We present an explicit unified stochastic model of fluctuations in population size due to random birth, death, density-dependent competition and environmental fluctuations. Stochastic dynamics provide insight into small populations, including processes such as extinction, that cannot be correctly treated by deterministic methods. We present exact analytical and simulation-based results for extinction times of our stochastic model and compare the different effects of environmental stochasticity and intrinsic demographic stochasticity. We use both the discrete master equation approach and an exact mapping to a Fokker-Planck equation (the Poisson method) and stochastic equation, showing they are precisely equivalent. We also calculate approximate extinction times using a steepest descent method. This model can readily be extended to accommodate metapopulation structure and genetic variation in the population and thus represents a step towards a microscopically explicit synthesis of population dynamics and population genetics.
Demographic stochasticity is important in determining extinction risks of small populations, but it is largely unknown how its effect depends on the life histories of species. We modeled effects of demographic stochasticity on extinction risk in a broad range of generalized life histories, using matrix models and branching processes. Extinction risks of life histories varied greatly in their sensitivity to demographic stochasticity. Comparing life histories, extinction risk generally increased with increasing fecundity and decreased with higher ages of maturation. Effects of adult survival dep…
In large but finite populations, weak demographic stochasticity due to random birth and death events can lead to population extinction. The process is analogous to the escaping problem of trapped particles under random forces. Methods widely used in studying such physical systems, for instance, Wentzel-Kramers-Brillouin (WKB) and Fokker-Planck methods, can be applied to solve similar biological problems. In this article, we comparatively analyse applications of WKB and Fokker-Planck methods to some typical stochastic population dynamical models, including the logistic growth, endemic SIR, pred…
Methods for predicting the probability and timing of a species' extinction are typically based on a combination of theoretical models and empirical data, and focus on single species population dynamics. Of course, species also interact with each other, forming more or less complex networks of interactions. Models to assess extinction risk often lack explicit incorporation of these interspecific interactions. We study a birth and death process in which the death rate includes an effect from predation. This predation rate is included via a general nonlinear expression for the functional response…
We consider non-demographic noise in the form of uncertainty in the reaction step size, and reveal a dramatic effect this noise may have on the stability of self-regulating populations. Employing the reaction scheme mA->kA, but allowing, e.g., the product number k to be a-priori unknown and sampled from a given distribution, we show that such non-demographic noise can greatly reduce the population's extinction risk compared to the fixed k case. Our analysis is tested against numerical simulations, and by using empirical data of different species, we argue that certain distributions may be more…
We study simple stochastic scenarios, based on birth-and-death Markovian processes, that describe populations with Allee effect, to account for the role of demographic stochasticity. In the mean-field deterministic limit we recover well-known deterministic evolution equations widely employed in population ecology. The mean-time to extinction is in general obtained by the Wentzel-Kramers-Brillouin (WKB) approximation for populations with strong and weak Allee effects. An exact solution for the mean time to extinction can be found via a recursive equation for special cases of the stochastic dyna…
May and Leonard (SIAM J. Appl. Math 1975) introduced a three-species Lotka-Volterra type population model that exhibits heteroclinic cycling. Rather than producing a periodic limit cycle, the trajectory takes longer and longer to complete each "cycle", passing closer and closer to unstable fixed points in which one population dominates and the others approach zero. Aperiodic heteroclinic dynamics have subsequently been studied in ecological systems (side-blotched lizards; colicinogenic E. coli), in the immune system, in neural information processing models ("winnerless competition"), and in mo…
In recent years non-demographic variability has been shown to greatly affect dynamics of stochastic populations. For example, non-demographic noise in the form of a bursty reproduction process with an a-priori unknown burst size, or environmental variability in the form of time-varying reaction rates, have been separately found to dramatically impact the extinction risk of isolated populations. In this work we investigate the extinction risk of an isolated population under the combined influence of these two types of non-demographic variation. Using the so-called momentum-space WKB approach we…
Theoretical ecologists have long sought to understand how the persistence of populations depends on biotic and abiotic factors. Classical work showed that demographic stochasticity causes the mean time to extinction to increase exponentially with population size, whereas variation in environmental conditions can lead to a power law scaling. Recent work has focused especially on the influence of the autocorrelation structure ("color") of environmental noise. In theoretical physics, there is a burst of research activity in analyzing large fluctuations in stochastic population dynamics. This rese…
