide at a fold bifurcation of limit cycles. Third, we relax a crucial assumption of the original model. To find evolutionary suicide, Matsuda and Abrams assumed a generalist predator, whose population size is fixed independently of the focal prey. We embed the original model into a three-species community of the focal prey, the predator and an alternative prey that can support the predator also alone, and investigate the effect of increasingly strong coupling between the focal prey and the predator's population dynamics. Our three-species model exhibits (1) evolutionary suicide via a subcritical Hopf bifurcation and (2) indirect evolutionary suicide, where the evolution of the focal prey first makes the community open to the invasion of the alternative prey, which in turn makes evolutionary suicide of the focal prey possible. These new phenomena highlight the importance of studying evolution in a broader community context.
  • Boldin, Barbara; Kisdi, Eva (2016)
    Evolutionary suicide is a riveting phenomenon in which adaptive evolution drives a viable population to extinction. Gyllenberg and Parvinen (Bull Math Biol 63(5):981-993, 2001) showed that, in a wide class of deterministic population models, a discontinuous transition to extinction is a necessary condition for evolutionary suicide. An implicit assumption of their proof is that the invasion fitness of a rare strategy is well-defined also in the extinction state of the population. Epidemic models with frequency-dependent incidence, which are often used to model the spread of sexually transmitted infections or the dynamics of infectious diseases within herds, violate this assumption. In these models, evolutionary suicide can occur through a non-catastrophic bifurcation whereby pathogen adaptation leads to a continuous decline of host (and consequently pathogen) population size to zero. Evolutionary suicide of pathogens with frequency-dependent transmission can occur in two ways, with pathogen strains evolving either higher or lower virulence.
  • Karisto, Petteri; Kisdi, Èva (2019)
    Functional connectivity, the realized flow of individuals between the suitable sites of a heterogeneous landscape, is a prime determinant of the maintenance and evolution of populations in fragmented habitats. While a large body of literature examines the evolution of dispersal propensity, it is less known how evolution shapes functional connectivity via traits that influence the distribution of the dispersers. Here, we use a simple model to demonstrate that, in a heterogeneous environment with clustered and solitary sites (i.e., with variable structural connectivity), the evolutionarily stable population contains strains that are strongly differentiated in their pattern of connectivity (local vs. global dispersal), but not necessarily in the fraction of dispersed individuals. Also during evolutionary branching, selection is disruptive predominantly on the pattern of connectivity rather than on dispersal propensity itself. Our model predicts diversification along a hitherto neglected axis of dispersal strategies and highlights the role of the solitary sites-the more isolated and therefore seemingly less important patches of habitat-in maintaining global dispersal that keeps all sites connected.
  • Gyllenberg, Mats; Kisdi, Eva; Weigang, Helene C. (2016)
    Empirical studies of dispersal indicate that decisions to immigrate are patch-type dependent; yet theoretical models usually ignore this fact. Here, we investigate the evolution of patch-type dependent immigration of a population inhabiting and dispersing in a heterogeneous landscape, which is structured by patches of low and high reward. We model the decision to immigrate in detail from a mechanistic underpinning. With the methods of adaptive dynamics, we derive both analytical and numerical results for the evolution of immigration when life-history traits are patch-type dependent. The model exhibits evolutionary branching in a wide parameter range and the subsequent coevolution can lead to a stable coexistence of a generalist, settling in patches of any type, and a specialist that only immigrates into patches of high reward. We find that individuals always settle in the patches of high reward, in which survival until maturation, relative fecundity and emigration probability are high. We investigate how the probability to immigrate into patches of low reward changes with model parameters. For example, we show that immigration into patches of low reward increases when the emigration probability in these patches increases. Further, immigration into patches of low reward decreases when the patches of high reward become less safe during the dispersal season. (C) 2016 Elsevier Ltd. All rights reserved.
