Costs of reproduction: Assessing responses to brood size manipulation on life-history and behavioural traits using multi-state capture-recapture models

Costs of reproduction are fundamental trade-offs shaping the evolution of life histories. There has been much interest, discussion and controversy about the nature and type of reproductive costs. The manipulation of reproductive effort (e.g. brood size manipulation) may alter not only life-history traits such as future adult survival rate and future reproductive effort, but also behavioural decisions affecting recapture/resighting and dispersal probabilities. We argue that many previous studies of the costs of reproduction may have erroneously concluded the existence or non-existence of such costs because of their use of local return rates to assess survival. In this paper, we take advantage of the modern multistate capture-recapture methods to highlight how the accurate assessment of the costs of reproduction requires incorporating not only recapture probability, but also behavioural 'state' variables, for example dispersal status and current reproductive investment. The inclusion of state-dependent decisions can radically alter the conclusions drawn regarding the costs of reproduction on future survival or reproductive investment. We illustrate this point by re-analysing data collected to address the question of the costs of reproduction in the collared flycatcher and the great tit. We discuss in some detail the methodological issues and implications of the analytical techniques.