Understanding the mechanisms governing population extinctions is of key importance to many problems in ecology and evolution. Stochastic factors are known to play a central role in extinction, but the interactions between a population's demographic stochasticity and environmental noise remain poorly understood. Here, we model environmental forcing as a stochastic fluctuation between two states, one with a higher death rate than the other. We find that in general there exists a rate of fluctuations that minimizes the mean time to extinction, a phenomenon previously dubbed "resonant activation."…
Population genetics struggles to model extinction; standard models track the relative rather than absolute fitness of genotypes, while the exceptions describe only the short-term transition from imminent doom to evolutionary rescue. But extinction can result from failure to adapt not only to catastrophes, but also to a backlog of environmental challenges. We model long-term evolution to long series of small challenges, where fitter populations reach higher population sizes. The population's long-term fitness dynamic is well approximated by a simple stochastic Markov chain model. Long-term pers…
Dispersal is an important strategy that allows organisms to locate and exploit favorable habitats. The question arises: given competition in a spatially heterogeneous landscape, what is the optimal rate of dispersal? Continuous population models predict that a species with a lower dispersal rate always drives a competing species to extinction in the presence of spatial variation of resources. However, the introduction of intrinsic demographic stochasticity can reverse this conclusion. We present a simple model in which competition between the exploitation of resources and stochastic fluctuatio…
We study the combined impact of a colored environmental noise and demographic noise on the extinction risk of a long-lived and well-mixed isolated stochastic population which exhibits the Allee effect. The environmental noise modulates the population birth and death rates. Assuming that the Allee effect is strong, and the environmental noise is positively correlated and Gaussian, we derive a Fokker-Planck equation for the joint probability distribution of the population sizes and environmental fluctuations. In WKB approximation this equation reduces to an effective two-dimensional Hamiltonian …
Environmental stochasticity is known to be a destabilizing factor, increasing abundance fluctuations and extinction rates of populations. However, the stability of a community may benefit from the differential response of species to environmental variations due to the storage effect. This paper provides a systematic and comprehensive discussion of these two contradicting tendencies, using the metacommunity version of the recently proposed time-average neutral model of biodiversity which incorporates environmental stochasticity and demographic noise and allows for extinction and speciation. We …
Community ecology has traditionally relied on the competitive exclusion principle, a piece of common wisdom in conceptual frameworks developed to describe species assemblages. Key concepts in community ecology, such as limiting similarity and niche partitioning, are based on competitive exclusion. However, this classical paradigm in ecology relies on implications derived from simple, deterministic models. Here we show how the predictions of a symmetric, deterministic model about the way extinctions proceed can be utterly different from the results derived from the same model when ecological dr…
We consider a stochastic population model where the intrinsic or demographic noise causes cycling between states before the population eventually goes extinct. A master equation approach coupled with a WKB (Wentzel-Kramers-Brillouin) approximation is used to construct the optimal path to extinction. In addition, a probabilistic argument is used to understand the pre-extinction dynamics and approximate the mean time to extinction. Analytical results agree well with numerical Monte Carlo simulations. A control method is implemented to decrease the mean time to extinction. Analytical results quan…
Understanding the coexistence of diverse species in a changing environment is an important problem in community ecology. Bet-hedging is a strategy that helps species survive in such changing environments. However, studies of bet-hedging have often focused on the expected long-term growth rate of the species by itself, neglecting competition with other coexisting species. Here we study the extinction risk of a bet-hedging species in competition with others. We show that there are three contributions to the extinction risk. The first is the usual demographic fluctuation due to stochastic reprodu…
Recent collapses of many fisheries across the globe have challenged the mathematical approach to these systems through classic bioeconomic models. Decimated populations did not recover as fast as predicted by these models and depensatory effects were introduced to better fit the dynamics at low population abundances. Alternative to depensation, modeling captures by non-linear harvesting functions produces equivalent outcomes at small abundances, and the dynamics undergoes a bifurcation leading to population collapse and recovery once catching efforts are above or below certain thresholds, resp…
