Abstract
In most evolutionary matrix game models, the payoffs are assumed to be constant. Here, we develop a toy model of hunting in pairs in which interaction outcomes are determined not only by the behavioural strategy choice of the players but also a second, evolvable physical trait of them. We analyse the evolutionary dynamics of this model in a limiting case of the classical monomorphic framework introduced by Maynard Smith and Price characterised by rare mutations in physical trait, moderately frequent mutations in behaviour, and very fast selection in both. We show that this process can transform the evolutionarily stable strategy and, as a result, even the nature of the game played in the population: We present an illustrative numerical example in which the concurrent evolution of the behavioural and trait components of the phenotype turns a social dilemma into a harmony game via an intermediate phase of coexistence.
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References
Traulsen, A. & Glynatsi, N. E. The future of theoretical evolutionary game theory. Phil. Trans. R. Soc. B 378, 20210508 (2023).
von Neumann, J. & Morgenstern, O. Theory of Games and Economic Behaviour (Princeton University Press, Princeton, NJ, 1944).
Nash, J. F. Jr. Equilibrium points in n-person games. Proc. Natl. Acad. Sci. USA 36, 48–49 (1950).
Axelrod, R. The Evolution of Cooperation. (Basic Books, New York, NY, 1984).
Poundstone, W. Prisoner’s Dilemma. (Anchor Books, New York, NY, 1993).
Maynard Smith, J. & Price, G. R. The logic of animal conflict. Nature 246, 15–18 (1973).
Nowak, M. A. Five rules for the evolution of cooperation. Science 314, 1560–1563 (2006).
Szabó, G. & Fáth, G. Evolutionary games on graphs. Phys. Rep. 446, 97–216 (2007).
Perc, M. & Szolnoki, A. Coevolutionary games – a mini review. BioSystems 99, 109–125 (2010).
Pires, D. L., Erovenko, I. V. & Broom, M. Network topology and movement cost, not updating mechanism, determine the evolution of cooperation in mobile structured populations. PLoS ONE 18, e0289366 (2023).
Erovenko, I. V. & Broom, M. The evolution of cooperation in a mobile population on random networks: Network topolgy matters only for low-degree networks. Dyn. Games Appl. (2024).
Trivers, R. L. The evolution of reciprocal altruism. Q. Rev. Biol. 46, 35–57 (1971).
Nowak, M. A. & Sigmund, K. The evolution of stochastic strategies in the prisoner’s dilemma. Acta Appl. Math. 20, 247–265 (1990).
Garay, J. & Móri, T. F. Best reply player against mixed evolutionarily stable strategy user. Bull. Math. Biol. 84, 23 (2022).
Huang, F., Cao, M. & Wang, L. Learning enables adaptation in cooperation for multi-player stochastic games. J. R. Soc. Interface 17, 20200639 (2020).
Huang, F., Cao, M. & Wang, L. Approximate policy iteration for robust stochastic control of multiagent Markov decision processes. IEEE Trans. Autom. Control 70, 3587–3602 (2025).
Szabó, G. & Hauert, C. Evolutionary prisoner’s dilemma games with voluntary participation. Phys. Rev. E 66, 062903 (2002).
Dreber, A., Rand, D. G., Fudenberg, D. & Nowak, M. A. Winners don’t punish. Nature 452, 348–351 (2008).
Perc, M. et al. Statistical physics of human cooperation. Phys. Rep. 687, 1–51 (2017).
Gokhale, C. S. & Hauert, C. Eco-evolutionary dynamics of social dilemmas. Theor. Popul. Biol. 111, 28–42 (2016).
Argasinski, K. & Broom, M. Evolutionary stability under limited population growth: Eco-evolutionary feedbacks and replicator dynamics. Ecol. Complex. 18, 198–212 (2018).
Croll, J. C., Egas, M. & Smallegange, I. M. An eco-evolutionary feedback loop between population dynamics and fighter expression affects the evolution of alternative reproductive tactics. J. Anim. Ecol. 88, 11–23 (2019).
Liu, L., Chen, X. & Szolnoki, A. Coevolutionary dynamics via adaptive feedback in collective-risk social dilemmas. eLife 12, e82954 (2023).
Sigmund, K. & Holt, R. D. Toward ecoevolutionary dynamics. Proc. Natl. Acad. Sci. USA 118, e2100200118 (2021).
Zhang, F. & Hui, C. Eco-evolutionary feedback and the invasion of cooperation in prisoner’s dilemma games. PLoS One 6, e27523 (2011).
Wang, X. & Fu, F. Eco-evolutionary dynamics with environmental feedback: Cooperation in a changing world. EPL 132, 10001 (2020).
Tilman, A. R., Plotkin, J. B. & Akçay, E. Evolutionary games with environmental feedbacks. Nat. Commun. 11, 915 (2020).
Weitz, J. S., Eksin, C., Paarporn, K., Brown, S. P. & Ratcliff, W. C. An oscillating tragedy of the commons in replicator dynamics with game-environment feedback. Proc. Natl. Acad. Sci. U.S.A. 113, E7518–E7525 (2016).
Taylor, C. & Nowak, M. A. Evolutionary game dynamics with non-uniform interaction rates. Theor. Popul. Biol. 69, 243–252 (2006).
