Effects of experimentally sustained elevated testosterone on incubation behaviour and reproductive success in female great tits (Parus major).

In many seasonally breeding birds, female and male testosterone (T) levels peak at the start of the breeding season, coinciding with pair bonding and nesting activities. Shortly after the onset of egg laying, T levels slowly decline to baseline levels in both sexes, but more rapidly so in females. During this period, T in males may still function to facilitate territorial behaviour, mate guarding and extra pair copulations, either via short lasting peaks or elevated basal levels of the hormone. In some species, however, males become insensitive to increased T after the onset of egg laying. It has been postulated that in these species bi-parental care is essential for offspring survival, as T is known to inhibit paternal care. However, only very few studies have analysed this for females. As females are heavily involved in parental care, they too might become insensitive to T after egg laying. Alternatively, because territorial defence, mate guarding and extra pair copulations are expected to be less important for females than for males, they may not have had the need to evolve a mechanism to become insensitive to T during the period of maternal care, because their natural T levels are never elevated during this part of the breeding season anyway. We tested these alternative hypotheses in female great tits (Parus major). Male great tits have previously been shown to be insensitive to T after egg laying with regard to nestling feeding behaviour (but not song rate). When females had started nest building, we experimentally elevated their T levels up to the nestling feeding phase, and measured incubation behaviour (only females incubate) and reproductive success. T did not significantly affect nest building or egg laying behaviour, although egg laying tended to be delayed in T females. Females with experimentally enhanced T maintained lower temperature during incubation but did not spend less time incubating. This might explain the reduced hatching success of their eggs, smaller brood size and lower number of fledglings we found in this study. As in this species T-dependent behaviour by females during the phase of parental care is not needed, the results support the hypothesis that in this species the need for selection in favour of T-insensitivity did not occur.

Bioaccumulation of heavy metals in eggshell and egg content of seabirds: Lesser (Thalasseus bengalensis) and Greater Crested Tern (Thalasseus bergii).

The Persian Gulf, the most important marine ecosystem, and its islands are negatively impacted by development and pollution. Different types of pollution, especially heavy metals in these habitats, may create risks to birds, influencing reproductive performance, survival, and behaviors. Heavy metals (Cd, Pb, Cr) were analyzed in two long-lived and top food web-level sea birds, Lesser (Thalasseus bengalensis) and Greater Crested Tern (Thalasseus bergii). The research was carried out during 2018 on Nakhiloo Island, located in Dayyer-Nakhiloo Marine National Park, northeastern Persian Gulf, Iran. The nests were selected randomly, and 40 eggs of colonially-nesting terns (20 eggs T. bengalensis and 20 eggs T. bergii; a single egg per nest) were collected. The concentration of heavy metals in eggshells and egg contents was studied to investigate whether bioaccumulation varies between the eggshells and egg contents and also varies in both species. There were significant differences in the amounts of Cd between eggshell and egg content in the Greater and Lesser Crested Tern. This difference was not significant in the concentrations of Pb between the eggshell and egg content in the Tern species. Concentration of Cr did not appear to accumulate in the egg content and was below detection limit. There were significant differences in the concentrations of Cd in the eggshells and egg content between the Greater and Lesser Crested Tern. The results serve to improve understanding and to evaluate the dynamics of heavy metals in eggshells and egg content in various avian species.

The effect of ambient temperature, habitat quality and individual age on incubation behaviour and incubation feeding in a socially monogamous songbird.

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 STATEMENT: Parents 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.