Biodiversity is essential to the viability of ecological systems. Species diversity in ecosystems is promoted by cyclic, non-hierarchical interactions among competing populations. Such non-transitive relations lead to an evolution with central features represented by the `rock-paper-scissors' game, where rock crushes scissors, scissors cut paper, and paper wraps rock. In combination with spatial dispersal of static populations, this type of competition results in the stable coexistence of all species and the long-term maintenance of biodiversity. However, population mobility is a central feature of real ecosystems: animals migrate, bacteria run and tumble. Here, we observe a critical influence of mobility on species diversity. When mobility exceeds a certain value, biodiversity is jeopardized and lost. In contrast, below this critical threshold all subpopulations coexist and an entanglement of travelling spiral waves forms in the course of temporal evolution. We establish that this phenomenon is robust, it does not depend on the details of cyclic competition or spatial environment. These findings have important implications for maintenance and evolution of ecological systems and are relevant for the formation and propagation of patterns in excitable media, such as chemical kinetics or epidemic outbreaks.
This work deals with a system of three distinct species that changes in time under the presence of mobility, selection, and reproduction, as in the popular rock-paper-scissors game. The novelty of the current study is the modification of the mobility rule to the case of directional mobility, in which the species move following the direction associated to a larger (averaged) number density of selection targets in the surrounding neighborhood. Directional mobility can be used to simulate eyes that see or a nose that smells, and we show how it may contribute to reduce the probability of coexisten…
We investigate the impact of cyclic competition on pattern formation in the rock-paper-scissors game. By separately considering random and prepared initial conditions, we observe a critical influence of the competition rate $p$ on the stability of spiral waves and on the emergence of biodiversity. In particular, while increasing values of $p$ promote biodiversity, they may act detrimental on spatial pattern formation. For random initial conditions, we observe a phase transition from biodiversity to an absorbing phase, whereby the critical value of mobility grows linearly with increasing values…
Rock-paper-scissors (RPS) game is a nice model to study the biodiversity in ecosystem. However, the previous studies only consider the nearest- neighbor- interaction among the species. In this paper, taking the long range migration into account, the effects of the interplay between nearest-neighbor-interaction and long-range-interaction of Levy flight obey the power law distance distribution with the exponent h (-0.3<h<-0.1) in spatial RPS game is investigated. Taking the probability of long range Levy flight and the power exponent as parameters, the coexistence conditions of three species are…
We study the role of the adaptive movement strategy in promoting biodiversity in cyclic models described by the rock-paper-scissors game rules. We assume that individuals of one out of the species may adjust their movement to escape hostile regions and stay longer in their comfort zones. Running a series of stochastic simulations, we calculate the alterations in the spatial patterns and population densities in scenarios where not all organisms are physically or cognitively conditioned to perform the behavioural strategy. Although the adaptive movement strategy is not profitable in terms of ter…
The dynamical mechanisms that can stabilize the coexistence of species (or strategies) are of substantial interest for the maintenance of biodiversity and in sociobehavioural dynamics. We investigate the mean extinction time in the coevolutionary dynamics of three cyclically invading strategies for different evolutionary processes on various classes of complex networks, including random graphs, scale-free and small world networks. We find that scale-free and random graphs lead to a strong stabilization of coexistence both for the Moran process and the Local Update process. The stabilization is…
We investigate a tritrophic system whose cyclic dominance is modelled by the rock-paper-scissors game. We consider that organisms of one or two species are affected by movement limitations, which unbalances the cyclic spatial game. Performing stochastic simulations, we show that mobility unevenness controls the population dynamics. In the case of one slow species, the predominant species depends on the level of mobility restriction, with the slow species being preponderant if the mobility limitations are substantial. If two species face mobility limitations, our outcomes show that being higher…
The rock-paper-scissors game is a paradigmatic model for biodiversity, with applications ranging from microbial populations to human societies. Research has shown, however, that mobility jeopardizes biodiversity by promoting the formation of spiral waves, especially if there is no conservation law in place for the total number of competing players. Firstly, we show that even if such a conservation law applies, mobility still jeopardizes biodiversity in the spatial rock-paper-scissors game if only a small fraction of links of the square lattice is randomly rewired. Secondly, we show that zealot…
This work reports on two related investigations of stochastic simulations which are widely used to study biodiversity and other related issues. We first deal with the behavior of the Hamming distance under the increase of the number of species and the size of the lattice, and then investigate how the mobility of the species contributes to jeopardize biodiversity. The investigations are based on the standard rules of reproduction, mobility and predation or competition, which are described by specific rules, guided by generalization of the rock-paper-scissors game, valid in the case of three spe…
If rock beats scissors and scissors beat paper, one might assume that rock beats paper too. But this is not the case for intransitive relationships that make up the famous rock-paper-scissors game. However, the sole presence of paper might prevent rock from beating scissors, simply because paper beats rock. This is the blueprint for the rock-paper-scissors game with protection spillovers, which has recently been introduced as a new paradigm for biodiversity in well-mixed microbial populations. Here we study the game in structured populations, demonstrating that protection spillovers give rise …
Non-transitive dominance and the resulting cyclic loop of three or more competing species provide a fundamental mechanism to explain biodiversity in biological and ecological systems. Both Lotka-Volterra and May-Leonard type model approaches agree that heterogeneity of invasion rates within this loop does not hazard the coexistence of competing species. While the resulting abundances of species become heterogeneous, the species who has the smallest invasion power benefits the most from unequal invasions. Nevertheless, the effective invasion rate in a predator and prey interaction can also be m…
