Chimpanzees and the mathematics of battle.

Recent experiments have demonstrated the importance of numerical assessment in animal contests. Nevertheless, few attempts have been made to model explicitly the relationship between the relative number of combatants on each side and the costs and benefits of entering a contest. One framework that may be especially suitable for making such explicit predictions is Lanchester's theory of combat, which has proved useful for understanding combat strategies in humans and several species of ants. We show, with data from a recent series of playback experiments, that a model derived from Lanchester's 'square law' predicts willingness to enter intergroup contests in wild chimpanzees (Pan troglodytes). Furthermore, the model predicts that, in contests with multiple individuals on each side, chimpanzees in this population should be willing to enter a contest only if they outnumber the opposing side by a factor of 1.5. We evaluate these results for intergroup encounters in chimpanzees and also discuss potential applications of Lanchester's square and linear laws for understanding combat strategies in other species.