How parasitism affects critical patch-size in a host-parasitoid model: application to the forest tent caterpillar

Habitat structure has broad impacts on many biological systems. In particular, habitat fragmentation can increase the probability of species extinction and on the other hand it can lead to population outbreaks in response to a decline in natural enemies. An extreme consequence of fragmentation is the isolation of small regions of suitable habitat surrounded by a large region of hostile matrix. This scenario can be interpreted as a critical patch-size problem, well studied in a continuous time framework, but relatively new to discrete time models. In this paper we present an integrodifference host-parasitoid model, discrete in time and continuous in space, to study how the critical habitat-size necessary for parasitoid survival changes in response to parasitoid life history traits, such as emergence time. We show that early emerging parasitoids may be able to persist in smaller habitats than late emerging species. The model predicts that these early emerging parasitoids lead to more severe host outbreaks. We hypothesise that promoting efficient late emerging parasitoids may be key in reducing outbreak severity, an approach requiring large continuous regions of suitable habitat. We parameterise the model for the host species of the forest tent caterpillar Malacosoma disstria Hbn., a pest insect for which fragmented landscape increases the severity of outbreaks. This host is known to have several parasitoids, due to paucity of data and as a first step in the modelling we consider a single generic parasitoid. The model findings are related to observations of the forest tent caterpillar offering insight into this host-parasitoid response to habitat structure.     

宿主-寄生物相互作用种群模型中的混沌与分形现象

自然界具有离散世代的宿主-寄生物相互作用的种群模型都以差分方程组来描述,由于宿主-寄生物相互关系具有不同的类型,模型的功能反应函数也具有多种形式.通过数值模拟试验,分析研究了聚集效应模型随参数变化时表现出的混沌动态行为以及吸引域的自相似分形属性.结果显示虽然在一定参数范围内种群数量显示严格的周期,但混沌动态是不可避免的,共存的多吸引子初值区域显示出自相似分形属性.说明混沌动态行为和分形属性是离散的种群互作用种群模型中必然出现的现象.     

Strange limits of stability in host-parasitoid systems

The classical Nicholson-Bailey model for a two species host-parasitoid system with discrete generations assumes random distributions of both hosts and parasitoids, randomly searching parasitoids, and random encounters between the individuals of the two species. Although unstable, this model induced many investigations into more complex host-parasitoid systems. Local linearized stability analysis shows that equilibria of host parasitoid systems within the framework of a generalized Nicholson-Bailey model are generally unstable. Stability is only possible if host fertility does not exceede ⁴=54.5982 and if superparasitism is unsuccessful. This special situation has already been discovered by Hassell et al. (1983) in their study of the effects of variable sex ratios on host parasitoid dynamics. We discuss global behaviour of the Hassell-Waage-May model using KAM-theory and illustrate its sensitivity to small perturbations, which can give rise to radically different patterns of the population dynamics of interacting hosts and parasitoids.     

Semi-discrete host-parasitoid models

Arthropod host-parasitoid interactions constitute a very important class of consumer resource dynamics. Discrete-time models are a tradition for such interactions and are characterized by an updating function, which relates the population densities at a fixed date in one year to those at the same date in the previous year. Previous workers have investigated the effects of functional response and density dependence on the stability of the host-parasitoid interaction by heuristically incorporating them in the updating function. Such an approach ignores the effects of population changing continuously within a year due to different processes (for example intraspecific competition, mortality from parasitism) that may act simultaneously. Their cumulative effect on the updating function is not obvious and a more systematic methodology is needed. This paper uses a hybrid approach to formulate the updating function. This is done by modeling the dynamics of various within-year processes in continuous-time, and reproduction as a discrete event. Using this formalism we derive results connecting the stability of the host-parasitoid interaction with different forms of density dependence and the form of the functional response. The latter results contradict previous conclusions from heuristically formulated models, and illustrate the need for such a hybrid approach in discrete-time host-parasitoid theory.     

Competition and dispersal in predator-prey waves

Dispersing predators and prey can exhibit complex spatio-temporal wave-like patterns if the interactions between them cause oscillatory dynamics. We study the effect of these predator-prey density waves on the competition between prey populations and between predator populations with different dispersal strategies. We first describe 1- and 2-dimensional simulations of both discrete and continuous predator-prey models. The results suggest that any population that diffuses faster, disperses farther, or is more likely to disperse will exclude slower diffusing, shorter dispersing, or less likely dispersing populations, everything else being equal. It also appears that it does not matter whether time, space, or state are discrete or continuous, nor what the exact interactions between the predators and prey are. So long as waves exist the competition between populations occurs in a similar fashion. We derive a theory that qualitatively explains the observed behaviour and calculate approximate analytical solutions that describe, to a reasonable extent, these behaviours. Predictions about the cost of dispersal are tested. If strong enough, cost can reverse the populations' relative competitive strengths or lead to coexistence because of the effect of spiral wave cores. The theory is also able to explain previous results of simulations of coexistence in host-parasitoid models (Comins, H. N., and Massell, M. P., 1996, J. Theor. Biol. 183, 19-28).     

