Fatal disease and demographic Allee effect: population persistence and extinction

If a healthy stable host population at the disease-free equilibrium is subject to the Allee effect, can a small number of infected individuals with a fatal disease cause the host population to go extinct? That is, does the Allee effect matter at high densities? To answer this question, we use a susceptible–infected epidemic model to obtain model parameters that lead to host population persistence (with or without infected individuals) and to host extinction. We prove that the presence of an Allee effect in host demographics matters even at large population densities. We show that a small perturbation to the disease-free equilibrium can eventually lead to host population extinction. In addition, we prove that additional deaths due to a fatal infectious disease effectively increase the Allee threshold of the host population demographics.     

A diffusive SI model with Allee effect and application to FIV

A minimal reaction-diffusion model for the spatiotemporal spread of an infectious disease is considered. The model is motivated by the Feline Immunodeficiency Virus (FIV) which causes AIDS in cat populations. Because the infected period is long compared with the lifespan, the model incorporates the host population growth. Two different types are considered: logistic growth and growth with a strong Allee effect. In the model with logistic growth, the introduced disease propagates in form of a travelling infection wave with a constant asymptotic rate of spread. In the model with Allee effect the spatiotemporal dynamics are more complicated and the disease has considerable impact on the host population spread. Most importantly, there are waves of extinction, which arise when the disease is introduced in the wake of the invading host population. These waves of extinction destabilize locally stable endemic coexistence states. Moreover, spatially restricted epidemics are possible as well as travelling infection pulses that correspond either to fatal epidemics with succeeding host population extinction or to epidemics with recovery of the host population. Generally, the Allee effect induces minimum viable population sizes and critical spatial lengths of the initial distribution. The local stability analysis yields bistability and the phenomenon of transient epidemics within the regime of disease-induced extinction. Sustained oscillations do not exist.     

Population collapse to extinction: the catastrophic combination of parasitism and Allee effect

Infectious diseases are responsible for the extinction of a number of species. In conventional epidemic models, the transition from endemic population persistence to extirpation takes place gradually. However, if host demographics exhibits a strong Allee effect (AE) (population decline at low densities), extinction can occur abruptly in a catastrophic population crash. This might explain why species suddenly disappear even when they used to persist at high endemic population levels. Mathematically, the tipping point towards population collapse is associated with a saddle-node bifurcation. The underlying mechanism is the simultaneous population size depression and the increase of the extinction threshold due to parasite pathogenicity and Allee effect. Since highly pathogenic parasites cause their own extinction but not that of their host, there can be another saddle-node bifurcation with the re-emergence of two endemic equilibria. The implications for control interventions are discussed, suggesting that effective management may be possible for ?(0)?1.     

Combined impacts of Allee effects and parasitism

Despite their individual importance for population dynamics and conservation biology, the combined impacts of Allee effects and parasitism have received little attention. We built a mathematical model to compare the dynamics of populations with or without Allee effects when infected by microparasites. We show that the influence of an Allee effect takes the form of a tradeoff. The presence of an Allee effect in host populations may protect them, by reducing the range of population sizes that allow parasite spread. Yet if infection spreads, the Allee effect weakens host populations by reducing their size and by widening the range of parasite species that lead them to extinction. These results have important implications for predicting the survival of threatened populations or the success of reintroductions, and may help define size ranges within which given populations should be maintained to prevent both epidemics and Allee effects driven extinctions.     

