Honesty in signalling games is maintained by trade-offs rather than costs.

Our results refute the claim that signals must be costly at the evolutionary equilibrium to be reliable, as predicted by the Handicap Principle and so-called 'costly signalling' theory. Thus, our results raise serious concerns about the handicap paradigm. We argue that the evolution of reliable signalling is better understood within a Darwinian life-history framework, and that the conditions for honest signalling are more clearly stated and understood by evaluating their trade-offs rather than their costs per se. We discuss potential shortcomings of equilibrium models and we provide testable predictions to help advance the field and establish a better explanation for honest signals. Last but not least, our results highlight why signals are expected to be efficient rather than wasteful.