Discrete-time host-parasitoid models with pest control

We propose a simple discrete-time host-parasitoid model to investigate the impact of external input of parasitoids upon the host-parasitoid interactions. It is proved that the input of the external parasitoids can eventually eliminate the host population if it is above a threshold and it also decreases the host population level in the unique interior equilibrium. It can simplify the host-parasitoid dynamics when the host population practices contest competition. We then consider a corresponding optimal control problem over a finite time period. We also derive an optimal control model using a chemical as a control for the hosts. Applying the forward-backward sweep method, we solve the optimal control problems numerically and compare the optimal host populations with the host populations when no control is applied. Our study concludes that applying a chemical to eliminate the hosts directly may be a more effective control strategy than using the parasitoids to indirectly suppress the hosts.     

Models for integrated pest control and their biological implications

Successful integrated pest management (IPM) control programmes depend on many factors which include host-parasitoid ratios, starting densities, timings of parasitoid releases, dosages and timings of insecticide applications and levels of host-feeding and parasitism. Mathematical models can help us to clarify and predict the effects of such factors on the stability of host-parasitoid systems, which we illustrate here by extending the classical continuous and discrete host-parasitoid models to include an IPM control programme. The results indicate that one of three control methods can maintain the host level below the economic threshold (ET) in relation to different ET levels, initial densities of host and parasitoid populations and host-parasitoid ratios. The effects of host intrinsic growth rate and parasitoid searching efficiency on host mean outbreak period can be calculated numerically from the models presented. The instantaneous pest killing rate of an insecticide application is also estimated from the models. The results imply that the modelling methods described can help in the design of appropriate control strategies and assist management decision-making. The results also indicate that a high initial density of parasitoids (such as in inundative releases) and high parasitoid inter-generational survival rates will lead to more frequent host outbreaks and, therefore, greater economic damage. The biological implications of this counter intuitive result are discussed.     

Spatial synchrony in host-parasitoid models using aggregation of variables

We consider a host-parasitoid system with individuals moving on a square grid of patches. We study the effects of increasing movement frequency of hosts and parasitoids on the spatial dynamics of the system. We show that there exists a threshold value of movement frequency above which spatial synchrony occurs and the dynamics of the system can be described by an aggregated model governing the total population densities on the grid. Numerical simulations show that this threshold value is usually small. This allows using the aggregated model to make valid predictions about global host-parasitoid spatial dynamics.     

Host-Parasitoid Dynamics and the Success of Biological Control When Parasitoids Are Prone to Allee Effects

In sexual organisms, low population density can result in mating failures and subsequently yields a low population growth rate and high chance of extinction. For species that are in tight interaction, as in host-parasitoid systems, population dynamics are primarily constrained by demographic interdependences, so that mating failures may have much more intricate consequences. Our main objective is to study the demographic consequences of parasitoid mating failures at low density and its consequences on the success of biological control. For this, we developed a deterministic host-parasitoid model with a mate-finding Allee effect, allowing to tackle interactions between the Allee effect and key determinants of host-parasitoid demography such as the distribution of parasitoid attacks and host competition. Our study shows that parasitoid mating failures at low density result in an extinction threshold and increase the domain of parasitoid deterministic extinction. When proned to mate finding difficulties, parasitoids with cyclic dynamics or low searching efficiency go extinct; parasitoids with high searching efficiency may either persist or go extinct, depending on host intraspecific competition. We show that parasitoids suitable as biocontrol agents for their ability to reduce host populations are particularly likely to suffer from mate-finding Allee effects. This study highlights novel perspectives for understanding of the dynamics observed in natural host-parasitoid systems and improving the success of parasitoid introductions.     

Migration frequency and the persistence of host-parasitoid interactions

This paper analyses the effect of migration frequency on the stability and persistence of a host-parasitoid system in a two-patch environment. The hosts and parasitoids are allowed to move from one patch to the other a certain number of times within a generation. When this number is low, i.e. when the time-scales associated with migration and demography are of the same order, host-parasitoid interactions are usually not persistent. When this number is high, however, persistence is more likely. Moreover, in this situation, aggregation methods can be used to simplify the proposed initial model into an aggregated model describing the dynamics of both the total host and parasitoid populations. Analysis of the aggregated model shows that the system reaches a stable steady state for some regions of the parameter domain. Persistence occurs when the movement of the parasitoids is asymmetrical, i.e. they move preferentially to one of the two patches. We show that the growth rate of the host population is a key parameter in determining which migration strategies of the parasitoids lead to persistent host-parasitoid interactions.     

