I examine the effect of exogenous spatial heterogeneity on the coexistence of competing species using a simple model of non-hierarchical competition for site occupancy on a lattice. The sites on the lattice are divided into two types representing two different habitats or spatial resources. The model features no temporal variability, hierarchical competition, type-dependent interactions or other features traditionally known to support more competing species than there are resources. Nonetheless, stable coexistence of two habitat specialists and a generalist is observed in this model for a range of parameter values. In the spatially implicit mean field approximation of the model, such coexistence is shown to be impossible, demonstrating that it indeed arises from the explicit spatial structure.
Resource competition theory predicts coexistence and exclusion patterns based on species R*s, the minimum resource values required for a species to persist. A central assumption of the theory is that all species have equal access to resources. However, many systems are characterized by preemption exploitation, where some species deplete resources before their competitors can access them (e.g., asymmetric light competition, contest competition among animals). We hypothesize that coexistence under preemption requires an R*-preemption tradeoff, i.e., the species with the priority access should ha…
We use the optimal foraging theory to study coexistence between two plant species and a generalist pollinator. We compare conditions for plant coexistence for non-adaptive vs. adaptive pollinators that adjust their foraging strategy to maximize fitness. When pollinators have fixed preferences, we show that plant coexistence typically requires both weak competition between plants for resources (e.g., space or nutrients) and pollinator preferences that are not too biased in favour of either plant. We also show how plant coexistence is promoted by indirect facilitation via the pollinator. When po…
Coexistence of plants depends on their competition for common resources and indirect interactions mediated by shared exploiters or mutualists. These interactions are driven either by changes in animal abundance (density-mediated interactions, e.g., apparent competition), or by changes in animal preferences for plants (behaviorally-mediated interactions). This article studies effects of behaviorally-mediated interactions on two plant population dynamics and animal preference dynamics when animal densities are fixed. Animals can be either adaptive exploiters or adaptive mutualists (e.g., herbivo…
Patterned vegetation is a characteristic feature of many dryland ecosystems. While plant densities on the ecosystem-wide scale are typically low, a spatial self-organisation principle leads to the occurrence of alternating patches of high biomass and patches of bare soil. Nevertheless, intraspecific competition dynamics other than competition for water over long spatial scales are commonly ignored in mathematical models for vegetation patterns. In this chapter, I address the impact of local intraspecific competition on a modelling framework for banded vegetation patterns. Firstly, I show that …
The properties of competition models where all individuals are identical are relatively well-understood; however, juveniles and adults can experience or generate competition differently. We study here less well-known structured competition models in discrete time that allow multiple life history parameters to depend on adult or juvenile population densities. A numerical study with Ricker density-dependence suggested that when competition coefficients acting on juvenile survival and fertility reflect opposite competitive hierarchies, stage structure could foster coexistence. We revisit and expa…
Ecological trade-offs between species are often invoked to explain species coexistence in ecological communities. However, few mathematical models have been proposed for which coexistence conditions can be characterized explicitly in terms of a trade-off. Here we present a model of a plant community which allows such a characterization. In the model plant species compete for sites where each site has a fixed stress condition. Species differ both in stress tolerance and competitive ability. Stress tolerance is quantified as the fraction of sites with stress conditions low enough to allow establ…
If two species exhibit different nonlinear responses to a single shared resource, and if each species modifies the resource dynamics such that this favors its competitor, they may stably coexist. This coexistence mechanism, known as relative nonlinearity of competition, is well understood theoretically, but less is known about its evolutionary properties and its prevalence in real communities. We address this challenge by using adaptive dynamics theory and individual-based simulations to compare community stabilization and evolutionary stability of species that coexist by relative nonlinearity…
Predators often consume multiple prey and by mutually subsidizing a shared predator, the prey may reciprocally harm each other. When predation levels are high, this apparent competition can culminate in a prey species being displaced. Coupling quantitative genetics and Lotka-Volterra models, we study how predator evolution alters this and other ecological outcomes. These models account for a trade-off between the predator's attack rates on two prey species. We provide a mathematical characterization of a strong form of persistence--permanence--for which there is a global attractor bounded away…
In plant-pollinator communities many pollinators are potential generalists and their preferences for certain plants can change quickly in response to changes in plant and pollinator densities. These changes in preferences affect coexistence within pollinator guilds as well as within plant guilds. Using a mathematical model, we study how adaptations of pollinator preferences influence population dynamics of a two-plant-two-pollinator community interaction module. Adaptation leads to coexistence between generalist and specialist pollinators, and produces complex plant population dynamics, involv…
We present properties of Lotka-Volterra equations describing ecological competition among a large number of competing species. First we extend to the case of a non-homogeneous niche space stability conditions for solutions representing species coexistence. Second, we discuss mechanisms leading to species clustering and obtain an analytical solution for a lumped state in a specific instance of the system. We also discuss how realistic ecological interactions may result in different types of competition coefficients.
