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In a diverse population, where many species are present, competitors can fight for surviving at individual and collective levels. In particular, species, which would beat each other individually, may form a specific alliance that ensures them stable coexistence against the invasion of an external species. Our principal goal is to identify those general features of a formation which determine its vitality. Therefore, we here study a traditional Lotka-Volterra model of eight-species where two four-species cycles can fight for space. Beside these formations, there are other solutions which may emerge when invasion rates are varied. The complete range of parameters is explored and we find that in most of the cases those alliances prevail which are formed by equally strong members. Interestingly, there are regions where the symmetry is broken and the system is dominated by a solution formed by seven species. Our work also highlights that serious finite-size effects may emerge which prevent observing the valid solution in a small system., Comment: 10 double-column pages, 11 figures

We investigate evolving dynamics of cyclically competing species on spatially extended systems with considering a specific region, which is called the “wildlife refuge,” one of the institutional ways to preserve species biodiversity. Through Monte-Carlo simulations, we found that the refuge can play not groundbreaking but an important role in species survival. Species coexistence is maintained at a moderate mobility regime, which traditionally leads to the collapse of coexistence, and eventually, the extinction is postponed depending on the competition rate rather than the portion of the refuge. Incorporating the extinction probability and Fourier transform supported our results in both stochastic and analogous ways. Our findings may provide valuable evidence to assist fields of ecological/biological sciences in understanding the presence and construction of refuges for wildlife with associated effects on species biodiversity.

Securing space for species breeding is important in the evolution and maintenance of life in ecological sciences, and an increase in the number of competing species may cause frequent competition and conflict among the population in securing such spaces in a given area. In particular, for cyclically competing species, which can be described by the metaphor of rock–paper–scissors game, most of the previous works in microscopic frameworks have been studied with the initially given three species without any formation of additional competing species, and the phase transition of biodiversity via mobility from coexistence to extinction has never been changed by a change of spatial scale. In this regard, we investigate the relationship between spatial scales and species coexistence in the spatial cyclic game by considering the emergence of a new competing group by mutation. For different spatial scales, our computations reveal that coexistence can be more sensitive to spatial scales and may require larger spaces for frequencies of interactions. By exploiting the calculation of the coexistence probability from Monte-Carlo simulations, we obtain that certain interaction ranges for coexistence can be affected by both spatial scales and mobility, and spatial patterns for coexistence can appear in different ways. Since the issue of spatial scale is important for species survival as competing populations increase, we expect our results to have broad applications in the fields of social and ecological sciences.

Multistability is common feature resulting in nonlinear dynamical systems, and its characteristic can be generally depicted by investigating basin structures of initial conditions for give parameter settings. In this paper, we explore the formation of extinction states according to the change of strength of competition levels in the toy model for three species. Through the linear stability analysis, we find that the extinction state can be stable which is persistent. For specific conditions between intensities of two different competitions, we also found that the extinction state can be either bistable or tristable. In each case, the final state of the system can be characterized sensitively depending on initial conditions. To validate our results, we investigate basin structures of parameters for interspecific competition associated to a strength of intraspecific competition. In addition, we found that coexistence becomes robust as intraspecific competition is intensified relatively to the interspecific competition level. We hope our results can be a chance to suggest the emergence of the multistability according to complex competition structures on systems of many populations.

Asymmetrical rock-paper-scissors (RPS) competition has been perceived as a crucial factor in shaping species biodiversity, and understanding this ecological issue in a multi-species paradigm is rather difficult because community dynamics usually depend on distinct factors such as abiotic environments, biotic interactions and symmetry-breaking phenomenon. To address this problem, we employ a Lotka-Volterra competitive system consisting of both symmetrical, asymmetrical interactions and abiotic environment components. We discover that that asymmetrical RPS competition in heterogeneous environments can yield much richer dynamical behaviors, compared to the symmetrical and asymmetrical competition in homogeneous environments. While it is observed that species coexistence outcomes and/or oscillatory solutions are maintained as in the case of homogeneous environments, the nonuniformity in the environmental carrying capacities may lead to extra dynamics with regards to the appearance of survival states; for instance, coexistence of any two-species and single-species persistence states, which are not evident in the previous modelling studies. By means of bifurcation analysis, various salient features of the dynamical systems, including the emergence of certain attractors (e.g., different steady states, stable limit cycles and heteroclinic cycles) and co-dimension one bifurcations (e.g., transcritical and supercritical Hopf bifurcations) are realized in this ecological model. Overall, this modelling work provides a novel attempt to simultaneously encompass not only symmetry-breaking phenomenon through RPS competition, but also heterogeneity in the environments. This framework can provide additional insights to better understand various mechanisms underlying the effects of distinct ecological processes on multi-species communities.

