Miloš Krist

Egg size and offspring quality: a meta‐analysis in birds

Parents affect offspring fitness by propagule size and quality, selection of oviposition site, quality of incubation, feeding of dependent young, and their defence against predators and parasites. Despite many case studies on each of these topics, this knowledge has not been rigorously integrated into individual parental care traits for any taxon. Consequently, we lack a comprehensive, quantitative assessment of how parental care modifies offspring phenotypes. This meta‐analysis of 283 studies with 1805 correlations between egg size and offspring quality in birds is intended to fill this gap. The large sample size enabled testing of how the magnitude of the relationship between egg size and offspring quality depends on a number of variables. Egg size was positively related to nearly all studied offspring traits across all stages of the offspring life cycle. Not surprisingly, the relationship was strongest at hatching but persisted until the post‐fledging stage. Morphological traits were the most closely related to egg size but significant relationships were also found with hatching success, chick survival, and growth rate. Non‐significant effect sizes were found for egg fertility, chick immunity, behaviour, and life‐history or sexual traits. Effect size did not depend on whether chicks were raised by their natural parents or were cross‐fostered to other territories. Effect size did not depend on species‐specific traits such as developmental mode, clutch size, and relative size of the egg, but was larger if tested in captive compared to wild populations and between rather than within broods. In sum, published studies support the view that egg size affects juvenile survival. There are very few studies that tested the relationship between egg size and the fecundity component of offspring fitness, and no studies on offspring survival as adults or on global fitness. More data are also needed for the relationships between egg size and offspring behavioural and physiological traits. It remains to be established whether the relationship between egg size and offspring performance depends on the quality of the offspring environment. Positive effect sizes found in this study are likely to be driven by a causal effect of egg size on offspring quality. However, more studies that control for potential confounding effects of parental post‐hatching care, genes, and egg composition are needed to establish firmly this causal link.

Are blue eggs a sexually selected signal of female collared flycatchers? A cross-fostering experiment

Impressive variation in egg colouration among birds has puzzled evolutionary biologists for a long time. The most frequently studied selective forces moulding egg colouration—predation and brood parasitism—have either received little empirical support or may play a role in only a minority of species. A novel hypothesis has suggested that egg colour may be significantly influenced by sexual selection. Females may deposit a blue-green pigment biliverdin into eggshells instead of using it for themselves as a powerful antioxidant. By this handicap, females may signal their quality to males, which are then hypothesized to increase their paternal effort. We experimentally tested the hypothesis in the collared flycatcher (Ficedula albicollis), a species laying blue-green eggs. We cross-fostered clutches between nests to disentangle effects of female/territory quality and egg colour on paternal effort and nestling quality. The results supported two assumptions of sexual signalling through egg colour hypothesis: Blue pigment seems to be a limited resource for females, and female quality is positively correlated with the intensity of the blue-green colour. However, we did not find support for the main prediction of the hypothesis, as male parental effort parameters (feeding frequencies to nestlings and intensity of nest defence) were unrelated to egg colour. We discuss possible reasons for the discrepancy between our results and previous correlative analyses that supported the hypothesis that blue egg colour may be a postmating, sexually selected signal in females.

Egg size and offspring performance in the collared flycatcher (Ficedula albicollis): a within-clutch approach

Adaptive within-clutch allocation of resources by laying females is an important focus of evolutionary studies. However, the critical assumption of these studies, namely that within-clutch egg-size deviations affect offspring performance, has been properly tested only rarely. In this study, we investigated effects of within-clutch deviations in egg size on nestling survival, weight, fledgling condition, structural size and offspring recruitment to the breeding population in the collared flycatcher (Ficedula albicollis). Besides egg-size effects, we also followed effects of hatching asynchrony, laying sequence, offspring sex and paternity. There was no influence of egg size on nestling survival, tarsus length, condition or recruitment. Initially significant effect on nestling mass disappeared as nestlings approached fledging. Thus, there seems to be limited potential for a laying female to exploit within-clutch egg-size variation adaptively in the collared flycatcher, which agrees with the majority of earlier studies on other bird species. Instead, we suggest that within-clutch egg-size variation originates from the effects of proximate constraints on laying females. If true, adaptive explanations for within-clutch patterns in egg size should be invoked with caution.