Explaining coexistence in species-rich communities of primary producers remains a challenge for ecologists because of their likely competition for shared resources. Following Hutchinson's seminal suggestion, many theoreticians have tried to create diversity through a fluctuating environment, which impairs or slows down competitive exclusion. However, fluctuating-environment models often only produce a dozen of coexisting species at best. Here, we investigate how to create richer communities in fluctuating environments, using an empirically parameterized model. Building on the forced Lotka-Volt…
We introduce and analyze a spatial Lotka-Volterra competition model with local and nonlocal interactions. We study two alternative classes of nonlocal competition that differ in how each species' characteristics determine the range of the nonlocal interactions. In both cases, nonlocal interactions can create spatial patterns of population densities in which highly populated clumps alternate with unpopulated regions. This non-populated regions provide spatial niches for a weaker competitor to establish in the community and persist in conditions in which local models predict competitive exclusio…
Similarity of competitors has been proposed to facilitate coexistence of species because it slows down competitive exclusion, thus making it easier for equalizing mechanisms to maintain diverse communities. On the other hand, chaos can promote coexistence of species. Here we link these two previously unrelated findings, by analyzing the dynamics of food web models. We show that near-neutrality of competition of prey, in the presence of predators, increases the chance of developing chaotic dynamics. Moreover we confirm that this results in a higher biodiversity. Our results suggest that near-ne…
Local coexistence of species in large ecosystems is traditionally explained within the broad framework of niche theory. However, its rationale hardly justifies rich biodiversity observed in nearly homogeneous environments. Here we consider a consumer-resource model in which a coarse-graining procedure accounts for a variety of ecological mechanisms and leads to effective spatial effects which favour species coexistence. Herein, we provide conditions for several species to live in an environment with very few resources. In fact, the model displays two different phases depending on whether the n…
RNA viruses exist as genetically diverse populations displaying different phenotypes, including diverse degrees of virulence. The evolution of virulence in viral populations is, however, poorly understood. Based on the experimental observation of an RNA virus clone in cell culture diversifying into two subpopulations of different virulence, we study the dynamics of heterogeneous virus populations and the evolution of virulence. We introduce a competition-colonization trade-off into standard mathematical models of intra-host viral infection. Colonizers are fast spreading, virulent strains, wher…
We introduce an asymmetric noisy voter model to study the joint effect of immigration and a competition-dispersal tradeoff in the dynamics of two species competing for space in regular lattices. Individuals of one species can invade a nearest-neighbor site in the lattice, while individuals of the other species are able to invade sites at any distance but are less competitive locally, i.e., they establish with a probability $g \le 1$. The model also accounts for immigration, modeled as an external noise that may spontaneously replace an individual at a lattice site by another individual of the …
The outcome of competition among species is influenced by the spatial distribution of species and effects such as demographic stochasticity, immigration fluxes, and the existence of preferred habitats. We introduce an individual-based model describing the competition of two species and incorporating all the above ingredients. We find that the presence of habitat preference --- generating spatial niches --- strongly stabilizes the coexistence of the two species. Eliminating habitat preference --- neutral dynamics --- the model generates patterns, such as distribution of population sizes, practi…
This letter introduced a new R package 'coexist' which can perform species coexistence simulation and analysis. The package was initially developed for understanding the role of different combinations of varying species growth rates, dispersal ability, habitat heterogeneity, and competition interaction on influencing species coexistence under fluctuating environment. Further versions could introduce more coexistence models under different ecological conditions. The source package is available at the Google Code (https://code.google.com/p/coexist/) and R CRAN repository (http://cran.r-project.o…