Interpatch migration between two environments is generally considered as a spatial concept and can affect species biodiversity in each patch by inducing flux of population such as inflow and outflow quantities of species. In this paper, we explore the effect of interpatch migration, which can be generally considered as a spatial concept and may affect species biodiversity between two different patches in the perspective of the macroscopic level by exploiting the coupling of two systems, where each patch is occupied by cyclically competing three species who can stably coexist by exhibiting periodic orbits. For two simple scenarios of interpatch migration either single or all species migration, we found that two systems with independently stable coexisting species in each patch are eventually synchronized, and oscillatory behaviors of species densities in two patches become identical, i.e., the synchronized coexistence emerges. In addition, we find that, whether single or all species interpatch migration occurs, the waiting time for the synchronization is exponentially decreasing as the coupling strength is intensified. Our findings suggest that the synchronized behavior of species as a result of migration between different patches can be easily predicted by the coupling of systems and additional information such as waiting times and sensitivity of initial densities.

Balancedness among species interactions may be an important key to understand species biodiversity. Biodiversity among species is usually promoted by competitions which can occur between two different species or among the same species. In this paper, we investigate how symmetry breaking of interspecific competitions can affect biodiversity on cyclic structured three species which may compete with themselves. From theoretical and numerical results of the deterministic system, we found that the symmetry breaking of interspecific competitions on the self-competitive species system can lead the emergence of new survival states in which are stable. Further, we figured out that these diverse survival states can be influenced by the moderate balance between interspecific and intraspecific competitions which is uncovered numerically.

One of the common assumptions in previous spatial dynamics of cyclic competition is that, regardless of competing structure and strength among species, the spatial size of a network is considered as large as possible to avoid finite size effect for species biodiversity. In real ecosystems, however, species richness, which can be defined by spatial size and competition strength, can sensitively affect species coexistence as a competition among individuals becomes complicated. In this paper, we investigate the structural stability of coexistence of mobile species in three cyclic competition games due to network complexity in which imposes a size of a square lattice and competition strength among species. By exploiting the coexistence probability, our computations quantitatively reveal that the network complexity due to changes in the competition rate and lattice size can strongly affect the structural stability of coexistence in each model. In particular, intense intraspecific competition can yield the robust coexistence at small-sized lattices regardless of mobility, and strengthening interspecific competition simultaneously induces changes in critical mobility that hampers coexistence and in spatial size for stable coexistence. Qualitatively, we find that such structural stability of coexistence relates to the degree of stability of fixed points in deterministic systems. Our finding can be useful to gain insights into species coexistence on spatially extended systems with respect to network complexity.

Classic frameworks of rock-paper-scissors game have been assumed in a closed community that a density of each group is only affected by internal factors such as competition interplay among groups and reproduction itself. In real systems in ecological and social sciences, however, the survival and a change of a density of a group can be also affected by various external factors. One of common features in real population systems in ecological and social sciences is population flow that is characterized by population inflow and outflow in a group or a society, which has been usually overlooked in previous works on models of rock-paper-scissors game. In this paper, we suggest the rock-paper-scissors system by implementing population flow and investigate its effect on biodiversity. For two scenarios of either balanced or imbalanced population flow, we found that the population flow can strongly affect group diversity by exhibiting rich phenomena. In particular, while the balanced flow can only lead the persistent coexistence of all groups which accompanies a phase transition through supercritical Hopf bifurcation on different carrying simplices, the imbalanced flow strongly facilitates rich dynamics such as alternative stable survival states by exhibiting various group survival states and multistability of sole group survivals by showing not fully covered but spirally entangled basins of initial densities due to local stabilities of associated fixed points. In addition, we found that, the system can exhibit oscillatory dynamics for coexistence by relativistic interplay of population flows which can capture the robustness of the coexistence state. Applying population flow in the rock-paper-scissors system can ultimately change a community paradigm from closed to open one, and our foundation can eventually reveal that population flow can be also a significant factor on a group density which is independent to fundamental interactions among groups.

