Species extinction is a core process that affects the diversity of life on Earth. Competition between species in a population is considered by ecological niche-based theories as a key factor leading to different severity of species extinctions. There are population dynamics models that describe a simple and easily understandable mechanism for resource competition. However, these models can not efficiently characterize and quantify new emergent extinctions in a large population appearing due to environmental forcing. To address this issue we develop a stochastic physics-inspired approach to analyze how environmental forcing influences the severity of species extinctions in such models. This approach is based on the large deviations theory of stochastic processes (the Freidlin-Wentzell theory). We show that there are three possible fundamentally different scenarios of extinctions, which we call catastrophic extinctions, asymmetric ones, and extinctions with exponentially small probabilities. The realization of those scenarios depends on environmental noise properties and the boundaries of niches, which define the domain, where species survive. Furthermore, we describe a hysteresis effect in species extinction showing that fluctuations can lead to dramatic consequences even if an averaged resource supply is sufficient to support population survival. Our stochastic physics-inspired approach generalizes niche theory by accounting environmental forcing and will be useful to find, by available data, which environmental perturbations may induce extinctions.
Most infectious diseases including more than half of known human pathogens are not restricted to just one host, yet much of the mathematical modeling of infections has been limited to a single species. We investigate consequences of a single epidemic propagating in multiple species and compare and contrast it with the endemic steady state of the disease. We use the two-species Susceptible-Infected-Recovered (SIR) model to calculate the severity of post-epidemic collapses in populations of two host species as a function of their initial population sizes, the times individuals remain infectious,…
Current theories about mechanisms promoting species co-existence in diverse communities assume that species only interact ecologically. Species are treated as discrete evolutionary entities, even though abundant empirical evidence indicates that patterns of gene flow such as selfing and hybridization frequently occur in plant and animal groups. Here, we allow mating behavior to respond to local species composition and abundance in a data-driven meta-community model of species co-existence. While individuals primarily out-cross, they also maintain some small capacity for selfing and hybridizati…
Understanding how an extinction event affects ecosystem is fundamental to biodiversity conservation. For this reason, food web response to species loss has been investigated in several ways in the last years. Several studies focused on secondary extinction due to biodiversity loss in a bottom-up perspective using in-silico extinction experiments in which a single species is removed at each step and the number of secondary extinctions is recorded. In these binary simulations a species goes secondarily extinct if it loses all its resource species, that is, when the energy intake is zero. This pu…
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…
The extinction of species is a core process that affects the diversity of life on Earth. One way of investigating the causes and consequences of extinctions is to build conceptual ecological models, and to use the dynamical outcomes of such models to provide quantitative formalization of changes to Earth's biosphere. In this paper we propose and study a conceptual resource model that describes a simple and easily understandable mechanism for resource competition, generalizes the well-known Huisman and Weissing model, and takes into account species self-regulation, extinctions, and time depende…
We consider a cyclically competing species model on a ring with global mixing at finite rate, which corresponds to the well-known Lotka-Volterra equation in the limit of infinite mixing rate. Within a perturbation analysis of the model from the infinite mixing rate, we provide analytical evidence that extinction occurs deterministically at sufficiently large but finite values of the mixing rate for any species number $N\ge3$. Further, by focusing on the cases of rather small species numbers, we discuss numerical results concerning the trajectories toward such deterministic extinction, includin…
Recent methodological advances are enabling better examination of speciation and extinction processes and patterns. A major open question is the origin of large discrepancies in species number between groups of the same age. Existing frameworks to model this diversity either focus on changes between lineages, neglecting global effects such as mass extinctions, or focus on changes over time which would affect all lineages. Yet it seems probable that both lineages differences and mass extinctions affect the same groups. Here we used simulations to test the performance of two widely used methods,…
Species extinction occurs regularly and unavoidably in ecological systems. The time scales for extinction can broadly vary and inform on the ecosystem's stability. We study the spatio-temporal extinction dynamics of a paradigmatic population model where three species exhibit cyclic competition. The cyclic dynamics reflects the non-equilibrium nature of the species interactions. While previous work focusses on the coarsening process as a mechanism that drives the system to extinction, we found that unexpectedly the dynamics to extinction is much richer. We observed three different types of dyna…
Economic incentives to harvest a species usually diminish as its abundance declines, because harvest costs increase. This prevents harvesting to extinction. A known exception can occur if consumer demand causes a declining species' harvest price to rise faster than costs. This threat may affect rare and valuable species, such as large land mammals, sturgeons, and bluefin tunas. We analyze a similar but underappreciated threat, which arises when the geographic area (range) occupied by a species contracts as its abundance declines. Range contractions maintain the local densities of declining pop…
