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Cyclic dominance models, such as the classic rock-paper-scissors (RPS) game, have found real-world applications in biology, ecology, and sociology. A key quantity of interest in such models is the coexistence time, i.e., the time until at least one population type goes extinct. Much recent research has considered conditions that lengthen coexistence times in an RPS model. A general finding is that coexistence is promoted by localized spatial interactions (low mobility), while extinction is fostered by global interactions (high mobility). That is, there exists a mobility threshold which separates a regime of long coexistence from a regime of rapid collapse of coexistence. The key finding of our paper is that if zealots (i.e., nodes able to defeat others while themselves being immune to defeat) of even a single type exist, then system coexistence time can be significantly prolonged, even in the presence of global interactions. This work thus highlights a crucial determinant of system survival time in cyclic dominance models. [ABSTRACT FROM AUTHOR]

When playing games in groups, it is an advantage for individuals to have accurate statistical information on the strategies of their opponents. Such information may be obtained by remembering previous interactions. We consider a rock-paper-scissors game in which agents are able to recall their last m interactions, used to estimate the behavior of their opponents. At critical memory length, a Hop! bifurcation leads to the formation of stable limit cycles. In a mixed population, agents with longer memories have an advantage, provided the system has a stable fixed point, and there is some asymmetry in the payoffs of the pure strategies. However, at a critical concentration of long memory agents, the appearance of limit cycles destroys their advantage. By introducing population dynamics that favors successful agents, we show that the system evolves toward the bifurcation point. [ABSTRACT FROM AUTHOR]

We analyze the replicator-mutator equations for the rock-paper-scissors game. Various graph-theoretic patterns of mutation are considered, ranging from a single unidirectional mutation pathway between two of the species, to global bidirectional mutation among all the species. Our main result is that the coexistence state, in which all three species exist in equilibrium, can be destabilized by arbitrarily small mutation rates. After it loses stability, the coexistence state gives birth to a stable limit cycle solution created in a supercritical Hopf bifurcation. This attracting periodic solution exists for all the mutation patterns considered, and persists arbitrarily close to the limit of zero mutation rate and a zero-sum game. [ABSTRACT FROM AUTHOR]

The spatiotemporal arrangement of interacting populations often influences the maintenance of species diversity and is a subject of intense research. Here, we study the spatiotemporal patterns arising from the cyclic competition between three species in two dimensions. Inspired by recent experiments, we consider a generic metapopulation model comprising "rock-paper-scissors" interactions via dominance removal and replacement, reproduction, mutations, pair exchange, and hopping of individuals. By combining analytical and numerical methods, we obtain the model's phase diagram near its Hopf bifurcation and quantitatively characterize the properties of the spiraling patterns arising in each phase. The phases characterizing the cyclic competition away from the Hopf bifurcation (at low mutation rate) are also investigated. Our analytical approach relies on the careful analysis of the properties of the complex Ginzburg-Landau equation derived through a controlled (perturbative) multiscale expansion around the model's Hopf bifurcation. Our results allow us to clarify when spatial "rock-paper-scissors" competition leads to stable spiral waves and under which circumstances they are influenced by nonlinear mobility. [ABSTRACT FROM AUTHOR]

We study the rock-paper-scissors game in structured populations, where the invasion rates determine individual payoffs that govern the process of strategy change. The traditional version of the game is recovered if the payoffs for each potential invasion stem from a single pairwise interaction. However, the transformation of invasion rates to payoffs also allows the usage of larger interaction ranges. In addition to the traditional pairwise interaction, we therefore consider simultaneous interactions with all nearest neighbors, as well as with all nearest and next-nearest neighbors, thus effectively going from single pair to group interactions in games of cyclic dominance. We show that differences in the interaction range affect not only the stationary fractions of strategies but also their relations of dominance. The transition from pairwise to group interactions can thus decelerate and even revert the direction of the invasion between the competing strategies. Like in evolutionary social dilemmas, in games of cyclic dominance, too, the indirect multipoint interactions that are due to group interactions hence play a pivotal role. Our results indicate that, in addition to the invasion rates, the interaction range is at least as important for the maintenance of biodiversity among cyclically competing strategies [ABSTRACT FROM AUTHOR]

In this work we investigate the development of stable dynamical structures along interfaces separating domains belonging to enemy partnerships in the context of cyclic predator-prey models with an even number of species N ≥ 8. We use both stochastic and field theory simulations in one and two spatial dimensions, as well as analytical arguments, to describe the association at the interfaces of mutually neutral individuals belonging to enemy partnerships and to probe their role in the development of the dynamical structures at the interfaces. We identify an interesting behavior associated with the symmetric or asymmetric evolution of the interface profiles depending on whether N/2 is odd or even, respectively. We also show that the macroscopic evolution of the interface network is not very sensitive to the internal structure of the interfaces. Although this work focuses on cyclic predator-prey models with an even number of species, we argue that the results are expected to be quite generic in the context of spatial stochastic May-Leonard models. [ABSTRACT FROM AUTHOR]

