The regulation of brood reduction in Booted Eagles Hieraaetus pennatus through habitat heterogeneity

Brood reduction, the death of one or more chicks through siblicide or starvation, can occur through density‐dependence in fecundity. Brood reduction may arise in territorial breeding systems either as a response to a high level of territorial interference in a situation of high density or as a result of habitat heterogeneity. To test the predictions of the two main hypotheses that attempt to explain how density‐dependent fecundity is generated, the Habitat Heterogeneity Hypothesis (HHH) and the Individual Adjustment Hypothesis (IAH), we analysed the relationship between density and fecundity in an expanding population of Booted Eagles in Doñana National Park, Spain, using an 18‐year data series. We also studied the occurrence and frequency of brood reduction in the same Booted Eagle population to appreciate further its effects and the factors that influence its occurrence and frequency. Our results support the HHH in the present situation of high density, as fecundity in the better territories (older and more frequently occupied) was higher than in low quality territories and was not affected by population density in high density periods. Nevertheless, the fecundity of high quality territories was affected (although not significantly) by population density in periods of low density, suggesting that the IAH was supported when only high quality territories were occupied. Older territories were used more frequently and chicks in these areas hatched earlier and suffered lower mortality than in new territories. We found a significant negative relationship between mean fecundity and its skewness, a finding that also supports HHH. During years of food shortage, less frequently occupied territories suffered higher rates of brood reduction. Brood reduction in this Booted Eagle population was a consequence of the heterogeneous structure of the habitat, with some territories having a higher probability of brood reduction than others. Parental nutritional condition did not affect brood reduction. The effect of brood reduction on nestling quality and population dynamics is also discussed.