Influence of parasitized adult reproduction on host-parasitoid dynamics: an age-structured model

In this work, we develop an age-structured model (based on delay-differential equations) to investigate the dynamics of host-parasitoid systems in which adults are the target of parasitism. The rare previous work dealing with such interactions assumes that hosts are functionally dead as soon as they are attacked. We relax this assumption and show that low reproduction rates of parasitized hosts can promote stability at the expense of cyclic behavior (either long term or generation cycles). Higher reproduction rates make the regulation of the host population by parasitoids impossible. As it is the case in models in which adults are subjected to attacks but do not reproduce, our model generates generation cycles for a larger set of parameter values than in models in which juveniles are attacked.     

Relating coupled map lattices to integro-difference equations: dispersal-driven instabilities in coupled map lattices

Recently there has been a great deal of interest within the ecological community about the interactions of local populations that are coupled only by dispersal. Models have been developed to consider such scenarios but the theory needed to validate model outcomes has been somewhat lacking. In this paper, we present theory which can be used to understand these types of interaction when population exhibit discrete time dynamics. In particular, we consider a spatial extension to discrete-time models, known as coupled map lattices (CMLs) which are discrete in space. We introduce a general form of the CML and link this to integro-difference equations via a special redistribution kernel. General conditions are then derived for dispersal-driven instabilities. We then apply this theory to two discrete-time models; a predator-prey model and a host-pathogen model.     

Epidemiological landscape models reproduce cyclic insect outbreaks

Forest insect outbreaks can have large impacts on ecosystems and understanding the underlying ecological processes is critical for their management. Current process-based modeling approaches of insect outbreaks are often based on population processes operating at small spatial scales (i.e. within individual forest stands). As such, they are difficult to parameterize and offer limited applicability when modeling and predicting outbreaks at the landscape level where management actions take place. In this paper, we propose a new process-based landscape model of forest insect outbreaks that is based on stand defoliation, the Forest-Infected-Recovering-Forest (FIRF) model. We explore both spatially-implicit (mean field equations with global dispersal) and spatially-explicit (cellular automata with limited dispersal between neighboring stands) versions of this model to assess the role of dispersal in the landscape dynamics of outbreaks. We show that density-dependent dispersal is necessary to generate cyclic outbreaks in the spatially-implicit version of the model. The spatially-explicit FIRF model with local and stochastic dispersal displays cyclic outbreaks at the landscape scale and patchy outbreaks in space, even without density-dependence. Our simple, process-based FIRF model reproduces large scale outbreaks and can provide an innovative approach to model and manage forest pests at the landscape scale.     

Dynamic heterogeneous spatio-temporal pattern formation in host-parasitoid systems with synchronised generations

In this paper we develop a general mathematical model describing the spatio-temporal dynamics of host-parasitoid systems with forced generational synchronisation, for example seasonally induced diapause. The model itself may be described as an individual-based stochastic model with the individual movement rules derived from an underlying continuum PDE model. This approach permits direct comparison between the discrete model and the continuum model. The model includes both within-generation and between-generation mechanisms for population regulation and focuses on the interactions between immobile juvenile hosts, adult hosts and adult parasitoids in a two-dimensional domain. These interactions are mediated, as they are in many such host-parasitoid systems, by the presence of a volatile semio-chemical (kairomone) emitted by the hosts or the hosts food plant. The model investigates the effects on population dynamics for different host versus parasitoid movement strategies as well as the transient dynamics leading to steady states. Despite some agreement between the individual and continuum models for certain motility parameter ranges, the model dynamics diverge when host and parasitoid motilities are unequal. The individual-based model maintains spatially heterogeneous oscillatory dynamics when the continuum model predicts a homogeneous steady state. We discuss the implications of these results for mechanistic models of phenotype evolution.P. Schofield gratefully acknowledges the financial support of the BBSRC and The Wellcome Trust.     