The rock-paper-scissors game and its generalizations with S > 3 species are well-studied models for cyclically interacting populations. Four is, however, the minimum number of species that, by allowing other interactions beyond the single, cyclic loop, breaks both the full intransitivity of the food graph and the one-predator, one-prey symmetry. Lütz et al. [J. Theor. Biol. 317, 286 (2013)] have shown the existence, on a square lattice, of two distinct phases, with either four or three coexisting species. In both phases, each agent is eventually replaced by one of its predators, but these strategy oscillations remain localized as long as the interactions are short ranged. Distant regions may be either out of phase or cycling through different food-web subloops (if any). Here we show that upon replacing a minimum fraction Q of the short-range interactions by long-range ones, there is a Hopf bifurcation, and global oscillations become stable. Surprisingly, to build such long-distance, global synchronization, the four-species coexistence phase requires fewer long-range interactions than the three-species phase, while one would naively expect the opposite to be true. Moreover, deviations from highly homogeneous conditions (x = 0 or 1) increase Qc, and the more heterogeneous is the food web, the harder the synchronization is. By further increasing Q, while the three-species phase remains stable, the four-species one has a transition to an absorbing, single-species state. The existence of a phase with global oscillations for S > 3, when the interaction graph has multiple subloops and several possible local cycles, leads to the conjecture that global oscillations are a general characteristic, even for large, realistic food webs. [ABSTRACT FROM AUTHOR]

We study adaptive dynamics in games where players abandon the population at a given rate and are replaced by naive players characterized by a prior distribution over the admitted strategies. We demonstrate how such a process leads macroscopically to a variant of the replicator equation, with an additional term accounting for player turnover. We study how Nash equilibria and the dynamics of the system are modified by this additional term for prototypical examples such as the rock-paper-scissors game and different classes of two-action games played between two distinct populations. We conclude by showing how player turnover can account for nontrivial departures from Nash equilibria observed in data from lowest unique bid auctions. [ABSTRACT FROM AUTHOR]

We investigate the population dynamics in generalized rock-paper-scissors models with an arbitrary number of species Ν. We show that spiral patterns with Ν arms may develop both for odd and even N, in particular in models where a bidirectional prédation interaction of equal strength between all species is modified to include one -cyclic predator-prey rule. While the former case gives rise to an interface network with Y-type junctions obeying the scaling law L ∝ t1/2, where L is the characteristic length of the network and t is the time, the latter can lead to a population network with N-armed spiral patterns, having a roughly constant characteristic length scale. We explicitly demonstrate the connection between interface junctions and spiral patterns in these models and compute the corresponding scaling laws. This work significantly extends the results of previous studies of population dynamics and could have profound implications for the understanding of biological complexity in systems with a large number of species. [ABSTRACT FROM AUTHOR]

We introduce a family of rock-paper-scissors-type models with ZN symmetry (N is the number of species), and we show that it has a very rich structure with many completely different phases. We study realizations that lead to the formation of domains, where individuals of one or more species coexist, separated by interfaces whose (average) dynamics is curvature driven. This type of behavior, which might be relevant for the development of biological complexity, leads to an interface network evolution and pattern formation similar to the ones of several other nonlinear systems in condensed matter and cosmology. [ABSTRACT FROM AUTHOR]

The rock-paper-scissors game is a model example of the ongoing cyclic turnover typical of many ecosystems, ranging from the terrestrial and aquatic to the microbial. Here we explore the evolution of a rock-paper-scissors system where three species compete for space. The species are allowed to mutate and change the speed by which they invade one another. In the case when all species have similar mutation rates, we observe a perpetual arms race where no single species prevails. When only two species mutate, their aggressions increase indefinitely until the ecosystem collapses and only the nonmutating species survives. Finally we show that when only one species mutates, group selection removes individual predators with the fastest growth rates, causing the growth rate of the species to stabilize. We explain this group selection quantitatively. [ABSTRACT FROM AUTHOR]

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 detrimentally 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 of p on a log-log scale but then saturates as p becomes large. For prepared initial conditions, we observe the formation of single-armed spirals, but only for values of p that are below a critical value. Once above that value, the spirals break up and form disordered spatial structures, mainly because of the percolation of vacant sites. Thus there exists a critical value of the competition rates pc for stable single-armed spirals in finite populations. Importantly though, pc increases with increasing system size because noise reinforces the disintegration of ordered patterns. In addition, we also find that pc increases with the mobility. These phenomena are reproduced by a deterministic model that is based on nonlinear partial differential equations. Our findings indicate that competition is vital for the sustenance of biodiversity and the emergence of pattern formation in ecosystems governed by cyclical interactions. [ABSTRACT FROM AUTHOR]