Quan, J. & Wang, X.-J. Some analytical properties of the model for stochastic evolutionary games in finite populations with non-uniform interaction rates. Commun. Theor. Phys. 60, 37–47 (2013).
Mei, J., Tao, Y., Li, C. & Zheng, X.-D. Evolutionary game dynamics with non-uniform interaction rates in finite population. J. Theor. Biol. 540, 111086 (2022).
Křivan, V. & Cressman, R. Interaction times change evolutionary outcomes: Two-player matrix games. J. Theor. Biol. 416, 199–207 (2017).
Garay, J., Csiszár, V. & Móri, T. F. Evolutionary stability for matrix games under time constraints. J. Theor. Biol. 415, 1–12 (2017).
Akçay, E. & Roughgarden, J. The evolution of payoff matrices: providing incentives to cooperate. Proc. R. Soc. B 278, 2198–2206 (2011).
Sadekar, O., Civilini, A., Gómez-Gardeñes, J., Latora, V. & Battiston, F. Evolutionary game selection creates cooperative environments. Phys. Rev. E 110, 014306 (2024).
Venkataraman, P., Mahilkar, A., Raj, N. & Saini, S. Empirical evidene of resource dependent evolution of payoff matrices in saccharomyces cerevisiae populations. J. Evol. Biol. 38, 122–128 (2024).
Király, B., Varga, T., Szabó, G. & Garay, J. Evolutionarily stable payoff matrix in hawk-dove games. BMC Ecol. Evol. 24, 65 (2024).
Yamaguchi, N., Cooper, A., Werdelin, L. & Macdonald, D. W. Evolution of the mane and group-living in the lion ( panthera leo: a review. J. Zool. 263, 329–342 (2004).
Schaller, G. B. The Serengeti Lion (University of Chicago Press, Chicago, IL, 1972).
Packer, C. & Pusey, A. E. Divided we fall: Cooperation among lions. Sci. Am. 276, 32–39 (1997).
Packer, C., Scheel, D. & Pusey, A. E. Why lions form groups: Food is not enough. Am. Nat. 136, 1–19 (1990).
Stander, P. E. Cooperative hunting in lions: The role of the individual. Behav. Ecol. Sociobiol. 29, 445–454 (1992).
Sunquist, M. & Sunquist, F. Wild Cats of the World (University of Chicago Press, Chicago, IL, 2002).
Fisher, D. O. & Dickman, C. R. Body size-prey relationships in insectivorous marsupials: Tests of three hypotheses. Ecology 74, 1871–1883 (1993).
Murphy, K. J. et al. Improving biological realism of predator-prey size relationships in food web models alters ecosystem dynamics. Biol. Lett. 19, 20230142 (2023).
Stander, P. E. Foraging dynamics of lions in a semi-arid environment. Can. J. Zool. 70, 8–21 (1992).
Hamilton, W. D. Geometry for the selfish herd. J. Theor. Biol. 31, 295–311 (1971).
Cressman, R. & Garay, J. The effects of opportunistic and intentional predators on the herding behavior of prey. Ecology 92, 432–440 (2011).
Bishop, T. & Cannings, C. A generalized war of attrition. J. Theor. Biol. 70, 85–124 (1978).
Haldane, J. B. S. A mathematical theory of natural and artificial selection. part i. Trans. Camb. Phil. Soc. 23, 19–41 (1924).
Garay, J., Csiszár, V. & Móri, T. F. Subsistence of sib altruism in different mating systems and Haldane’s arithmetic. J. Theor. Biol. 557, 111330 (2023).
Garay, J., López, I., Varga, Z., Csiszár, V. & Móri, T. F. Survival cost of sharing among altruistic full siblings in Mendelian population. BMC Ecol. Evol. 24, 142 (2024).
Garay, J., Varga, T., Csiszár, V., Móri, T. F. & Szilágyi, A. Matrix game between full siblings in Mendelian populations. PLoS ONE 20, e0331044 (2025).
Acknowledgements
T.V. discloses the following: This work was supported by the Hungarian National Research, Development and Innovation Office NKFIH, Hungary KKP 129877. The study was funded by the National Research, Development and Innovation Office in Hungary (RRF-2.3.1-21-2022-00006). This work was supported by project TKP2021-NVA-09. Project no TKP2021-NVA-09 has been implemented with the support provided by the Ministry of Culture and Innovation of Hungary from the National Research, Development and Innovation Fund, financed under the TKP2021-NVA funding scheme.
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J.G. and B.K. conceptualized and designed the study. B.K. and T.V. carried out the necessary investigations and analyses. B.K. prepared the figures. All authors contributed to the writing of, read, and approved the final manuscript.
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Király, B., Varga, T. & Garay, J. Evolutionary path from a prisoner’s dilemma to a harmony game via hawk–dove games.
Sci Rep (2026). https://doi.org/10.1038/s41598-026-47565-9
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DOI: https://doi.org/10.1038/s41598-026-47565-9
Keywords
- Matrix game
- Trait evolution
- Payoff evolution
- Social dilemma
- Evolutionarily stable strategy transition
Source: Ecology - nature.com
