 The handicap principle has proven hard to test empirically, partly because of inconsistent interpretations of Zahavi ’ s metaphor and Grafen ’ s marginal fitness model, and partly because of conflicting empirical results: in some studies individuals with bigger signals seem to pay higher costs, in other studies they seem to be paying lower costs. A possible explanation for the inconsistent empirical results is given in a series of papers by Getty, who shows that Grafen ’ s proof of the handicap principle is based on the critical simplifying assumption that signallers trade off costs for benefits in an additive fashion, the way humans invest money to increase income in the same currency. But the assumption that costs and benefits trade off in an additive fashion is not valid for the survival cost – reproduction benefit tradeoff that is assumed to mediate the evolution of sexually selected signals. Fitness depends on producing offspring, which is a multiplicative function of reproductive success given an individual is still alive times the probability of still being alive, given investment in signals.