Lost in time, lonely, and single: reproductive asynchrony and the Allee effect

Identifying linkages between life-history traits and small population processes is essential to effective multispecies conservation. Reproductive asynchrony, which occurs when individuals are reproductively active for only a portion of the population-level breeding period, may provide one such link. Traditionally, reproductive asynchrony has been considered from evolutionary perspectives as an advantageous bet-hedging strategy in temporally unpredictable environments. Here, we explore the dynamic consequences of reproductive asynchrony as a density-dependent life-history trait. To examine how asynchrony affects population growth rate and extinction risk, we used a general model of reproductive timing to quantify the temporal overlap of opposite-sex individuals and to simulate population dynamics over a range of initial densities and empirical estimates of reproductive asynchrony. We also considered how protandry, a sexually selected life-history strategy that often accompanies asynchrony, modulates the population-level effects of reproductive asynchrony. We found that asynchrony decreases the number of males a female overlaps with, decreases the average probability of mating per male/female pair that does overlap, and leaves some females completely isolated in time. This loss of reproductive potential, which is exacerbated by protandry, reduces population growth rate at low density and can lead to extinction via an Allee effect. Thus reproductive asynchrony and protandry, both of which can be evolutionarily advantageous at higher population densities, may prove detrimental when population density declines.     

Backward bifurcations and strong Allee effects in matrix models for the dynamics of structured populations

In nonlinear matrix models, strong Allee effects typically arise when the fundamental bifurcation of positive equilibria from the extinction equilibrium at r=1 (or R₀=1) is backward. This occurs when positive feedback (component Allee) effects are dominant at low densities and negative feedback effects are dominant at high densities. This scenario allows population survival when r (or equivalently R₀) is less than 1, provided population densities are sufficiently high. For r>1 (or equivalently R₀>1) the extinction equilibrium is unstable and a strong Allee effect cannot occur. We give criteria sufficient for a strong Allee effect to occur in a general nonlinear matrix model. A juvenile–adult example model illustrates the criteria as well as some other possible phenomena concerning strong Allee effects (such as positive cycles instead of equilibria).     

Allee effects,extinctions, and chaotic transients in simple population models

Discrete time single species models with overcompensating density dependence and an Allee effect due to predator satiation and mating limitation are investigated. The models exhibit four behaviors: persistence for all initial population densities, bistability in which a population persists for intermediate initial densities and otherwise goes extinct, extinction for all initial densities, and essential extinction in which "almost every" initial density leads to extinction. For fast-growing populations, these models show populations can persist at high levels of predation even though lower levels of predation lead to essential extinction. Alternatively, increasing the predator's handling time, the population's carrying capacity, or the likelihood of mating success may lead to essential extinction. In each of these cases, the mechanism behind these disappearances are chaotic dynamics driving populations below a critical threshold determined by the Allee effect. These disappearances are proceeded by chaotic transients that are proven to be approximately exponentially distributed in length and highly sensitive to initial population densities.     

Habitat selection reduces extinction of populations subject to Allee effects

Theoretical studies indicate that a single population under an Allee effect will decline to extinction if reduced below a particular threshold, but the existence of multiple local populations connected by random dispersal improves persistence of the global population. An additional process that can facilitate persistence is the existence of habitat selection by dispersers. Using analytic and simulation models of population change, I found that when habitat patches exhibiting Allee effects are connected by dispersing individuals, habitat selection by these dispersers increases the likelihood that patches persist at high densities, relative to results expected by random settlement. Populations exhibiting habitat selection also attain equilibrium more quickly than randomly dispersing populations. These effects are particularly important when Allee effects are large and more than two patches exist. Integrating habitat selection into population dynamics may help address why some studies have failed to find extinction thresholds in populations, despite well-known Allee effects in many species.     

捕食者具有厌食性反应且食饵具有Allee效应的捕食系统

在Dubis动力系统的基础上,建立了具有Allee效应的捕食系统模型。对系统的稳定性进行了分析,受Allee效应的影响,食饵种群可能因为种群大小处于临界点以下而趋于灭绝。通过对系统进行模拟,结果表明:不受Allee效应的影响,系统的演化属于一种理想化的情形系统到达P(平衡)点的时间较不受Allee效应影响时系统到达P点的时间短,不利于生物的进化,而在Allee效应的影响下,系统的演化将达到一个平衡状态。由此,说明Allee效应为濒临灭绝物种的管理提供了重要的理论依据,对管理部门的决策有参考指导作用。     