We study the role of the noise in the dynamics of two competing species. We consider generalized Lotka-Volterra equations in the presence of a multiplicative noise, which models the interaction between the species and the environment. The interaction parameter between the species is a random process which obeys a stochastic differential equation with a generalized bistable potential in the presence of a periodic driving term, which accounts for the environment temperature variation. We find noise-induced periodic oscillations of the species concentrations and stochastic resonance phenomenon. W…
We investigate the formation of stable ecological networks where many species share the same resource. We show that such stable ecosystem naturally occurs as a result of extinctions. We obtain an analytical relation for the number of coexisting species and find a relation describing how many species that may go extinct as a result of a sharp environmental change. We introduce a special parameter that is a combination of species traits and resource characteristics used in the model formulation. This parameter describes the pressure on system to converge, by extinctions. When that stress paramet…
We use a classical combinatorial inequality to establish a Markov inequality for multivariate binary Markov processes on trees. We then apply this result, alongside with the FKG inequality, to compare the expected loss of biodiversity under two models of species extinction. One of these models is the generalized version of an earlier model in which extinction is influenced by some trait that can be classified into two states and which evolves on a tree according to a Markov process. Since more than one trait can affect the rates of species extinction, it is reasonable to allow, in the generali…
When four species compete stochastically in a cyclic way, the formation of two teams of mutually neutral partners is observed. In this paper we study through numerical simulations the extinction processes that can take place in this system both in the well mixed case as well as on different types of lattices. The different routes to extinction are revealed by the probability distribution of the domination time, i.e. the time needed for one team to fully occupy the system. If swapping is allowed between neutral partners, then the probability distribution is dominated by very long-lived states w…
We propose a generic model of eco-systems, with a {\it hierarchical} food web structure. In our computer simulations we let the eco-system evolve continuously for so long that that we can monitor extinctions as well as speciations over geological time scales. {\it Speciation} leads not only to horizontal diversification of species at any given trophic level but also to vertical bio-diversity that accounts for the emergence of complex species from simpler forms of life. We find that five or six trophic levels appear as the eco-system evolves for sufficiently long time, starting initially from j…
We use interacting particle systems to investigate survival and extinction of a species with colonies located on each site of $\mathbb {Z}^d$. In each of the four models studied, an individual in a local population can reproduce, die or migrate to neighboring sites. We prove that an increase of the death rate when the local population density is small (the Allee effect) may be critical for survival, and that the migration of large flocks of individuals is a possible solution to avoid extinction when the Allee effect is strong. We use attractiveness and comparison with oriented percolation, eit…
Coextinction models are useful to understand community robustness to species loss and resilience to disturbances. We simulated pollinator extinctions in pollination networks by using a hybrid model that combined a recently developed stochastic coextinction model (SCM) for plant extinctions and a topological model (TCM) for animal extinctions. Our model accounted for variation in interaction strengths and included empirical estimates of plant dependence on pollinators to set seeds. The stochastic nature of such model allowed us determining plant survival to single (and multiple) extinction even…
If predictions for species extinctions hold, then the `tree of life' today may be quite different to that in (say) 100 years. We describe a technique to quantify how much each species is likely to contribute to future biodiversity, as measured by its expected contribution to phylogenetic diversity. Our approach considers all possible scenarios for the set of species that will be extant at some future time, and weights them according to their likelihood under an independent (but not identical) distribution on species extinctions. Although the number of extinction scenarios can typically be very…
Planning for the protection of species often involves difficult choices about which species to prioritize, given constrained resources. One way of prioritizing species is to consider their "evolutionary distinctiveness", i.e. their relative evolutionary isolation on a phylogenetic tree. Several evolutionary isolation metrics or phylogenetic diversity indices have been introduced in the literature, among them the so-called Fair Proportion index (also known as the "evolutionary distinctiveness" score). This index apportions the total diversity of a tree among all leaves, thereby providing a simp…
Our paper computationally explores the extinction dynamics of an animal species effected by a sudden spike in mortality due to an extreme event. In our study, the animal species has a 2-year life cycle and is endowed with a high survival probability under normal circumstances. Our proposed approach does not involve any restraining assumptions concerning environmental variables or predator-prey relationships. Rather it is based on the simple premise that if observed on an year-to-year basis, the population size will be noted to either have gone up or come down as compared to last year. The conc…
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…