The extinction of a single species due to demographic stochasticity is analyzed. The discrete nature of the individual agents and the Poissonian noise related to the birth-death processes result in local extinction of a metastable population, as the system hits the absorbing state. The Fokker-Planck formulation of that problem fails to capture the statistics of large deviations from the metastable state, while approximations appropriate close to the absorbing state become, in general, invalid as the population becomes large. To connect these two regimes, a master equation based on a real space…
Neutral dynamics, where taxa are assumed to be demographically equivalent and their abundance is governed solely by the stochasticity of the underlying birth-death process, has proved itself as an important minimal model that accounts for many empirical datasets in genetics and ecology. However, the restriction of the model to demographic [${\cal{O}} ({\sqrt N})$] noise yields relatively slow dynamics that appears to be in conflict with both short-term and long-term characteristics of the observed systems. Here we analyze two of these problems - age size relationships and species extinction ti…
The scarcity of water characterising drylands forces vegetation to adopt appropriate survival strategies. Some of these generate water-vegetation feedback mechanisms that can lead to spatial self-organisation of vegetation, as it has been shown with models representing plants by a density of biomass, varying continuously in time and space. However, although plants are usually quite plastic they also display discrete qualities and stochastic behaviour. These features may give rise to demographic noise, which in certain cases can influence the qualitative dynamics of ecosystem models. In the pre…
Immigration can rescue local populations from extinction, helping to stabilise a metapopulation. Local population dynamics is important for determining the strength of this rescue effect, but the mechanistic link between local demographic parameters and the rescue effect at the metapopulation level has received very little attention by modellers. We develop an analytical framework that allows us to describe the emergence of the rescue effect from interacting local stochastic dynamics. We show this framework to be applicable to a wide range of spatial scales, providing a powerful and convenient…
Extinction is the ultimate absorbing state of any stochastic birth-death process, hence the time to extinction is an important characteristic of any natural population. Here we consider logistic and logistic-like systems under the combined effect of demographic and bounded environmental stochasticity. Three phases are identified: an inactive phase where the mean time to extinction $T$ increases logarithmically with the initial population size, an active phase where $T$ grows exponentially with the carrying capacity $N$, and temporal Griffiths phase, with power-law relationship between $T$ and …
Deterministic evolutionary theory robustly predicts that populations displaying altruistic behaviours will be driven to extinction by mutant cheats that absorb common benefits but do not themselves contribute. Here we show that when demographic stochasticity is accounted for, selection can in fact act in the reverse direction to that predicted deterministically, instead favouring cooperative behaviors that appreciably increase the carrying capacity of the population. Populations that exist in larger numbers experience a selective advantage by being more stochastically robust to invasions than …
Recreational fishing is a highly socio-ecological process. Although recreational fisheries are self-regulating and resilient, changing anthropogenic pressure drives these fisheries to overharvest and collapse. Here, we evaluate the effect of demographic and environmental stochasticity for a social-ecological two-species fish model. In the presence of noise, we find that an increase in harvesting rate drives a critical transition from high-yield/low-price fisheries to low-yield/high-price fisheries. To calculate stochastic trajectories for demographic noise, we derive the master equation corres…
Frequency dependent selection and demographic fluctuations play important roles in evolutionary and ecological processes. Under frequency dependent selection, the average fitness of the population may increase or decrease based on interactions between individuals within the population. This should be reflected in fluctuations of the population size even in constant environments. Here, we propose a stochastic model, which naturally combines these two evolutionary ingredients by assuming frequency dependent competition between different types in an individual-based model. In contrast to previous…
We investigate extinction of a long-lived self-regulating stochastic population, caused by intrinsic (demographic) noise. Extinction typically occurs via one of two scenarios depending on whether the absorbing state n=0 is a repelling (scenario A) or attracting (scenario B) point of the deterministic rate equation. In scenario A the metastable stochastic population resides in the vicinity of an attracting fixed point next to the repelling point n=0. In scenario B there is an intermediate repelling point n=n_1 between the attracting point n=0 and another attracting point n=n_2 in the vicinity o…