This work deals with the influence of the neighborhood in simple rock-paper-scissors models of biodiversity. We consider the case of three distinct species which evolve under the standard rules of mobility, reproduction and competition. The rule of competition follows the guidance of the rock-paper-scissors game, with the prey being annihilated, leaving an empty site in accordance with the May-Leonard proposal for the predator and prey competition. We use the von Neumann neighborhood, but we consider mobility under the presence of the first, second and third neighbors in three distinct environ…
The spatial rock-paper-scissors ecosystem, where three species interact cyclically, is a model example of how spatial structure can maintain biodiversity. We here consider such a system for a broad range of interaction rates. When one species grows very slowly, this species and its prey dominate the system by self-organizing into a labyrinthine configuration in which the third species propagates. The cluster size distributions of the two dominating species have heavy tails and the configuration is stabilized through a complex, spatial feedback loop. We introduce a new statistical measure that …
Stochastic simulations of cyclic three-species spatial predator-prey models are usually performed in square lattices with nearest neighbor interactions starting from random initial conditions. In this Letter we describe the results of off-lattice Lotka-Volterra stochastic simulations, showing that the emergence of spiral patterns does occur for sufficiently high values of the (conserved) total density of individuals. We also investigate the dynamics in our simulations, finding an empirical relation characterizing the dependence of the characteristic peak frequency and amplitude on the total de…
When faced with an imminent risk of predation, many animals react to escape consumption. Antipredator strategies are performed by individuals acting as a group to intimidate predators and minimize the damage when attacked. We study the antipredator prey response in spatial tritrophic systems with cyclic species dominance using the rock-paper-scissors game. The impact of the antipredator behavior is local, with the predation probability reducing exponentially with the number of preys in the predator's neighborhood. In contrast to the standard Lotka-Volterra implementation of the rock-paper-scis…
Dynamical mechanisms that can stabilize the coexistence or diversity in biology are generally of fundamental interest. In contrast to many two-strategy evolutionary games, games with three strategies and cyclic dominance like the rock-paper-scissors game (RPS) stabilize coexistence and thus preserve biodiversity in this system. In the limit of infinite populations, resembling the traditional picture of evolutionary game theory, replicator equations predict the existence of a fixed point in the interior of the phase space. But in finite populations, strategy frequencies will run out of the fixe…
Biodiversity widely observed in ecological systems is attributed to the dynamical balance among the competing species. The time-varying populations of the interacting species are often captured rather well by a set of deterministic replicator equations in the evolutionary game theory. However, intrinsic fluctuations arisen from the discreteness of populations lead to stochastic derivations from the smooth evolution trajectories. The role of these fluctuations is shown to be critical at causing extinction and deteriorating the biodiversity of ecosystem. We use children's rock-paper-scissors gam…
We study the cyclic dominance of three species in two-dimensional constrained Newman-Watts networks with a four-state variant of the rock-paper-scissors game. By limiting the maximal connection distance $R_{max}$ in Newman-Watts networks with the long-rang connection probability $p$, we depict more realistically the stochastic interactions among species within ecosystems. When we fix mobility and vary the value of $p$ or $R_{max}$, the Monte Carlo simulations show that the spiral waves grow in size, and the system becomes unstable and biodiversity is lost with increasing $p$ or $R_{max}$. Thes…
Decoding species coexistence: A reinforcement learning perspective
School of Physics and Information Technology, Shaanxi Normal University（陕西师范大学物理与信息技术学院）；School of Physics and Electronic Science, East China Normal University（华东师范大学物理与电子科学学院）；School of Computer Science, Soochow University（苏州大学计算机科学与技术学院）；School of Physics, Ningxia University（宁夏大学物理学院）
Antipredator behaviour is a self-preservation strategy present in many biological systems, where individuals join the effort in a collective reaction to avoid being caught by an approaching predator. We study a nonhierarchical tritrophic system, whose predator-prey interactions are described by the rock-paper-scissors game rules. We performed a set of spatial stochastic simulations where organisms of one out of the species can resist predation in a collective strategy. The drop in predation capacity is local, which means that each predator faces a particular opposition depending on the prey gr…
Cyclic, nonhierarchical interactions among biological species represent a general mechanism by which ecosystems are able to maintain high levels of biodiversity. However, species coexistence is often possible only in spatially extended systems with a limited range of dispersal, whereas in well-mixed environments models for cyclic competition often lead to a loss of biodiversity. Here we consider the dispersal of biological species in a fluid environment, where mixing is achieved by a combination of advection and diffusion. In particular, we perform a detailed numerical analysis of a model comp…
Neutral species facilitate coexistence among cyclically competing species under birth and death processes
School of Statistics and Mathematics, Yunnan University of Finance and Economics（云南财经大学统计与数学学院）；School of Mathematics and Statistics, Xidian University（西安电子科技大学数学与统计学院）；Department of Applied Mathematics, Colleage of Applied Sciences, Kyung Hee University（韩国庆熙大学应用科学学院应用数学系）；G-LAMP NEXUS Institute, Kyung Hee University（韩国庆熙大学G-LAMP NEXUS研究所）
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…
We study the generalised rock-paper-scissors game with five species whose organisms face local epidemic outbreaks. As an evolutionary behavioural survival strategy, organisms of one out of the species move in the direction with more enemies of their enemies to benefit from protection against selection. We consider that each organism scans the environment, performing social distancing instead of agglomerating when perceiving that the density of sick organisms is higher than a tolerable threshold. Running stochastic simulations, we study the interference of the adaptive movement survival strateg…
Population structure induced by both spatial embedding and more general networks of interaction, such as model social networks, have been shown to have a fundamental effect on the dynamics and outcome of evolutionary games. These effects have, however, proved to be sensitive to the details of the underlying topology and dynamics. Here we introduce a minimal population structure that is described by two distinct hierarchical levels of interaction. We believe this model is able to identify effects of spatial structure that do not depend on the details of the topology. We derive the dynamics gove…
