Determinants of brood defence in the great tit Parus major L.

Summary Great tits (Parus major) tending nestlings reacted defensively to a live predator (Glaucidium perlatum; domestic cat) and the playback of a mixed species mobbing chorus, or to the latter alone. Defensive behaviour, mainly mobbing, reflected the risk taken and is assessed by five measures. Multivariate and contingency analyses revealed that at least 11 of 16 contextual independent variables affected the risk taken. Incremental effects are due to: Age of young, sex of the defending bird, the expected number of neighbouring mobbers, low temperature, wet canopy, the raptor's distance from cover, coniferous forest, advancing season. A decremental effect is exerted by a large brood that is older. Annual differences in defence arise probably from demographic factors such as fecundity, which in turn affect the parent's benefit-cost ratio (number of young of the same sex as the parent/residual reproductive value of the parent).While the effects of annual fecundity, age of young and season were predicted on the basis of this benefit-cost ratio, the failure to verify an incremental effect of brood size runs counter to established theory. We conclude that parents gear their defence efforts to energy investment, past or future, and are mal-adapted to brood size as a promotor of risk taken. The influence of the habitat is poorly understood. At least three factors (age and number of young, parent's sex) act additively on part of the response. Despite the large number of variables examined, about 43% of the total response variance remains unexplained.While four defence measures are determined by at least 10 contextual factors, a fifth measure, the male's minimum distance from the raptor, is determined by one other factor, the appearance of the male. The latter leads us to assume an additional, social rôle of brood defence.Risk-assessment by great tits leading to risk-aversive defence behaviour is governed by evolved restraints rather than by momentary constraints. Examples are provided by the effects of weather and cover.