We analyse metapopulation dynamics in terms of an individual-based, stochastic model of a finite metapopulation. We suggest a new approach, using the number of patches in the population as a large parameter. This approach does not require that the number of individuals per patch is large, neither is it necessary to assume a time-scale separation between local population dynamics and migration. Our approach makes it possible to accurately describe the dynamics of metapopulations consisting of many small patches. We focus on metapopulations on the brink of extinction. We estimate the time to ext…
Chloroplast microsatellites are becoming increasingly popular markers for population genetic studies in plants, but there has been little focus on their potential for demographic inference. In this work the utility of chloroplast microsatellites for the study of population expansions was explored. First, we investigated the power of mismatch distribution analysis and the F(S) test with coalescent simulations of different demographic scenarios. We then applied these methods to empirical data obtained for the Canary Island pine (Pinus canariensis). The results of the simulations showed that chlo…
Understanding the conditions ensuring the persistence of a population is an issue of primary importance in population biology. The first theoretical approach to the problem dates back to the 50's with the KiSS (after Kierstead, Slobodkin and Skellam) model, namely a continuous reaction-diffusion equation for a population growing on a patch of finite size $L$ surrounded by a deadly environment with infinite mortality -- i.e. an oasis in a desert. The main outcome of the model is that only patches above a critical size allow for population persistence. Here, we introduce an individual-based anal…
Multiple-scale mobility is ubiquitous in nature and has become instrumental for understanding and modeling animal foraging behavior. However, the impact of individual movements on the long-term stability of populations remains largely unexplored. We analyze deterministic and stochastic Lotka Volterra systems, where mobile predators consume scarce resources (prey) confined in patches. In fragile systems (that is, those unfavorable to species coexistence), the predator species has a maximized abundance and is resilient to degraded prey conditions when individual mobility is multiple scaled. With…
Stochasticity can play an important role in the dynamics of biologically relevant populations. These span a broad range of scales: from intra-cellular populations of molecules to population of cells and then to groups of plants, animals and people. Large deviations in stochastic population dynamics - such as those determining population extinction, fixation or switching between different states - are presently in a focus of attention of statistical physicists. We review recent progress in applying different variants of dissipative WKB approximation (after Wentzel, Kramers and Brillouin) to thi…
An ecological system with multiple stable equilibria is prone to undergo catastrophic change or regime shift from one steady-state to another. It should be noted that, if one of the steady states is an extinction state, the catastrophic change may lead to extinction. A suitable manual measure may control the prevention of catastrophic changes of different species from one equilibrium to another. We consider two stochastic models with linear and nonlinear harvesting terms. We inspect either density regulation or Allee type density regulated models [Saha et al., Ecological Modelling, 2013], whic…
We present a general framework to describe the evolutionary dynamics of an arbitrary number of types in finite populations based on stochastic differential equations (SDE). For large, but finite populations this allows to include demographic noise without requiring explicit simulations. Instead, the population size only rescales the amplitude of the noise. Moreover, this framework admits the inclusion of mutations between different types, provided that mutation rates, $μ$, are not too small compared to the inverse population size 1/N. This ensures that all types are almost always represented i…
The abundance of a species' population in an ecosystem is rarely stationary, often exhibiting large fluctuations over time. Using historical data on marine species, we show that the year-to-year fluctuations of population growth rate obey a well-defined double-exponential (Laplace) distribution. This striking regularity allows us to devise a stochastic model despite seemingly irregular variations in population abundances. The model identifies the effect of reduced growth at low population density as a key factor missed in current approaches of population variability analysis and without which …
A strong demographic Allee effect in which the expected population growth rate is negative below a certain critical population size can cause high extinction probabilities in small introduced populations. However, many species are repeatedly introduced to the same location and eventually one population may overcome the Allee effect by chance. With the help of stochastic models, we investigate how much genetic diversity such successful populations harbour on average and how this depends on offspring-number variation, an important source of stochastic variability in population size. We find that…
Population dynamics reflects an underlying birth-death process, where the rates associated with different events may depend on external environmental conditions and on the population density. A whole family of simple and popular deterministic models (like logistic growth) support a transcritical bifurcation point between an extinction phase and an active phase. Here we provide a comprehensive analysis of the phases of that system, taking into account both the endogenous demographic noise (random birth and death events) and the effect of environmental stochasticity that causes variations in bir…
