Abstract

Incubation is an important aspect of avian life history. The behaviour is energetically costly, and investment in incubation strategies within species, like female nest attentiveness and the feeding by the non-incubating partner during incubation, can therefore vary depending on environmental and individual characteristics. However, little is known about the combined effect of these characteristics. We investigated the importance of ambient temperature, habitat quality, and bird age on female incubation behaviour and male feeding of the incubating female (incubation feeding) in blue tits Cyanistes caeruleus, a socially monogamous songbird. An increase in ambient temperature resulted in a higher nest temperature, and this enabled females to increase the time off the nest for self-maintenance activities. Probably as a consequence of this, an increase in ambient temperature was associated with fewer incubation feedings by the male. Moreover, in areas with more food available (more deciduous trees), females had shorter incubation recesses and males fed females less often. Additionally, males fed young females more, presumably to increase such females’ investment in their eggs, which were colder on average (despite the length of recesses and female nest attentiveness being independent of female age). Male age did not affect incubation feeding rate. In conclusion, the patterns of incubation behaviour were related to both environmental and individual characteristics, and male incubation feeding was adjusted to females’ need for food according these characteristics, which can facilitate new insights to the study of avian incubation energetics.Significance statementParents often invest a substantial amount of energy in raising offspring. How much they do so depends on several environmental factors and on the extent they cooperate to raise the offspring. In birds, males can feed incubating females, which may allow females to stay longer on the nest, which, in turn, may ultimately improve reproductive success. The interplay between environmental factors and such incubation feeding on incubation attendance has, however, received little attention. Here, we show that favourable circumstances (higher ambient temperature and food availability) allowed incubating blue tit females to increase the time off the nest to improve self-maintenance and males to feed them less, whereas males also fed inexperienced partners more often. Thus, we show a concerted effect of several environmental and intrinsic factors on parental effort during incubation, which will help to improve the general understanding of avian incubation and parental care.

Highlights

  • Understanding why species differ in patterns of parental care is important in evolutionary ecology studies (Clutton-Brock 1991)

  • We investigate the simultaneous effect of these factors on different measures of female incubation behaviour and male incubation feeding in blue tits Cyanistes caeruleus, a socially monogamous songbird with female-only incubation and male incubation feeding behaviour

  • Estimate ± SE t value p value Estimate ± SE t value p value Estimate ± SE t value p value (Intercept) Ambient temperature (°C) Proportion deciduous trees Female age Estimated effect sizes of each term (Estimate) with associated standard errors, t and p values were presented based on the minimal adequate model in italics, and parameters of non-significant predictors were obtained immediately before they were removed from the model a Among 92 nest-boxes with temperature logger data, the age of females was unknown in four nest-boxes and these four were not included

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Summary

Introduction

Understanding why species differ in patterns of parental care is important in evolutionary ecology studies (Clutton-Brock 1991). Incubation behaviour is an important aspect of avian parental care. Incubating females usually face a trade-off between spending time on the nest to incubate the eggs to promote embryo development and time off the nest for investment in selfmaintenance (Ardia et al 2009). The incubation rhythm consists of alternated periods, where females alternate leaving the clutch to obtain food (off-bouts) and returning to re-warm the eggs (onbouts; Boulton et al 2010). The incubation rhythm may affect females’ condition and the egg temperature and, the embryo development (Lyon and Montgomerie 1987; Williams 1991)

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