The efficiency of institutionalized punishment is studied by evaluating the stationary states in the spatial public goods game comprising unconditional defectors, cooperators, and cooperating pool punishers as the three competing strategies. Fines and costs of pool punishment are considered as the two main parameters determining the stationary distributions of strategies on the square lattice. Each player collects a payoff from five five-person public goods games, and the evolution of strategies is subsequently governed by imitation based on pairwise comparisons at a low level of noise. The impact of pool punishment on the evolution of cooperation in structured populations is significantly different from that reported previously for peer punishment. Representative phase diagrams reveal remarkably rich behavior, depending also on the value of the synergy factor that characterizes the efficiency of investments payed into the common pool. Besides traditional single- and two-strategy stationary states, a rock-paper-scissors type of cyclic dominance can emerge in strikingly different ways. [ABSTRACT FROM AUTHOR]

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 coexistence. [ABSTRACT FROM AUTHOR]

This work deals with the effects of an apex predator on the cyclic competition among three distinct species that follow the rules of the rock-paper-scissors game. The investigation develops standard stochastic simulations but is motivated by a procedure which is explained in the work. We add the apex predator as the fourth species in a system that contains three species that evolve following the standard rules of migration, reproduction, and predation, and study how the system evolves in this new environment, in comparison with the case in the absence of the apex predator. The results show that the apex predator engenders the tendency to spread uniformly in the lattice, contributing to destroy the spiral patterns, keeping biodiversity but diminishing the average size of the clusters of the species that compete cyclically. [ABSTRACT FROM AUTHOR]

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. First, 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 zealots are very effective in taming the amplitude of oscillations that emerge due to mobility and/or interaction randomness, and this regardless of whether the later is quenched or annealed. While even a tiny fraction of zealots brings significant benefits, at 5% occupancy zealots practically destroy all oscillations regardless of the intensity of mobility, and regardless of the type and strength of randomness in the interaction structure. Interestingly, by annealed randomness the impact of zealots is qualitatively the same as by mobility, which highlights that fast diffusion does not necessarily destroy the coexistence of species, and that zealotry thus helps to recover the stable mean-field solution. Our results strengthen the important role of zealots in models of cyclic dominance, and they reveal fascinating evolutionary outcomes in structured populations that are a unique consequence of such uncompromising behavior. [ABSTRACT FROM AUTHOR]

The classical game of rock-paper-scissors has inspired experiments and spatial model systems that address the robustness of biological diversity. In particular, the game nicely illustrates that cyclic interactions allow multiple strategies to coexist for long-time intervals. When formulated in terms of a one-dimensional cellular automata, the spatial distribution of strategies exhibits coarsening with algebraically growing domain size over time, while the two-dimensional version allows domains to break and thereby opens the possibility for long-time coexistence. We consider a quasi-one-dimensional implementation of the cyclic competition, and study the long-term dynamics as a function of rare invasions between parallel linear ecosystems. We find that increasing the complexity from two to three parallel subsystems allows a transition from complete coarsening to an active steady state where the domain size stays finite. We further find that this transition happens irrespective of whether the update is done in parallel for all sites simultaneously or done randomly in sequential order. In both cases, the active state is characterized by localized bursts of dislocations, followed by longer periods of coarsening. In the case of the parallel dynamics, we find that there is another phase transition between the active steady state and the coarsening state within the three-line system when the invasion rate between the subsystems is varied. We identify the critical parameter for this transition and show that the density of active boundaries has critical exponents that are consistent with the directed percolation universality class. On the other hand, numerical simulations with the random sequential dynamics suggest that the system may exhibit an active steady state as long as the invasion rate is finite. [ABSTRACT FROM AUTHOR]

We study an evolutionary game model based on a transition matrix approach, in which the total change in the proportion of a population playing a given strategy is summed directly over contributions from all other strategies. This general approach combines aspects of the traditional replicator model, such as preserving unpopulated strategies, with mutation-type dynamics, which allow for nonzero switching to unpopulated strategies, in terms of a single transition function. Under certain conditions, this model yields an endemic population playing non-Nash-equilibrium strategies. In addition, a Hopf bifurcation with a limit cycle may occur in the generalized rock-scissors-paper game, unlike the replicator equation. Nonetheless, many of the Folk Theorem results are shown to hold for this model. [ABSTRACT FROM AUTHOR]

In spatial evolutionary games the payoff matrices are used to describe pair interactions among neighboring players located on a lattice. Now we introduce a way how the payoff matrices can be built up as a sum of payoff components reflecting basic symmetries. For the two-strategy games this decomposition reproduces interactions characteristic to the Ising model. For the three-strategy symmetric games the Fourier components can be classified into four types representing games with self-dependent and cross-dependent payoffs, variants of three-strategy coordinations, and the rock-scissors-paper (RSP) game. In the absence of the RSP component the game is a potential game. The resultant potential matrix has been evaluated. The general features of these systems are analyzed when the game is expressed by the linear combinations of these components. [ABSTRACT FROM AUTHOR]