Effects of changes in functional composition of soil communities on nutrient cycling are still not well understood. Models simulating community dynamics overcome the technical challenges of conducting species removal experiments in the field. However, to date, available soil food web models do not adequately represent the organic matter processing chain which is key for soil dynamics. Here, we present a new model of soil food web dynamics accounting for allometric scaling of metabolic rate, ontogeny of organic matter, and explicit representation of nitrogen and carbon flows. We use this model …
Cyclic dominance between species may yield spiral waves that are known to provide a mechanism enabling persistent species coexistence. This observation holds true even in presence of spatial heterogeneity in the form of quenched disorder. In this work we study the effects on spatio-temporal patterns and species coexistence of structured spatial heterogeneity in the form of habitats that locally provide one of the species with an advantage. Performing extensive numerical simulations of systems with three and six species we show that these structured habitats destabilize spiral waves. Analyzing …
In a complex community, species continuously adapt to each other. On rare occasions, the adaptation of a species can lead to the extinction of others, and even its own. "Adaptive dynamics" is the standard mathematical framework to describe evolutionary changes in community interactions, and in particular, predict adaptation driven extinction. Unfortunately, most authors implement the equations of adaptive dynamics through computer simulations, that require assuming a large number of questionable parameters and fitness functions. In this study we present analytical solutions to adaptive dynamic…
Transitions to absorbing states are of fundamental importance in non-equilibrium physics as well as ecology. In ecology, absorbing states correspond to the extinction of species. We here study the spatial population dynamics of three cyclically interacting species. The interaction scheme comprises both direct competition between species as in the cyclic Lotka-Volterra model, and separated selection and reproduction processes as in the May-Leonard model. We show that the dynamic processes leading to the transient maintenance of biodiversity are closely linked to attractors of the nonlinear dyna…
We study a model of a multi-species ecosystem described by Lotka-Volterra-like equations. Interactions among species form a network whose evolution is determined by the dynamics of the model. Numerical simulations show power-law distribution of intervals between extinctions, but only for ecosystems with sufficient variability of species and with networks of connectivity above certain threshold that is very close to the percolation threshold of the network. Effect of slow environmental changes on extinction dynamics, degree distribution of the network of interspecies interactions, and some emer…
Simulations of the coevolution of many interacting species are performed using the Webworld model. The model has a realistic set of predator-prey equations that describe the population dynamics of the species for any structure of the food web. The equations account for competition between species for the same resources, and for the diet choice of predators between alternative prey according to an evolutionarily stable strategy. The set of species present undergoes long-term evolution due to speciation and extinction events. We summarize results obtained on the macro-evolutionary dynamics of sp…
Consumer demand for plant and animal products threatens many populations with extinction. The anthropogenic Allee effect (AAE) proposes that such extinctions can be caused by prices for wildlife products increasing with species rarity. This price-rarity relationship creates financial incentives to extract the last remaining individuals of a population, despite higher search and harvest costs. The AAE has become a standard approach for conceptualizing the threat of economic markets on endangered species. Despite its potential importance for conservation, AAE theory is based on a simple graphica…
We analyze the long term behavior of interacting populations which can be controlled through harvesting. The dynamics is assumed to be discrete in time and stochastic due to the effect of environmental fluctuations. We present extinction and coexistence criteria when there are one or two interacting species. We then use these tools in order to see when harvesting leads to extinction or persistence of species, as well as what the optimal harvesting strategies, which maximize the expected long term yield, look like. For single species systems, we show under certain conditions that the optimal ha…
In this work we suggest and consider an original, simple mathematical model of a "quasi-rapid" extinction population dynamics. It describes a decrease and final extinction of the population of one prey species by a "quasi-rapid" interaction with one predator species with increasing population. This "quasi-rapid" interaction means ecologically that prey species behaves practically quite passively (since there is no time for any reaction, i.e. defense), like an appropriate environment, in respect to "quasi-rapid" activity of the predator species that can have different "quasi-rapid" hunting abil…
The current rate of species extinction is rapidly approaching unprecedented highs and life on Earth presently faces a sixth mass extinction event driven by anthropogenic activity, climate change and ecological collapse. The field of conservation genetics aims at preserving species by using their levels of genetic diversity, usually measured as neutral genome-wide diversity, as a barometer for evaluating population health and extinction risk. A fundamental assumption is that higher levels of genetic diversity lead to an increase in fitness and long-term survival of a species. Here, we argue aga…
Mathematical models of population extinction have a variety of applications in such areas as ecology, paleontology and conservation biology. Here we propose and investigate two types of sub-exponential models of population extinction.The results of performed analysis show that the principle of minimum of information loss is the underlying law for the evolution of a broad class of models of population extinction. Finally, we propose a possible application of this modeling framework to mechanisms underlying time perception.