Barrier crossing in small avian migrants: individual tracking reveals prolonged nocturnal flights into the day as a common migratory strategy

Over decades it has been unclear how individual migratory songbirds cross large ecological barriers such as seas or deserts. By deploying light-level geolocators on four songbird species weighing only about 12 g, we found that these otherwise mainly nocturnal migrants seem to regularly extend their nocturnal flights into the day when crossing the Sahara Desert and the Mediterranean Sea. The proportion of the proposed diurnally flying birds gradually declined over the day with similar landing patterns in autumn and spring. The prolonged flights were slightly more frequent in spring than in autumn, suggesting tighter migratory schedules when returning to breeding sites. Often we found several patterns for barrier crossing for the same individual in autumn compared to the spring journey. As only a small proportion of the birds flew strictly during the night and even some individuals might have flown non-stop, we suggest that prolonged endurance flights are not an exception even in small migratory species. We emphasise an individual’s ability to perform both diurnal and nocturnal migration when facing the challenge of crossing a large ecological barrier to successfully complete a migratory journey.

Paternity covaries with laying and hatching order in the collared flycatcher Ficedula albicollis

Females in most bird species engage in extra-pair copulations. Although this behaviour is widespread, benefits for females of doing so are less understood. The most widely accepted hypothesis is that they improve their previous choice of social partner and gain genetic benefits for their offspring. Some evidence for this comes from studies that find that extra-pair young (EPY) have greater fitness than their half-sibs. However, this might be also caused by maternal, non-genetic effect, a possibility that remains largely untested. Here we test whether EPY are laid in larger eggs or eggs laid early in the laying sequence in the collared flycatcher (Ficedula albicollis). The size of eggs bearing EPY and within-pair young (WPY) did not differ, however, EPY were laid in early eggs and consequently hatched earlier than WPY. As hatching asynchrony is a strong determinant of offspring size and survival in many species, including collared flycatcher, our results suggest that a caution is needed when paternal genetic effects are to be inferred from comparison of naturally occurring half-sibs.

Short- and long-term effects of egg size and feeding frequency on offspring quality in the collared flycatcher (Ficedula albicollis)

1. Despite the central importance for life-history theory, egg-size effects on offspring fitness are still considered ambiguous. Most previous studies were only observational and consequently might suffer from uncontrolled correlations between egg size and parental/territory quality. Even after cross-fostering is performed, direct genetic effects and parental adjustment of post-natal care might confound our estimates of egg-size effects per se. 2. I performed a full cross-fostering experiment in the collared flycatcher (Ficedula albicollis) exchanging the whole clutches between pairs of nests. I statistically controlled for direct genetic effects and parental feeding frequencies. I followed young until recruitment to estimate the long-term effects of egg size and parental provisioning. In addition, I compared the effects obtained in the cross-fostering experiment with those obtained from a set of unmanipulated nests. 3. Egg size per se affected offspring morphology in both the short and long term, while having no effect on offspring survival and immunity. Egg-size effects were not confounded by parental post-natal care and direct genetic effects. 4. The number of care-givers was an influential predictor of nestling performance. Apart from the variation caused by this factor, feeding frequencies had no consistent effect on offspring performance. 5. Fitness benefits of large eggs may be difficult to establish due to variation of egg-size effects between years and habitats. Feeding frequency may affect offspring state but offspring state may also affect feeding frequency. Varying causality between feeding rate and offspring state may preclude the detection of a positive effect of the former on the latter.

Cross‐fostering eggs reveals that female collared flycatchers adjust clutch sex ratios according to parental ability to invest in offspring

Across animal taxa, reproductive success is generally more variable and more strongly dependent upon body condition for males than for females; in such cases, parents able to produce offspring in above‐average condition are predicted to produce sons, whereas parents unable to produce offspring in good condition should produce daughters. We tested this hypothesis in the collared flycatcher (Ficedula albicollis) by cross‐fostering eggs among nests and using the condition of foster young that parents raised to fledging as a functional measure of their ability to produce fit offspring. As predicted, females raising heavier‐than‐average foster fledglings with their social mate initially produced male‐biased primary sex ratios, whereas those raising lighter‐than‐average foster fledglings produced female‐biased primary sex ratios. Females also produced male‐biased clutches when mated to males with large secondary sexual characters (wing patches), and tended to produce male‐biased clutches earlier within breeding seasons relative to females breeding later. However, females did not adjust the sex of individuals within their clutches; sex was distributed randomly with respect to egg size, laying order and paternity. Future research investigating the proximate mechanisms linking ecological contexts and the quality of offspring parents are able to produce with primary sex‐ratio variation could provide fundamental insight into the evolution of context‐dependent sex‐ratio adjustment.

