Signalling among relatives. I. Is costly signalling too costly?

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Abstract
Zahavi's handicap principle,originally proposed as an explanation for sexual selection ofelaborate male traits, suggests that a sufficient cost to dishonest signals can outweigh the rewards of deception and allow individuals to communicate honestly. Maynard Smith (1991) and Johnstone and Grafen (1992) introduce the Sir Philip Sidney game in order to extend the handicap principle to interactions among related individuals, and to demonstrate that stable costly signalling systems can exist among relatives. In this paper we demonstrate that despite the benefits associated with honest information transfer, the costs incurred in a stable costly signalling system may leave all participants worse off than they would be in a system with no signalling at all. In both the discrete and continuous forms of the Sir Philip Sidney game, there exist conditions under which costly signalling among relatives, while stable, is so costly that it is disadvantageous compared with no signalling at all. We determine the factors which dictate signal cost and signal benefit in a generalized version of this game, and explain how signal cost can exceed signal value. Such results raise concerns about theevolutionary pathways which could have led to the existence of signalling equilibria in nature. The paper stresses the importance of comparing signalling equilibria with other possible strategies, beforedrawing conclusions regarding the optimality of signalling.