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…
Food-web perturbations stemming from climate change, overexploitation, invasive species, and habitat degradation often cause an initial loss of species that results in a cascade of secondary extinctions, posing considerable challenges to ecosystem conservation efforts. Here we devise a systematic network-based approach to reduce the number of secondary extinctions using a predictive modeling framework. We show that the extinction of one species can often be compensated by the concurrent removal or population suppression of other specific species, which is a counterintuitive effect not previous…
Phylogenetic diversity is a measure for describing how much of an evolutionary tree is spanned by a subset of species. If one applies this to the (unknown) subset of current species that will still be present at some future time, then this `future phylogenetic diversity' provides a measure of the impact of various extinction scenarios in biodiversity conservation. In this paper we study the distribution of future phylogenetic diversity under a simple model of extinction (a generalized `field of bullets' model). We show that the distribution of future phylogenetic diversity converges to a norma…
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…
We study the stochastic evolution of four species in cyclic competition in a well mixed environment. In systems composed of a finite number $N$ of particles these simple interaction rules result in a rich variety of extinction scenarios, from single species domination to coexistence between non-interacting species. Using exact results and numerical simulations we discuss the temporal evolution of the system for different values of $N$, for different values of the reaction rates, as well as for different initial conditions. As expected, the stochastic evolution is found to closely follow the me…
Populations of competing biological species exhibit a fascinating interplay between the nonlinear dynamics of evolutionary selection forces and random fluctuations arising from the stochastic nature of the interactions. The processes leading to extinction of species, whose understanding is a key component in the study of evolution and biodiversity, are influenced by both of these factors. In this paper, we investigate a class of stochastic population dynamics models based on generalized Lotka-Volterra systems. In the case of neutral stability of the underlying deterministic model, the impact o…
We investigate stochastic extinction in an epidemic model and the impact of random vaccinations in large populations formulated in terms of an optimal escape path. We find that different random vaccination strategies can have widely different results in decreasing expected time till extinction, for the same total amount of vaccines used. Vaccination strategies are considered in terms of two parameters: average frequency of vaccinations, given by $γ$, and the amplitude of the vaccinations, $ε$, where $ε\ll 1$ refers to the proportion of the population being vaccinated at some particular instant…
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…
We study inhomogeneous host-pathogen dynamics to model the global amphibian population extinction in a lake basin system. The lake basin system is modeled as quenched disorder. In this model we show that once the pathogen arrives at the lake basin it spreads from one lake to another, eventually spreading to the entire lake basin system in a wave like pattern. The extinction time has been found to depend on the steady state host population and pathogen growth rate. Linear estimate of the extinction time is computed. The steady state host population shows a threshold behavior in the interaction …
We study the ABC model (A + B --> 2B, B + C --> 2C, C + A --> 2A), and its counterpart: the three--component neutral drift model (A + B --> 2A or 2B, B + C --> 2B or 2C, C + A --> 2C or 2A.) In the former case, the mean field approximation exhibits cyclic behaviour with an amplitude determined by the initial condition. When stochastic phenomena are taken into account the amplitude of oscillations will drift and eventually one and then two of the three species will become extinct. The second model remains stationary for all initial conditions in the mean field approximation, and drifts when sto…
Cretaceous-Cenozoic history of angiosperms led to the a certain character of the distribution of taxa of different levels (the number of species and genera in families, species/genera ratio in families, the number of species in the genera). In most cases, these distributions are satisfactorily described by a power law (Pareto distribution). In logarithmic coordinates power function is a straight line. Empirical curves repeat this line is good enough, but on the right side of the graph (at low volumes taxa), there is a marked deviation of theoretical from the empirical curves. This suggests tha…
Many populations in nature are fragmented: they consist of local populations occupying separate patches. A local population is prone to extinction due to the shot noise of birth and death processes. A migrating population from another patch can dramatically delay the extinction. What is the optimal migration rate that minimizes the extinction risk of the whole population? Here we answer this question for a connected network of model habitat patches with different carrying capacities.