Nest design and the abundance of parasitic Protocalliphora blow flies in two hole-nesting passerines

ABSTRACT Ectoparasites dwelling in bird nests regularly reduce reproductive success and condition of breeding birds. Thus, establishing the factors that determine the abundance of ectoparasites is important for better understanding of reproductive trade-offs and life history evolution in birds. A recent hypothesis states that interspecific differences in the abundance of ectoparasites may be caused by nest composition. For example, great tits (Parus major) have nests made of mosses and fur, whereas Ficedula flycatchers have nests made of grasses, bast, and bark, and tits are more infested by nest-dwelling ectoparasites than flycatchers. We swapped nests between pairs of great tits and collared flycatchers (F. albicollis) during egg-laying or early incubation and counted parasitic Protocalliphora blow flies at the end of breeding to test this hypothesis experimentally. We controlled statistically for habitat (oak versus spruce forest), brood size, season, year, and mean nestling weight before fledging. We found a significant effect of bird species (tit > flycatcher), habitat (oak > spruce), and year. There was no effect of nest type. Consequently, the hypothesis ascribing the different abundance of ectoparasites in great tits and collared flycatchers to different nest composition was not supported by our study.

Superiority of extra-pair offspring: maternal but not genetic effects as revealed by a mixed cross-fostering design

Extra‐pair copulations (EPC) are the rule rather than an exception in socially monogamous birds, but despite widespread occurrences, the benefits of female infidelity remain elusive. Most attention has been paid to the possibility that females gain genetic benefits from EPC, and fitness comparisons between maternal half‐siblings are considered to be a defining test of this hypothesis. Recently, it was shown that these comparisons may be confounded by within‐brood maternal effects where one such effect may be the distribution of half‐siblings in the laying order. However, this possibility is difficult to study as it would be necessary to detect the egg from which each chick hatched. In this study, we used a new approach for egg‐chick assignment and cross‐fostered eggs on an individual basis among a set of nests of the collared flycatcher Ficedula albicollis. After hatching, chicks were ascribed to mothers and therefore to individual eggs by molecular genetic methods. Extra‐pair young predominated early in the laying order. Under natural conditions, this should give them a competitive advantage over their half‐siblings, mediated by hatching asynchrony. However, we experimentally synchronized hatching, and after this treatment, extra‐pair young did not outperform within‐pair young in any studied trait including survival up to recruitment and several indicators of reproductive success and attractiveness. We obtained only modest sample sizes for the last two traits and did not test for extra‐pair success of male offspring. Thus, we cannot exclude the possibility of advantages of extra‐pair young during the adult phase of life. However, our data tentatively suggest that the more likely reason for females’ EPCs is the insurance against the infertility of a social mate.

Context dependence of maternal effects: testing assumptions of optimal egg size, differential, and sex allocation models

If offspring develop in adverse conditions, the maternal component of their phenotypic variation might increase due to the stronger dependence of offspring traits on parental investment. This should result in increased parental investment to individual offspring, as assumed by the model of optimal egg size. The opposite pattern, i.e., stronger dependence of offspring fitness on parental investment and consequently larger parental investment under good conditions is assumed by both the theory of differential allocation if attractive males provide material benefits, and reproductive compensation if they invest less into paternal care. Another influential idea is the Trivers-Willard model, which assumes sex-specific dependence of offspring fitness on parental investment. Here we tested these ideas by examining the effects of egg size on offspring fitness across many postnatal contexts in the Collared Flycatcher Ficedula albicollis. We employed a cross-fostering design that generated variation in egg size within nests and used brood means of fledgling mass as a functional measure of the quality of rearing conditions. Effects of egg size on three offspring traits, including lifetime reproductive success of recruits, were more pronounced in low-quality broods. These results support the assumption of the model of optimal egg size. Based on female preference for males providing material benefits, this pattern could support differential allocation, if attractive males invest less in paternal care, or reproductive compensation, if they invest more. By comparison, we did not find any evidence for sex specificity of fitness returns that might explain sex monomorphism of egg size in this species. The challenge for future studies will be the integration of components of parental investment and offspring fitness into their global measures and testing how the former affects the latter across gradients of postnatal conditions.