Nests of cavity‐nesting birds usually harbor some species of haematophagous ectoparasites that feed on the incubating adults and nestlings. Given the negative impact of ectoparasites on nestlings there will be selection on hosts to reduce parasite infestations through behavioural means. We have experimentally reduced the abundance of all ectoparasites in nests of pied flycatchers Ficedula hypoleuca to explore both whether there are changes in the frequency and duration of putative anti‐parasite behaviours by tending adults, as well as whether such anti‐parasite behaviours are able to compensate for the deleterious effects that parasites may have on nestlings. Heat treatment of nests substantially decreased the density of ectoparasites, and thereby positively affected nestling growth. The frequency and intensity of female grooming and nest sanitation behaviours during the incubation and nestling periods decreased as a consequence of the experimental reduction of ectoparasite infestation. Although nestlings begged more intensely in infested nests, the experiment had no significant effect on parental provisioning effort. Reduction of parasites resulted in larger nestlings shortly before fledging and increased fledging success. This study shows a clear effect of a complete natural nest ectoparasite fauna on parental behaviour at the nest and nestling growth in a cavity‐nesting bird. Although ectoparasites induce anti‐parasite behaviours in females, these behaviours are not able to fully remove parasite's deleterious effects on nestling growth and survival.

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ABSTRACT. The 'female nutrition' hypothesis proposes that food provided by males during incubation is an important energy source for females in bird species in which females alone incubate. Females should be able to communicate their needs through begging signals to mates and males may compensate for the energetic limitations of females through their feeding visits, owing to their overlapping reproductive interests. To test whether female begging during incubation is an honest signal of energetic need and whether mates respond to it we experimentally handicapped female pied flycatchers at the beginning of incubation by clipping two primary flight feathers on each wing. Experimental manipulation led females to intensify begging displays arising from condition impairment and males accordingly increased their incubation feeding rates.Female begging intensity explained more than half of the variation in male incubation feeding rate, thereby showing that female nutrition is the main factor explaining male incubation feeding. Moreover, handicapped females consumed a higher proportion of male food deliveries during the first few days after hatching and weighed less at the end of the nestling period than control females.

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Nesting holes are a scarce resource for obligated cavity-nesting birds and an important selective force for the evolution of aggressive female behaviours, which may be mediated by testosterone (T) levels. It is known that during periods of intense intrasexual competition such as initial breeding stages, females are highly aggressive towards intruding females. Here, we studied the implications of T levels for female-female competition by comparing levels of aggressiveness towards simulated female intruders (decoys) in two populations of the pied flycatcher (Ficedula hypoleuca) with a marked difference in breeding density. To this end, we exposed free-living females to simulated territorial intrusions during 30 min when nest construction was almost complete. T levels of females were measured at the beginning of incubation under the assumption that they are positively associated with T levels during nest building. We also related aggressiveness to T levels in both populations. Furthermore, we aimed at detecting whether variation of T levels may explain female incubation attendance. Females showed higher T levels in the populations where pied flycatchers were exposed to a higher likelihood of conspecific interactions (high breeding density) than in the population with low breeding density. Female territorial presence, vigilance at the nest box and proximity to decoys were negatively related to circulating T levels in the high-density population, but not in the low-density population. Differences in T levels between populations did not result in differences in female incubation attendance, but T levels were negatively related to the incubation attendance in females from the population showing high T levels. In our populations, T levels in females prior to laying reflect the need to defend nesting cavities which is higher at high breeding density and in subdominant females. High T levels are costly in terms of incubation attendance.Ethology 121 (2015) 946-957