Species sharing a prey or a predator species may go extinct due to exploitative or apparent competition. We examine whether evolution of the shared species acts as a coexistence mechanism and to what extent the answer depends on the genetic architecture underlying trait evolution. In our models of exploitative and apparent competition, the shared species evolves its defense or prey use. Evolving species are either haploid or diploid. A single locus pleiotropically determines prey nutritional quality and predator attack rates. When pleiotropy is sufficiently antagonistic (e.g. nutritional prey …
Reaction-diffusion systems with a Lotka-Volterra-type reaction term, also known as competition-diffusion systems, have been used to investigate the dynamics of the competition among $m$ ecological species for a limited resource necessary to their survival and growth. Notwithstanding their rather simple mathematical structure, such systems may display quite interesting behaviours. In particular, while for $m=2$ no coexistence of the two species is usually possible, if $m \ge 3$ we may observe coexistence of all or a subset of the species, sensitively depending on the parameter values. Such coex…
Vegetation patterns are a ubiquitous feature of water-deprived ecosystems. Despite the competition for the same limiting resource, coexistence of several plant species is commonly observed. We propose a two-species reaction-diffusion model based on the single-species Klausmeier model, to analytically investigate the existence of states in which both species coexist. Ecologically, the study finds that coexistence is supported if there is a small difference in the plant species' average fitness, measured by the ratio of a species' capabilities to convert water into new biomass to its mortality r…
The storage effect is a general explanation for coexistence in a variable environment. The generality of the storage effect is both a strength - it can be quantified in many systems - and a challenge - there is not a clear relationship between the abstract conditions for storage effect and species' life-history traits (e.g., dormancy, stage-structure, non-overlapping generations), thus precluding a simple ecological interpretation of the storage effect. Our goal here is to provide a clearer understanding of the conditions for the storage effect as a step towards a better general explanation fo…
We examine the two-dimensional extension of the model of Kessler and Sander of competition between two species identical except for dispersion rates. In this class of models, the spatial inhomogeneity of reproduction rates gives rise to an implicit cost of dispersal, due to the tendency to leave favorable locations. Then, as in the Hamilton-May model with its explicit dispersal cost, the tradeoff between dispersal case and the beneficial role of dispersal in limiting fluctuations, leads to an advantage of one dispersal rate over another, and the eventual extinction of the disadvantaged species…
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…
When three species compete cyclically in a well-mixed, stochastic system of $N$ individuals, extinction is known to typically occur at times scaling as the system size $N$. This happens, for example, in rock-paper-scissors games or conserved Lotka-Volterra models in which every pair of individuals can interact on a complete graph. Here we show that if the competing individuals also have a "social temperament" to be either introverted or extroverted, leading them to cut or add links respectively, then long-living state in which all species coexist can occur when both introverts and extroverts a…
The far-reaching consequences of ecological interactions in the dynamics of biological communities remain an intriguing subject. For decades, competition has been a cornerstone in ecological processes, but mounting evidence shows that cooperation does also contribute to the structure of biological communities. Here, we propose a simple deterministic model for the study of the effects of facilitation and competition in the dynamics of such systems. The simultaneous inclusion of both effects produces rich dynamics and captures the context-dependence observed in the formation of ecological commun…
Dynamic interspecific interactions are thought to contribute to fundamental structures of ecological communities. Using the Yodzis framework of dominance control versus founder control versus niche control, the importance of intransitive loops in sub-communities in a source/sink or metacommunity competition context is examined. It is found that a three-species sub-community sampled from a species pool composed of nothing but dominance controlled species pairs can result in a stable three species coexistence. It is shown that if effect and response competition are balanced, the overall sub-comm…
In order to model real ecological systems one has to consider many species that interact in complex ways. However, most of the recent theoretical studies have been restricted to few species systems with rather trivial interactions. The few studies dealing with larger number of species and/or more complex interaction schemes are mostly restricted to numerical explorations. In this paper we determine, starting from the deterministic mean-field rate equations, for large classes of systems the space of coexistence fixed points at which biodiversity is maximal. For systems with a single coexistence…