Extinction of a long-lived isolated stochastic population can be described as an exponentially slow decay of quasi-stationary probability distribution of the population size. We address extinction of a population in a two-population system in the case when the population turnover -- renewal and removal -- is much slower than all other processes. In this case there is a time scale separation in the system which enables one to introduce a short-time quasi-stationary extinction rate W_1 and a long-time quasi-stationary extinction rate W_2, and develop a time-dependent theory of the transition bet…
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 exa…
We propose and study {a generalized Beverton-Holt competition model} subject to Allee effects to obtain insights on how the interplay of Allee effects and contest competition affects the persistence and the extinction of two competing species. By using {the theory of monotone dynamics} and the properties of critical curves for non-invertible maps, our analysis shows that our model has simple equilibrium dynamics for most conditions. The coexistence of two competing species occurs only if the system has four interior equilibria.} We provide an {approximation to} the basins of the boundary attra…
Established populations often exhibit oscillations in their sizes. If a population is isolated, intrinsic stochasticity of elemental processes can ultimately bring it to extinction. Here we study extinction of oscillating populations in a stochastic version of the Rosenzweig-MacArthur predator-prey model. To this end we extend a WKB approximation (after Wentzel, Kramers and Brillouin) of solving the master equation to the case of extinction from a limit cycle in the space of population sizes. We evaluate the extinction rates and find the most probable paths to extinction by applying Floquet th…
We study species abundance in the empirical plant-pollinator mutualistic networks exhibiting broad degree distributions, with uniform intra-group competition assumed, by the Lotka-Volterra equation. The stability of a fixed point is found to be identified by the signs of its non-zero components and those of its neighboring fixed points. Taking the annealed approximation, we derive the non-zero components to be formulated in terms of degrees and the rescaled interaction strengths, which lead us to find different stable fixed points depending on parameters, and we obtain the phase diagram. The s…
The current understanding of supernova and gamma-ray burst events suggests important effects on the biosphere if one of more of them happened to strike the earth in the past. In this paper we evaluate the possibility that life extinctions which probably occurred due to excess of radiation occurring in the geologic past might have left a genetic signature on surviving species. We emphasize the signatures of these extinctions, proposing a quantitative model to evaluate the surviving probability of the species, based on kinetic aspects of the frequency of mutations and the DNA repair rate.
We study a lattice model of a multi-species prey-predator system. Numerical results show that for a small mutation rate the model develops irregular long-period oscillatory behavior with sizeable changes in a number of species. The periodicity of extinctions on Earth was suggested by Raup and Sepkoski but so far is lacking a satisfactory explanation. Our model indicates that this is a natural consequence of the ecosystem dynamics, not the result of any extraterrestrial cause.
Cyclic dominance of species has been identified as a potential mechanism to maintain biodiversity, see e.g. B. Kerr, M. A. Riley, M. W. Feldman and B. J. M. Bohannan [Nature {\bf 418}, 171 (2002)] and B. Kirkup and M. A. Riley [Nature {\bf 428}, 412 (2004)]. Through analytical methods supported by numerical simulations, we address this issue by studying the properties of a paradigmatic non-spatial three-species stochastic system, namely the `rock-paper-scissors' or cyclic Lotka-Volterra model. While the deterministic approach (rate equations) predicts the coexistence of the species resulting i…
Isolated populations ultimately go extinct because of the intrinsic noise of elementary processes. In multi-population systems extinction of a population may occur via more than one route. We investigate this generic situation in a simple predator-prey (or infected-susceptible) model. The predator and prey populations may coexist for a long time but ultimately both go extinct. In the first extinction route the predators go extinct first, whereas the prey thrive for a long time and then also go extinct. In the second route the prey go extinct first causing a rapid extinction of the predators. A…
Within large taxonomic assemblages, the number of species with adult body mass M is characterized by a broad but asymmetric distribution, with the largest mass being orders of magnitude larger than the typical mass. This canonical shape can be explained by cladogenetic diffusion that is bounded below by a hard limit on viable species mass and above by extinction risks that increase weakly with mass. Here we introduce and analytically solve a simplified cladogenetic diffusion model. When appropriately parameterized, the diffusion-reaction equation predicts mass distributions that are in good ag…
This paper deals with extinction of an isolated population caused by intrinsic noise. We model the population dynamics in a "refuge" as a Markov process which involves births and deaths on discrete lattice sites and random migrations between neighboring sites. In extinction scenario I the zero population size is a repelling fixed point of the on-site deterministic dynamics. In extinction scenario II the zero population size is an attracting fixed point, corresponding to what is known in ecology as Allee effect. Assuming a large population size, we develop WKB (Wentzel-Kramers-Brillouin) approx…
