No abstract

Summary1. The habitat heterogeneity (HHH) and individual adjustment (IAH) hypotheses are commonly proposed to explain a decrease in reproduction rate with increasing population density. Higher numbers of low-quality territories with low reproductive success as density increases lead to a decrease in reproduction under the HHH, while more competition at high density decreases reproduction across all territories under the IAH. 2. We analyse the influence of density and habitat heterogeneity on reproductive success in eight populations of long-lived territorial birds of prey belonging to four species. Sufficient reliability in distinguishing between population-wide, site-specific and individual quality effects on reproduction was granted through the minimal duration of 20 years of all data sets and the ability to control for individual quality in five of them. 3. Density increased in five populations but reproduction did not decrease in these. Territory occupancy as a surrogate of territory quality correlated positively with reproductive success but only significantly so in large data sets with more than 100 territories. 4. Reproductive success was always best explained by measures of territory quality in multivariate models. Direct or delayed (t)1) population density entered very few of the best models. Mixed models controlling for individual quality showed an increasing reproductive performance in older individuals and in those laying earlier, but measures of territory quality were also always retained in the best models. 5. We find strong support for the habitat heterogeneity hypothesis but weak support for the individual adjustment hypothesis. Both individual and site characteristics are crucial for reproductive performance in long-lived birds. Proportional occupancy of territories enables recognition of highquality territories as preferential conservation targets.

Density-dependent reproduction is commonly explained by either the habitat heterogeneity (HHH) or individual adjustment (IAH) hypothesis. Under the HHH, high quality territories are assumed to be occupied fi rst. At higher density, occupation of low-quality territories increases due to lower availability of high-quality territories, which reduces mean reproductive success. Alternatively, the IAH assumes that increased competition at higher densities reduces reproductive success in all territories. For birds of prey, HHH plays an important role in territorial species, and IAH in socially breeding species. To test the generality of this hypothesis, we studied the mechanism behind density dependence in raven Corvus corax , a long-lived passerine bird, using long-term population data from a large number of territories.Population density decreased reproduction, which was explained by increased usage of low quality territories at higher density, supporting the HHH. Density reduced reproduction in low quality territories, but not in high and intermediate quality territories. We additionally compared the explanatory power of diff erent models describing brood size, representing IAH, HHH, or a combination of both. Th e best model represented a combination of both hypotheses, in which the eff ect of density depended on territory quality. Our conclusion that both IAH and HHH are supported can be explained by the biology of ravens, where territorial adults not only experience interference competition with other territorial adults, but also with social groups of juveniles and fl oaters. We conclude that the relative importance of IAH and HHH may depend on variation in territory quality and social structure.

Parents may adapt their offspring sex ratio in response to their own phenotype and environmental conditions. The most significant causes for adaptive sex-ratio variation might express themselves as different distributions of fitness components between sexes along a given variable. Several causes for differential sex allocation in raptors with reversed sexual size dimorphism have been suggested. We search for correlates of fledgling sex in an extensive dataset on common buzzards Buteo buteo, a long-lived bird of prey. Larger female offspring could be more resource-demanding and starvation-prone and thus the costly sex. Prominent factors such as brood size and laying date did not predict nestling sex. Nonetheless, lifetime sex ratio (LSR, potentially indicative of individual sex allocation constraints) and overall nestling sex were explained by territory quality with more females being produced in better territories. Additionally, parental plumage morphs and the interaction of morph and prey abundance tended to explain LSR and nestling sex, indicating local adaptation of sex allocation However, in a limited census of nestling mortality, not females but males tended to die more frequently in prey-rich years. Also, although females could have potentially longer reproductive careers, a subset of our data encompassing full individual life histories showed that longevity and lifetime reproductive success were similarly distributed between the sexes. Thus, a basis for adaptive sex allocation in this population remains elusive. Overall, in common buzzards most major determinants of reproductive success appeared to have no effect on sex ratio but sex allocation may be adapted to local conditions in morph-specific patterns.

By 2040, roughly two-thirds of humanity are expected to live in urban areas. As cities expand, humans irreversibly transform natural ecosystems, creating both opportunities and challenges for wildlife. Here, we investigate how the Northern Goshawk ( Accipiter gentilis ) is adjusting to urban environments. We measured a variety of behavioural and ecological parameters in three urban and four rural study sites. City life appeared related to all parameters we measured. Urban female goshawks were overall 21.7 (CI 95% 5.13–130) times more likely to defend their nestlings from humans than rural females. Urban goshawks were 3.64 (CI 95% 2.05–6.66) times more likely to feed on pigeons and had diets exhibiting lower overall species richness and diversity. Urban females laid eggs 12.5 (CI 95% 7.12–17.4) days earlier than rural individuals and were 2.22 (CI 95% 0.984–4.73) times more likely to produce a brood of more than three nestlings. Nonetheless, urban goshawks suffered more from infections with the parasite Trichomonas gallinae , which was the second most common cause of mortality (14.6%), after collisions with windows (33.1%). In conclusion, although city life is associated with significant risks, goshawks appear to thrive in some urban environments, most likely as a result of high local availability of profitable pigeon prey. We conclude that the Northern Goshawk can be classified as an urban exploiter in parts of its distribution.