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…
Networks provide a mathematically rich framework to represent social contacts sufficient for the transmission of disease. Social networks are often highly clustered and fail to be locally tree-like. In this paper, we study the effects of clustering on the spread of sequential strains of a pathogen using the generating function formulation under a complete cross-immunity coupling, deriving conditions for the threshold of coexistence of the second strain. We show that clustering reduces the coexistence threshold of the second strain and its outbreak size in Poisson networks, whilst exhibiting th…
Compartmentalization of self-replicating molecules (templates) in protocells is a necessary step towards the evolution of modern cells. However, coexistence between distinct template types inside a protocell can be achieved only if there is a selective pressure favoring protocells with a mixed template composition. Here we study analytically a group selection model for the coexistence between two template types using the diffusion approximation of population genetics. The model combines competition at the template and protocell levels as well as genetic drift inside protocells. At the steady s…
In this paper we consider a competition system in which two diseases spread by contact. We characterize the system behavior, establishing that only some configurations are possible. In particular we discover that coexistence of the two strains is not possible, under the assumptions of the model. A number of transcritical bifurcations relate the more relevant system's equilibria. Bistability is shown between a situation in which only the disease-unaffected population thrives and another one containing only the second population with endemic disease. An accurate computation of the separating sur…
We perform individual-based Monte Carlo simulations in a community consisting of two predator species competing for a single prey species, with the purpose of studying biodiversity stabilization in this simple model system. Predators are characterized with predation efficiency and death rates, to which Darwinian evolutionary adaptation is introduced. Competition for limited prey abundance drives the populations' optimization with respect to predation efficiency and death rates. We study the influence of various ecological elements on the final state, finding that both indirect competition and …
Intransitivity is a property of connected, oriented graphs representing species interactions that may drive their coexistence even in the presence of competition, the standard example being the three species Rock-Paper-Scissors game. We consider here a generalization with four species, the minimum number of species allowing other interactions beyond the single loop (one predator, one prey). We show that, contrary to the mean field prediction, on a square lattice the model presents a transition, as the parameter setting the rate at which one species invades another changes, from a coexistence t…
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…
Resource competition is a fundamental interaction in natural communities.However little is known about competition in spatial environments where organisms are able to regulate resource distributions. Here, we analyze the competition of two consumers for two resources in a one-dimensional habitat in which the resources are supplied from opposite sides. We show that the success of an invading species crucially depends on the slope of the resource gradients shaped by the resident. Our analysis reveals that parameter combinations which lead to coexistence in a uniform environment may favor alterna…
We consider the dynamics of a virus spreading through a population that produces a mutant strain with the ability to infect individuals that were infected with the established strain. Temporary cross-immunity is included using a time delay, but is found to be a harmless delay. We provide some sufficient conditions that guarantee local and global asymptotic stability of the disease-free equilibrium and the two boundary equilibria when the two strains outcompete one another. It is shown that, due to the immune evasion of the emerging strain, the reproduction number of the emerging strain must be…
The dynamics of the spread of contagions such as viruses, infectious diseases or even rumors/opinions over contact networks (graphs) have effectively been captured by the well known \textit{Susceptible-Infected-Susceptible} ($SIS$) epidemic model in recent years. When it comes to competition between two such contagions spreading on overlaid graphs, their propagation is captured by so-called \textit{bi-virus} epidemic models. Analysis of such dynamical systems involve the identification of equilibrium points and its convergence properties, which determine whether either of the viruses dies out,…