具有Allee效应的合作进化

Allee效应对物种的续存是潜在的干扰因素,在很大程度上将增加种群局部甚至全局灭绝的可能性。对许多物种,尤其是濒临物种更容易受其影响。将Allee效应引入囚徒困境博弈模型,通过理论分析与数值模拟相结合的方法分析讨论了Allee效应对合作进化的影响。研究结果表明:在恶劣的环境条件下,Allee效应极易使物种灭绝,不利于合作进化;在相对优越的环境条件下(死亡率较低),Allee效应促进合作进化,且Allee效应强度越强,更有利于合作进化,不过种群的空间斑块占有率也会随着Allee效应强度的增强而降低,使物种最终灭绝。     

局域种群的Allee效应和集合种群的同步性

从包含Allee效应的局域种群出发,建立了耦合映像格子模型,即集合种群模型.通过分析和计算机模拟表明:(1)当局域种群受到Allee效应强度较大时,集合种群同步灭绝;(2)而当Allee效应强度相对较弱时,通过稳定局域种群动态(减少混沌)使得集合种群发生同步波动,而这种同步波动能够增加集合种群的灭绝风险;(3)斑块间的连接程度对集合种群同步波动的发生有很大的影响,适当的破碎化有利于集合种群的续存.全局迁移和Allee效应结合起来增加了集合种群同步波动的可能,从而增加集合种群的灭绝风险.这些结果对理解同步性的机理、利用同步机理来制定物种保护策略和害虫防治都有重要的意义.     

Allee effects can both conserve and create spatial heterogeneity in population densities

In order to determine conditions which allow the Allee effect (caused by biparental reproduction) to conserve and create spatial heterogeneity in population densities, we studied a deterministic model of a symmetric two-patch metapopulation. We proved that under certain conditions there exist stable equilibria with unequal population densities in the two patches, a situation which can be interpreted as conserved heterogeneity. Furthermore, the Allee effect can lead to instability of the equilibrium with equal population densities if some degree of competition is assumed to occur between the subpopulations (non-local competition). This indicates the potential of the Allee effect to create spatial heterogeneity. Neither of these effects appear under biologically realistic parameter values in a model where uniparental reproduction is assumed. We proved that both the between-patch migration intensity and the degree of non-local competition are decisive in determining boundaries between these types of behaviour of the spatial system with Allee effect. Therefore, we propose that the Allee effect, migration intensity, and non-local competition should be considered jointly in studies focusing on problems like pattern formation in space and invasions of spreading species.     

Impact of Allee effect on an eco-epidemiological system

In this paper, we propose a general ratio-dependent prey-predator model with disease in predator subject to the strong Allee effect in prey. We obtain the complete dynamics of both models: (a) full model with Allee effect; (b) full model without Allee effect. Model (a) may have more than one interior equilibrium point, but model (b) has only one interior equilibrium point. Numerical results reveal that the coexistence of all the populations at the endemic state is possible for both the models. But for the model with Allee effect, the coexistence can be destroyed by an increased supply of alternative food for the predators. It can also be proved that for the full model with Allee effect, the disease can be suppressed under certain parametric conditions. Also by comparing models (a) and (b), we conclude that Allee effect can create or destroy the interior attractor. Finally, we have studied the disease free-submodel (prey and susceptible predator model) with and without Allee effect. The comparative study between these two submodels leads to the following conclusions: 1) In the presence of Allee effect, the number of interior equilibrium points can change from zero to two whereas the submodel without Allee effect has unique interior equilibrium point; 2) Both with and without Allee effect, initial conditions play an important role on the survival and extinction of prey as well as its corresponding predator; 3) In the presence of Allee effect, bi-stability occurs with stable or periodic coexistence of prey and susceptible predator and the extinction of prey and susceptible predator; 4) Allee effect can generate or destroy the interior equilibrium points.     