In this study, first we expand the cubic population model under Allee effect with a quadratic harvest function that represents harvest effect. Then, using four reaction equations representing the micro-interactions within the population under the influence of demographic noise and harvest, we obtain the mean field equations containing the effect of the harvest from the solution of the master equation. In this way, a relationship is established between the micro and macro parameters of the population. As a result, we calculate Mean Time to Extinction (MTE) by using WKB approximation for single …
Classical stochastic demography predicts that environmental stochasticity reduces population growth rates and, thereby, can increase extinction risk. In contrast, the SAS-CFF model demonstrates that environmental stochasticity can promote genetic diversity. Extending the SAS-CFF to account for demography, I examine the simultaneous effects of environmental stochasticity on genetic diversity and population persistence. Consistent with Gillespie's analysis, if the log-fitness function is concave and allelic responses to the environment are not perfectly correlated, then per-capita growth rates o…
The dynamics of two competing species in a finite size community is one of the most studied problems in population genetics and community ecology. Stochastic fluctuations lead, inevitably, to the extinction of one of the species, but the relevant timescale depends on the underlying dynamics. The persistence time of the community has been calculated for neutral models, where the only drive of the system is drift (demographic stochasticity) and for models with strong selection. Following recent analyses that stress the importance of environmental stochasticity in empirical systems, we present he…
The "Kill the Winner" hypothesis is an attempt to address the problem of diversity in biology. It argues that host-specific predators control the population of each prey, preventing a winner from emerging and thus maintaining the coexistence of all species in the system. We develop a stochastic model for the "Kill the Winner" paradigm and show that the stable coexistence state of the deterministic "Kill the Winner" model is destroyed by demographic stochasticity, through a cascade of extinction events. We formulate an individual-level stochastic model in which predator-prey coevolution promote…
Over the past century, nonlinear difference and differential equations have been used to understand conditions for species coexistence. However, these models fail to account for random fluctuations due to demographic and environmental stochasticity which are experienced by all populations. I review some recent mathematical results about persistence and coexistence for models accounting for each of these forms of stochasticity. Demographic stochasticity stems from populations and communities consisting of a finite number of interacting individuals, and often are represented by Markovian models …
Discrete time, spatially extended models play an important role in ecology, modelling population dynamics of species ranging from micro-organisms to birds. An important question is how 'bottom up', individual-based models can be approximated by 'top down' models of dynamics. Here, we study a class of spatially explicit individual-based models with contest competition: where species compete for space in local cells and then disperse to nearby cells. We start by describing simulations of the model, which exhibit large-scale discrete oscillations and characterise these oscillations by measuring s…
Structure, composition and stability of ecological populations are shaped by the inter- and intra-species interactions within these communities. It remains to be fully understood how the interplay of these interactions with other factors, such as immigration, control the structure, diversity and the long term stability of ecological systems in the presence of noise and fluctuations. We address this problem using a minimal model of interacting multi-species ecological communities that incorporates competition, immigration and demographic noise. We find that the complete phase diagram exhibits r…
Competition is the main driver of population dynamics, which shapes the genetic composition of populations and the assembly of ecological communities. Neutral models assume that all the individuals are equivalent and that the dynamics is governed by demographic (shot) noise, with a steady state species abundance distribution (SAD) that reflects a mutation-extinction equilibrium. Recently, many empirical and theoretical studies emphasized the importance of environmental variations that affect coherently the relative fitness of entire populations. Here we consider two generic time-averaged neutr…
Positive density-dependence occurs when individuals experience increased survivorship, growth, or reproduction with increased population densities. Mechanisms leading to these positive relationships include mate limitation, saturating predation risk, and cooperative breeding and foraging. Individuals within these populations may differ in age, size, or geographic location and thereby structure these populations. Here, I study structured population models accounting for positive density-dependence and environmental stochasticity i.e. random fluctuations in the demographic rates of the populatio…