Cyclically competition models have been successful to gain an insight of biodiversity mechanism in ecosystems. There are, however, still limitations to elucidate complex phenomena arising in real competition. In this paper, we report that a multistability occurs in a simple rock-paper-scissor cyclically competition model by assuming that intraspecific competition depends on the logistic growth of each species density. This complex stability is absent in any cyclically competition model, and we investigate how the proposed intraspecific competition affects biodiversity in the existing society of three species through macroscopic and microscopic approaches. When the system is multistable, we show basins of the asymptotically stable heteroclinic cycle and stable attractors to demonstrate how the survival state is determined by initial densities of three species. Also, we find that the multistability is associated with a subcritical Hopf bifurcation. This surprising finding will give an opportunity to interpret rich dynamical phenomena in ecosystems which may occur in cyclic competition systems with different types of interactions.

Understanding mechanisms which promote or hinder existing ecosystems are important issues in ecological sciences. In addition to fundamental interactions such as competition and migration among native species, existing ecosystems can be easily disturbed by external factors, and the emergence of new species may be an example in such cases. The new species which does not exist in a current ecosystem can be regarded as either alien species entered from outside or mutant species born by mutation in existing normal species. Recently, as existing ecosystems are getting influenced by various physical/chemical external factors, mutation due to anthropogenic and environmental factors can occur more frequently and is thus attracting much attention for the maintenance of ecosystems. In this paper, we consider emergences of mutant species among self-competing three species in the cyclic dominance. By defining mutation as the birth of mutant species, we investigate how mutant species can affect biodiversity in the existing ecosystem. Through microscopic and macroscopic approaches, we have found that the society of existing normal species can be disturbed by mutant species either the society is maintained accompanying with the coexistence of all species or jeopardized by occupying of mutant species. Due to the birth of mutant species, the existing society may be more complex by constituting two different groups of normal and mutant species, and our results can be contributed to analyze complex ecosystems of many species. We hope our findings may propose a new insight on mutation in cyclic competition systems of many species.

We study the dynamics of cyclic competing mobile five species on spatially extended systems originated from asymmetric initial populations and investigate the basins for the three possible asymptotic states, coexistence of all species, existences of only two independent species, and the extinction. Through extensive numerical simulations, we find a prosperous dependence on initial conditions for species biodiversity. In particular, for fixed given equal densities of two relevant species, we find that only five basins for the existence of two independent species exist and they are spirally entangled for high mobility. A basin of coexistence is outbreaking when the mobility parameter is decreased through a critical value and surrounded by the other five basins. For fixed given equal densities of two independent species, however, we find that basin structures are not spirally entangled. Further, final states of two independent species are totally different. For all possible considerations, the extinction state is not witnessed which is verified by the survival probability. To provide the validity of basin structures from lattice simulations, we analyze the system in mean-field manners. Consequently, results on macroscopic levels are matched to direct lattice simulations for high mobility regimes. These findings provide a good insight into the fundamental issue of the biodiversity among many species than previous cases.

Alternative strategy is common in animal populations to promote reproductive fitness by obtaining resources. In spatial dynamics of cyclic competition, reproduction can occur when individuals obtain vacant rooms and, in this regard, empty sites should be resources for reproduction which can be induced by interspecific competition. In this paper, we study the role of alternative competition in the spatial system of cyclically competing five species by utilizing rock-paper-scissors-lizard-spock game. From Monte-Carlo simulations, we found that strong alternative competition can lead to the reemergence of coexistence of five species regardless of mobility, which is never reported in previous works under the symmetric competition structure. By investigating the coexistence probability, we also found that coexistence alternates by passing certain degrees of alternative competition in combination with mobility. In addition, we provided evidences in the opposite scenario by strengthening spontaneous competition, which exhibits the reemergence of coexistence similarly. Our findings may suggest more comprehensive perspectives to interpret mechanisms for biodiversity by alternative strategies in spatially extended systems than previously reported.

