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Game-theoretic arguments are used to derive two new hypotheses to explain why territorial residents so consistently defeat potential usurpers. Both hypotheses are based on help from established, familiar neighbors. The first hypothesis follows simply from Krebs' (1982) assertion that the value of a territory to a usurper must be decremented by the costs of negotiating dear-enemy relationships with the remaining neighbors. An implication is that the remaining neighbors will also have to pay these renegotiation costs if the usurper succeeds. The first hypothesis is that it may benefit a territorial animal to help its established neighbors defend so it can avoid having to renegotiate territorial boundaries with a new, unfamiliar neighbor. This hypothesis assumes net positive benefits to helping without requiring reciprocation.The second hypothesis requires reciprocation to compensate for immediate net costs of helping. An animal should help its neighbors fight off usurpers only if the neighbors will reciprocate. This hypothesis is based on the prisoner's dilemma game and builds on Axelrod's (1984) work. Cooperative defense (reciprocal help) can be an evolutionarily stable strategy (ESS) if several conditions are met. One critical condition is that the relationship between neighbors is relatively stable. Cooperative defense should help established neighbors retain their territories, and should therefore be a cause, as well as a consequence of stability. It is suggested that the necessary conditions are not very restrictive, that they are often met in nature, and that shared defense is observed but not recognized as such.

Dear Enemies and the Prisoner's Dilemma: Why Should Territorial Neighbors Form Defensive Coalitions?