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    Broom, Mark Johanis, Michal and Rychtář, Jan 2018. The effect of fight cost structure on fighting behaviour involving simultaneous decisions and variable investment levels. Journal of Mathematical Biology, Vol. 76, Issue. 1-2, p. 457.

    Snart, Charles J. P. Kapranas, Apostolos Williams, Huw Barrett, David A. and Hardy, Ian C. W. 2018. Sustenance and Performance: Nutritional Reserves, Longevity, and Contest Outcomes of Fed and Starved Adult Parasitoid Wasps. Frontiers in Ecology and Evolution, Vol. 6, Issue. ,

    Christensen, Charlotte and Radford, Andrew N 2018. Dear enemies or nasty neighbors? Causes and consequences of variation in the responses of group-living species to territorial intrusions. Behavioral Ecology,

    Sykes, David and Rychtář, Jan 2017. Optimal aggression in kleptoparasitic interactions. Involve, a Journal of Mathematics, Vol. 10, Issue. 5, p. 735.

    Takeuchi, Tsuyoshi 2017. Agonistic display or courtship behavior? A review of contests over mating opportunity in butterflies. Journal of Ethology, Vol. 35, Issue. 1, p. 3.

    Palaoro, Alexandre V. and Briffa, Mark 2017. Weaponry and defenses in fighting animals: how allometry can alter predictions from contest theory. Behavioral Ecology, Vol. 28, Issue. 1, p. 328.

    Mesterton-Gibbons, Mike Dai, Yao Goubault, Marlène and Hardy, Ian C. W. 2017. Volatile Chemical Emission as a Weapon of Rearguard Action: A Game-Theoretic Model of Contest Behavior. Bulletin of Mathematical Biology, Vol. 79, Issue. 11, p. 2413.

    Sherratt, Thomas N and Mesterton-Gibbons, Mike 2017. eLS. p. 1.

    Mesterton-Gibbons, Mike Karabiyik, Tugba and Sherratt, Tom N. 2016. On the Evolution of Partial Respect for Ownership. Dynamic Games and Applications, Vol. 6, Issue. 3, p. 359.

    Edmonds, Elizabeth and Briffa, Mark 2016. Weak rappers rock more: hermit crabs assess their own agonistic behaviour. Biology Letters, Vol. 12, Issue. 1, p. 20150884.

    Carvalho, M. R. M. Peixoto, P. E. C. and Benson, W. W. 2016. Territorial clashes in the Neotropical butterfly Actinote pellenea (Acraeinae): do disputes differ when contests get physical?. Behavioral Ecology and Sociobiology, Vol. 70, Issue. 1, p. 199.

    Bishop, Amanda M. Pomeroy, Paddy and Twiss, Sean D. 2015. Variability in individual rates of aggression in wild gray seals: fine-scale analysis reveals importance of social and spatial stability. Behavioral Ecology and Sociobiology, Vol. 69, Issue. 10, p. 1663.

    Whiting, Martin J. Noble, Daniel W.A. and Somaweera, Ruchira 2015. Sexual dimorphism in conspicuousness and ornamentation in the enigmatic leaf-nosed lizardCeratophora tennentiifrom Sri Lanka. Biological Journal of the Linnean Society, Vol. 116, Issue. 3, p. 614.

    Broom, Mark Johanis, Michal and Rychtář, Jan 2015. The effect of fight cost structure on fighting behaviour. Journal of Mathematical Biology, Vol. 71, Issue. 4, p. 979.

    Pietraszewski, David and Shaw, Alex 2015. Not by Strength Alone. Human Nature, Vol. 26, Issue. 1, p. 44.

    Sherratt, T. N. and Mesterton-Gibbons, M. 2015. The evolution of respect for property. Journal of Evolutionary Biology, Vol. 28, Issue. 6, p. 1185.

    Hasegawa, Masaru and Kutsukake, Nobuyuki 2015. Bayesian competitiveness estimation predicts dominance turnover among wild male chimpanzees. Behavioral Ecology and Sociobiology, Vol. 69, Issue. 1, p. 89.

    Candolin, Ulrika and Tukiainen, Iina 2015. The sexual selection paradigm: have we overlooked other mechanisms in the evolution of male ornaments?. Proceedings of the Royal Society B: Biological Sciences, Vol. 282, Issue. 1816, p. 20151987.

    Mesterton-Gibbons, Mike Karabiyik, Tugba and Sherratt, Tom N. 2014. The Iterated Hawk–Dove Game Revisited: The Effect of Ownership Uncertainty on Bourgeois as a Pure Convention. Dynamic Games and Applications, Vol. 4, Issue. 4, p. 407.

    Couchoux, Christelle and van Nouhuys, Saskya 2014. Effects of Intraspecific Competition and Host-Parasitoid Developmental Timing on Foraging Behaviour of a Parasitoid Wasp. Journal of Insect Behavior, Vol. 27, Issue. 3, p. 283.

  • Print publication year: 2013
  • Online publication date: June 2013

2 - Dyadic contests: modelling fights between two individuals



Animal contests were the focal topic that brought game theory to the attention of behavioural ecologists, giving rise to evolutionary game theory. Game theory has remained by far the most popular method of deriving theoretical predictions ever since, although it nowadays coexists with other methods of analysis. Here I review the developments to date and highlight similarities and differences between models. There is a clear progression from simple two-player models with fixed payoffs to explicit tracking of fitness consequences in a population context. In many cases this development has helped to discover that some of the early predictions may have been misleading. Despite the large number of current models, there are still gaps in the theoretical literature: sometimes simplifying assumptions have been relaxed in one context but not another. I hope that by highlighting these gaps theoreticians will be provided with new research ideas, and empiricists will be encouraged not only to distinguish between existing models but to be able to point out assumptions that are essential for deriving a result yet may be violated in existing systems, thus directing new modelling in the most useful direction.


Until the mid 1960s, animal contests were viewed using group selectionist thinking. Julian Huxley (1966) thought that ritualised fights evolved to limit intraspecific damage, partly based on Konrad Lorenz's (1964, 1965) ideas that species need to evolve mechanisms that limit aggression in species that possess dangerous weapons for other reasons, e.g. as adaptations for capturing prey. Following George C. Williams’ (1966) book Adaptation and Natural Selection: A Critique of Some Current Thought, however, biologists became aware of the need to distinguish between explanations that are based on benefits to the individual versus those that rely on benefits accruing to a group (or a species).

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Animal Contests
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