Demographic and Component Allee Effects in Southern Lake Superior Gray Wolves

Recovering populations of carnivores suffering Allee effects risk extinction because positive population growth requires a minimum number of cooperating individuals. Conservationists seldom consider these issues in planning for carnivore recovery because of data limitations, but ignoring Allee effects could lead to overly optimistic predictions for growth and underestimates of extinction risk. We used Bayesian splines to document a demographic Allee effect in the time series of gray wolf (Canis lupus) population counts (1980–2011) in the southern Lake Superior region (SLS, Wisconsin and the upper peninsula of Michigan, USA) in each of four measures of population growth. We estimated that the population crossed the Allee threshold at roughly 20 wolves in four to five packs. Maximum per-capita population growth occurred in the mid-1990s when there were approximately 135 wolves in the SLS population. To infer mechanisms behind the demographic Allee effect, we evaluated a potential component Allee effect using an individual-based spatially explicit model for gray wolves in the SLS region. Our simulations varied the perception neighborhoods for mate-finding and the mean dispersal distances of wolves. Simulation of wolves with long-distance dispersals and reduced perception neighborhoods were most likely to go extinct or experience Allee effects. These phenomena likely restricted population growth in early years of SLS wolf population recovery.     

Modelling the mating system of polar bears: a mechanistic approach to the Allee effect

Allee effects may render exploited animal populations extinction prone, but empirical data are often lacking to describe the circumstances leading to an Allee effect. Arbitrary assumptions regarding Allee effects could lead to erroneous management decisions so that predictive modelling approaches are needed that identify the circumstances leading to an Allee effect before such a scenario occurs. We present a predictive approach of Allee effects for polar bears where low population densities, an unpredictable habitat and harvest-depleted male populations result in infrequent mating encounters. We develop a mechanistic model for the polar bear mating system that predicts the proportion of fertilized females at the end of the mating season given population density and operational sex ratio. The model is parametrized using pairing data from Lancaster Sound, Canada, and describes the observed pairing dynamics well. Female mating success is shown to be a nonlinear function of the operational sex ratio, so that a sudden and rapid reproductive collapse could occur if males are severely depleted. The operational sex ratio where an Allee effect is expected is dependent on population density. We focus on the prediction of Allee effects in polar bears but our approach is also applicable to other species.     

Habitat destruction and the extinction debt revisited: The Allee effect

Habitat destruction, often caused by anthropogenic disturbance, can lead to the extinction of species at an unprecedented rate. It is important, therefore, to consider habitat destruction when assessing population viability. Another factor often ignored in population viability analysis, is the Allee effect that adds to the risk of populations already on the verge of extinction. Understanding the Allee effect on species dynamics and response to habitat destruction has intrinsic value in conservation prioritization. Here, the Allee effect was considered in a multi-species hierarchical competition model. Results showed that species persistence declines dramatically due to the Allee effect, and certain species become more susceptible to habitat destruction than others. Two extinction orders emerged under habitat destruction: either the best competitor becomes extinct first or the best colonizer first. The extinction debt and order, as well as the time lag between habitat destruction and species extinction, were found to be determined by species abundance and the intensity of the Allee effect.     

Allee effects in stochastic populations

The Allee effect, or inverse density dependence at low population sizes, could seriously impact preservation and management of biological populations. The mounting evidence for widespread Allee effects has lately inspired theoretical studies of how Allee effects alter population dynamics. However, the recent mathematical models of Allee effects have been missing another important force prevalent at low population sizes: stochasticity. In this paper, the combination of Allee effects and stochasticity is studied using diffusion processes, a type of general stochastic population model that accommodates both demographic and environmental stochastic fluctuations. Including an Allee effect in a conventional deterministic population model typically produces an unstable equilibrium at a low population size, a critical population level below which extinction is certain. In a stochastic version of such a model, the probability of reaching a lower size a before reaching an upper size b , when considered as a function of initial population size, has an inflection point at the underlying deterministic unstable equilibrium. The inflection point represents a threshold in the probabilistic prospects for the population and is independent of the type of stochastic fluctuations in the model. In particular, models containing demographic noise alone (absent Allee effects) do not display this threshold behavior, even though demographic noise is considered an "extinction vortex". The results in this paper provide a new understanding of the interplay of stochastic and deterministic forces in ecological populations.     

