Territoriality is widespread in the animal kingdom, but its evolutionary origins are less understood than the movement patterns that generate it. Previous studies attempted to understand the evolution of these movement patterns by assuming that certain movement patterns comprised competing strategies and using evolutionary game theory to find which strategies would provide greater fitness and give rise to evolutionary stability. Four strategies were identified: Cautious, an individual that moves through the landscape avoiding all places where it experienced conflicts with other animals; Common-sense, an individual that avoids areas where it lost conflicts and selects areas where it won conflicts; Paradoxical, an individual that does the opposite of Common-sense; and Daring, an individual that seeks areas where it won conflicts, but is unaffected by loss. Previous models used a combination of individual-based models and algebraic calculations, but they did not incorporate recent developments in mechanistic models. Here, we developed a stochastic individual-based model using fluid dynamics to represent space use and used invasion tests to determine evolutionary stability. Using this model, we tested which (if any) of the previous strategies would be evolutionarily stable in two scenarios: one with inconclusive territorial fights and another without. We found that the cautious strategy was evolutionarily stable, in disagreement with previous literature. Cautious individuals show a more homogeneous use of space, when compared to the other three strategies. Cautious individuals tend to win most resources across the environment, which results in greater fitness than other strategies and allows it to invade populations comprised of any of the others. Daring also invades paradox and common-sense strategies, while paradox invades common sense. There was no qualitative difference between scenarios. We discuss how the evolutionary stability of this strategy depends on whether individuals employ their maximal effort when foraging.

地域性在动物界很普遍，但是其进化起源远不及产生它的运动方式。先前的研究试图通过假设某些运动模式包含竞争策略并使用进化博弈论来发现哪些策略将提供更大的适应性并产生进化稳定性，从而理解这些运动模式的演化。确定了四个策略：谨慎，一个人在景观中穿行，避免与其他动物发生冲突的所有地方；常识，一个避免回避冲突的地区并选择赢得冲突的地区的人；矛盾的，与常识相反的个体；大胆，一个寻求赢得冲突的地区的人，但不受损失的影响。先前的模型结合了基于个人的模型和代数计算，但是没有将机械模型的最新发展纳入考虑范围。在这里，我们开发了一个基于随机个体的模型，使用流体动力学来表示空间的使用，并使用入侵测试来确定进化的稳定性。使用此模型，我们测试了在两种情况下哪一种（如果有的话）先前的策略在进化上是稳定的：一种是无确定性的领土斗争，另一种则没有。我们发现，谨慎策略在进化上是稳定的，与先前的文献不同。与其他三种策略相比，谨慎的个人显示出更统一的空间使用方式。谨慎的人往往会赢得整个环境中的大部分资源，与其他策略相比，它的适应性更高，并且可以入侵其他策略组成的人群。大胆还侵犯了悖论和常识策略，而悖论则侵犯了常识。场景之间没有质的差异。我们讨论了该策略的进化稳定性如何取决于个体在觅食时是否尽最大努力。