The adaptive significance of brood size and time of breeding in the dipper Cinclus cinclus (Aves: Passeriformes) as seen from post-fledging survival

We assessed the post-fledging survival of dippers Cinclus cinclus from 743 broods in relation to brood size, time of hatching and territory quality. We paid particular attention to assessing whether contrasting breeding performance along unproductive (i.e. acidic) and productive (i.e. circumneutral) rivers represented strategies which optimized the number of surviving young.
For all brood sizes, post-fledging survival varied significantly through the breeding season, with most survivors coming from attempts in the peak period of hatching. After correcting for these seasonal effects, the most common brood size overall, of four, was also the most productive as seen from post-fledging survival; differences in the frequency of occurrence and survival between broods of four and five were marginal. Moreover, a change in the modal brood size from five to four occurred as the season progressed. consistent with a shift in brood productivity.
Broods at acidic sites were significantly smaller than at circumneutral sites; while brood size four was the most productive at both types of site, brood size three was the second most productive at acidic sites, while brood size five was the second most productive at circumneutral sites. Dippers at acidic sites bred significantly later than at circumneutral sites, but post-fledging survival declined most rapidly through the season at the former.
These survival data provide evidence from both seasonal and spatial patterns that brood sizes in the dipper may be optimized in ways consistent with the enhancement of productivity. By contrast, delayed breeding at acidic sites contrasted with the patterns expected from optimization, instead reflecting resource scarcity.