On some theory of monostable and bistable pure birth-jump integro-differential equations

We study an integral-differential equation that models a pure birth-jump process, where birth and dispersal cannot be decoupled. A case has been made that these processes are more suitable for phenomena such as plant dynamics, fire propagation, and cancer cell dynamics. We contrast the dynamics of this equation with those of the classical reaction-diffusion equation, where the reaction term models either logistic growth or a strong Allee effect. Recent evidence of an Allee effect has been found in plant dynamics during the germination process (due to seed predation) but not in the generation of seeds. This motivates where the Allee effect is included in our model. We prove the global existence and uniqueness of solutions with bounded initial data and analyze some properties of the solutions. Additionally, we prove results related to the persistence or extinction of a species, which are analogous to those of the classical reaction-diffusion equation. A key finding is that in some cases a population which is initially below the Allee threshold in some area, even if small, will actually survive. This is in contrast to solutions of the classical reaction-diffusion with the same initial data. Another difference of note is the lack of regularization and an infinite number of discontinuous equilibrium solutions to the birth-jump model.     

Expansion or extinction: deterministic and stochastic two-patch models with Allee effects

We investigate the impact of Allee effect and dispersal on the long-term evolution of a population in a patchy environment. Our main focus is on whether a population already established in one patch either successfully invades an adjacent empty patch or undergoes a global extinction. Our study is based on the combination of analytical and numerical results for both a deterministic two-patch model and a stochastic counterpart. The deterministic model has either two, three or four attractors. The existence of a regime with exactly three attractors only appears when patches have distinct Allee thresholds. In the presence of weak dispersal, the analysis of the deterministic model shows that a high-density and a low-density populations can coexist at equilibrium in nearby patches, whereas the analysis of the stochastic model indicates that this equilibrium is metastable, thus leading after a large random time to either a global expansion or a global extinction. Up to some critical dispersal, increasing the intensity of the interactions leads to an increase of both the basin of attraction of the global extinction and the basin of attraction of the global expansion. Above this threshold, for both the deterministic and the stochastic models, the patches tend to synchronize as the intensity of the dispersal increases. This results in either a global expansion or a global extinction. For the deterministic model, there are only two attractors, while the stochastic model no longer exhibits a metastable behavior. In the presence of strong dispersal, the limiting behavior is entirely determined by the value of the Allee thresholds as the global population size in the deterministic and the stochastic models evolves as dictated by their single-patch counterparts. For all values of the dispersal parameter, Allee effects promote global extinction in terms of an expansion of the basin of attraction of the extinction equilibrium for the deterministic model and an increase of the probability of extinction for the stochastic model.     

Linking the allee effect, sexual reproduction, and temperature-dependent sex determination via spatial dynamics

We develop a spatially explicit, two-sex, individual-based model (IBM) and a derived spatially homogeneous model (SHM) to describe the Allee effect due to scarcity of mating possibilities at low population sizes or densities. The SHM, based on coupled difference equations, represents the first spatially homogeneous approach to this phenomenon, which differentiates between sexes and relies only on measurable population parameters. The IBM reinforces the findings of the SHM by adopting more realistic mate search strategies of diffusive movement and active search. Both models are characterized by a hyperbolic-shaped extinction boundary in the male-female state space, which contrasts with a linear boundary in one-dimensional models of the Allee effect. We examine how the position of the extinction boundary depends on population demography (primary sex ratio, reproduction and mortality probabilities) and adopted mate search strategies. The investigation of different phases in the IBM dynamics emphasizes the differences between local and global densities and shows the importance of scale when assessing the Allee effect. To demonstrate the potential application of our models, we combine the SHM and available data to predict the impact of environmental temperature changes on two turtle species with temperature-dependent sex determination.