The role of mutation, which is an error process in gene evolution, in systems of cyclically competing species has been studied from various perspectives, and it is regarded as one of the key factors for promoting coexistence of all species. In addition to naturally occurring mutations, many experiments in genetic engineering have involved targeted mutation techniques such as recombination between DNA and somatic cell sequences and have studied genetic modifications through loss or augmentation of cell functions. In this paper, we investigate nonlinear dynamics with targeted mutation in cyclically competing species. In different ways to classic approaches of mutation in cyclic games, we assume that mutation may occur in targeted individuals who have been removed from intraspecific competition. By investigating each scenario depending on the number of objects for targeted mutation analytically and numerically, we found that targeted mutation can lead to persistent coexistence of all species. In addition, under the specific condition of targeted mutation, we found that targeted mutation can lead to emergences of bistable states for species survival. Through the linear stability analysis of rate equations, we found that those phenomena are accompanied by Hopf bifurcation which is supercritical. Our findings may provide more global perspectives on understanding underlying mechanisms to control biodiversity in ecological/biological sciences, and evidences with mathematical foundations to resolve social dilemmas such as a turnover of group members by resigning with intragroup conflicts in social sciences.

In the past decade, there have been many efforts to understand the species interplay with biodiversity in cyclic games within the macro and microscopic levels. In this direction, mobility and intraspecific competition have been found to be the main factors promoting coexistence in spatially extended systems. In this paper, we explore the relevant effect of asymmetric competitions coupled with mobility on the coexistence of cyclically competing species. By examining the coexistence probability, we have found that mobility can facilitate coexistence in the limited cases of asymmetric competition and can be well predicted by the basin structure of the deterministic system. In addition, it is found that mobility can have beneficial and harmful effects on coexistence when all competitions occur asymmetrically. We also found that the coexistence in the spatial dynamics ultimately becomes a global attractor. We hope to provide insights into the associated effects of asymmetric interplays on species coexistence in a spatially extended system and understand the biodiversity of asymmetrically competitive species under more complex competition structures.

Conflict that arises between two groups of different paradigms is an inevitable phenomenon, and a representative example of the conflict among different groups is a conflict phenomenon caused by competition among political parties. In this paper, we study the dynamical behavior of a political party system. Considering three major political parties, we investigate how political party systems can be changed by employing a mathematical model. By considering the transfer mechanism of recruitment as well as conflict of competition between political parties, we found that all parties are likely to coexist when both the competition and transfer between the parties are weak, or if either mechanism can occur at a relatively low level. Otherwise, a political party system is changed to a single-party system. In addition, we found that when a party system was changed into a single-party system, it appeared to be either bistable or multistable, and has been elucidate by linear stability analysis. Our results may provide insights to understand mechanisms how political party systems can be changed by conflict and transfer.

A fundamental result in the evolutionary-game paradigm of cyclic competition in spatially extended ecological systems, as represented by the classic Reichenbach-Mobilia-Frey (RMF) model, is that high mobility tends to hamper or even exclude species coexistence. This result was obtained under the hypothesis that individuals move randomly without taking into account the suitability of their local environment. We incorporate local habitat suitability into the RMF model and investigate its effect on coexistence. In particular, we hypothesize the use of "basic instinct" of an individual to determine its movement at any time step. That is, an individual is more likely to move when the local habitat becomes hostile and is no longer favorable for survival and growth. We show that, when such local habitat suitability is taken into account, robust coexistence can emerge even in the high-mobility regime where extinction is certain in the RMF model. A surprising finding is that coexistence is accompanied by the occurrence of substantial empty space in the system. Reexamination of the RMF model confirms the necessity and the important role of empty space in coexistence. Our study implies that adaptation/movements according to local habitat suitability are a fundamental factor to promote species coexistence and, consequently, biodiversity.