Dynamical consequences of optimal host feeding on host-parasitoid population dynamics

This study examines the influence of various host-feeding patterns on host-parasitoid population dynamics. The following types of host-feeding patterns are considered: concurrent and non-destructive, non-concurrent and non-destructive, and non-concurrent and destructive. The host-parasitoid population dynamics is described by the Lotka-Volterra continuous-time model. This study shows that when parasitoids behave optimally, i.e. they maximize their fitness measured by the instantaneous per capita growth rate, the non-destructive type of host feeding stabilizes host-parasitoid dynamics. Other types of host feeding, i.e. destructive, concurrent, or non-concurrent, do not qualitatively change the neutral stability of the Lotka-Volterra model. Moreover, it is shown that the pattern of host feeding which maximizes parasitoid fitness is either non-concurrent and destructive, or concurrent and non-destructive host feeding, depending on the host abundance and parameters of the model. The effects of the adaptive choice of host-feeding patterns on host-parasitoid population dynamics are discussed.     

Stability analysis of the time delay in a host-parasitoid model

The effect of a time delay on the local stability of a host-parasitoid model is analyzed. The delay is between the time of parasitization of the host and the emergence of the parasitoid from the host. Both analytic methods and computer simulations are used in this study. By linearizing and transforming the original equations, sufficient conditions for the local stability are found. In the case of the parameters considered, the results illustrate the destabilizing effect of the time delay. As the lag increases the number of stable points decreases and the points become more scattered in the parameter space. Simulations of the original model are also produced. The region of stability indicated by the simulations is greater than that predicted by the use of the analytic technique. The analysis also reveals the impact of the population parameters upon the stability of the time delay model.The importance of understanding time lags is discussed with reference to population regulation.     

Stability and invariant manifolds of a generalized Beddington host-parasitoid model

We will investigate the stability and invariant manifolds of a new discrete host-parasitoid model. It is a generalization of the Beddington–Nicholson–Bailey model. Our study establishes analytically, for the first time, the stability of the coexistence fixed point.     

Modelling density-dependent resistance in insect-pathogen interactions.

We consider a mathematical model for a host-pathogen interaction where the host population is split into two categories: those susceptible to disease and those resistant to disease. Since the model was motivated by studies on insect populations, we consider a discrete-time model to reflect the discrete generations which are common among insect species. Whether an individual is born susceptible or resistant to disease depends on the local population levels at the start of each generation. In particular, we are interested in the case where the fraction of resistant individuals in the population increases as the total population increases. This may be seen as a positive feedback mechanism since disease is the only population control imposed upon the system. Moreover, it reflects recent experimental observations from noctuid moth-baculovirus interactions that pathogen resistance may increase with larval density. We find that the inclusion of a resistant class can stabilise unstable host-pathogen interactions but there is greatest regulation when the fraction born resistant is density independent. Nonetheless, inclusion of density dependence can still allow intrinsically unstable host-pathogen dynamics to be stabilised provided that this effect is sufficiently small. Moreover, inclusion of density-dependent resistance to disease allows the system to give rise to bistable dynamics in which the final outcome is dictated by the initial conditions for the model system. This has implications for the management of agricultural pests using biocontrol agents-in particular, it is suggested that the propensity for density-dependent resistance be determined prior to such a biocontrol attempt in order to be sure that this will result in the prevention of pest outbreaks, rather than their facilitation. Finally we consider how the cost of resistance to disease affects model outcomes and discover that when there is no cost to resistance, the model predicts stable periodic outbreaks of the insect population. The results are interpreted ecologically and future avenues for research to address the shortfalls in the present model system are discussed.     

Host-parasitoid spatial dynamics in heterogeneous landscapes

This paper explores the effect of spatial processes in a heterogeneous environment on the dynamics of a host-parasitoid interaction. The environment consists of a lattice of favourable (habitat) and hostile (matrix) hexagonal cells, whose spatial distribution is measured by habitat proportion and spatial autocorrelation (inverse of fragmentation). At each time step, a fixed fraction of both populations disperses to the adjacent cells where it reproduces following the Nicholson-Bailey model. Aspects of the dynamics analysed include extinction, stability, cycle period and amplitude, and the spatial patterns emerging from the dynamics.
We find that, depending primarily on the fraction of the host population that disperses in each generation and on the landscape geometry, five classes of spatio-temporal dynamics can be objectively distinguished: spatial chaos, spirals, metapopulation, mainland-island and spiral fragments. The first two are commonly found in theoretical studies of homogeneous landscapes. The other three are direct consequences of the heterogeneity and have strong similarities to dynamic patterns observed in real systems (e.g. extinction-recolonisation, source-sink, outbreaks, spreading waves).
We discuss the processes that generate these patterns and allow the system to persist. The importance of these results is threefold: first, our model merges into a same theoretical framework dynamics commonly observed in the field that are usually modelled independently. Second, these dynamics and patterns are explained by dispersal rate and common landscape statistics, thus linking in a practical way population ecology to landscape ecology. Third, we show that the landscape geometry has a qualitative effect on the length of the cycles and, in particular, we demonstrate how very long periods can be produced by spatial processes.