  • Cai, Yuhua; Geritz, Stefanus (2020)
    We study resident-invader dynamics in fluctuating environments when the invader and the resident have close but distinct strategies. First we focus on a class of continuous-time models of unstructured populations of multi-dimensional strategies, which incorporates environmental feedback and environmental stochasticity. Then we generalize our results to a class of structured population models. We classify the generic population dynamical outcomes of an invasion event when the resident population in a given environment is non-growing on the long-run and stochastically persistent. Our approach is based on the series expansion of a model with respect to the small strategy difference, and on the analysis of a stochastic fast-slow system induced by time-scale separation. Theoretical and numerical analyses show that the total size of the resident and invader population varies stochastically and dramatically in time, while the relative size of the invader population changes slowly and asymptotically in time. Thereby the classification is based on the asymptotic behavior of the relative population size, and which is shown to be fully determined by invasion criteria (i.e., without having to study the full generic dynamical system). Our results extend and generalize previous results for a stable resident equilibrium (particularly, Geritz in J Math Biol 50(1):67-82, 2005; Dercole and Geritz in J Theor Biol 394:231-254, 2016) to non-equilibrium resident population dynamics as well as resident dynamics with stochastic (or deterministic) drivers.
  • Cai, Yuhua; Geritz, Stefanus (2021)
    To better understand the environmental factors and ecological processes underlying the evolution of the irreversible transition from a free-swimming state to an immobile sessile state as seen in many aquatic invertebrates, we study the adaptive dynamics of the settling rate of a hypothetical microorganism onto the wall of a chemostat. The two states, floating or settled, differ in their nutrient ingestion, reproduction and death rate. We consider three different settling mechanisms involving competition for space on the wall: (i) purely exploitative competition where free-swimming individuals settle in vacant space only, (ii) mixed exploitative and interference competition where individuals attempt to settle in any place but fail and die if the space is already occupied, and (iii) mixed exploitative and interference competition, but now settling in occupied space is successful and the former occupant dies. In the simplified environment of the chemostat, the input concentration of nutrients and the dilution rate of the tank are the main environmental control variables. Using the theory of adaptive dynamics, we find that the settling mechanisms and environmental control variables have qualitatively different effects on the evolution of the settling rate in terms of the direction of evolution as well as on species diversity. In the case of purely exploitative competition a small change in the settings of the environmental control variables can lead to an abrupt reversal of the direction of evolution, while in the case of mixed exploitative and interference competition the effect is gradual. For all three settling mechanisms, periodic fluctuations in the nutrient input open the possibility of evolutionary branching leading to the long-term coexistence of an intermediate and an infinitely high settling rates (in the case of low-frequency fluctuations), and an intermediate and a zero settling rates (in the case of high-frequency fluctuations). (C) 2021 The Author(s). Published by Elsevier Ltd.
  • Fan, Ruili; Geritz, Stefan A H (2021)
    We study the evolution of virulence of an endemic pathogen in response to healthcare interventions which affect host recovery and pathogen transmission. By anticipating the evolutionary response of the pathogen we may develop effective long-term management strategies for controlling the impact of the endemic on the society. To that end, we use standard Adaptive Dynamics techniques in an SIS model. The recovery rate and the transmission rate, both of which can be affected by healthcare interventions, are used as evolutionary control variables. The effect of interventions may be density-independent (self-help based on healthcare instructions) or density-dependent (when assistance of a healthcare worker is required). We consider the evolutionary response of the pathogen both to abrupt changes and to gradual changes in the level of healthcare intervention. Healthcare intervention is optimised for three alternative objectives: minimisation of virulence, minimisation of the probability that an infected individual dies of the disease, and total eradication of the endemic. We find that the optimal strategy may depend on the objective. High levels of healthcare intervention may eradicate the pathogen, but this option may not be available for budgetary reasons or otherwise. Counterintuitively, to minimise virulence, one should keep healthcare interventions at a minimum, while to minimise the probability for an infected individual to die of the disease, both low and high levels of healthcare intervention suffice. Changes in the level of healthcare intervention should be implemented fast (not gradually) in order to avoid sudden changes in pathogen evolution and the possible emergence of multiple simultaneously coexisting pathogen strains. (C) 2021 The Author(s). Published by Elsevier Ltd.