Variable environments impose constraints on adaptation by modifying selection gradients unpredictably. Optimal bird development requires an adequate thermal range, outside which temperatures can alter nestling physiology, condition and survival. We studied the effect of temperature and nest heat exposure on the reproductive success of a population of double‐brooded Spotless Starlings Sturnus unicolor breeding in a nestbox colony in central Spain with a marked intra‐seasonal variation in temperature. We assessed whether the effect of temperature differed between first and second broods, thus constraining optimal nest‐site choice. Ambient temperature changed greatly during the chick‐rearing period and had a strong influence on nestling mass and all body size measures we recorded, although patterns of clutch size or nestling mortality were not influenced. This effect differed between first and second broods: nestlings were found to have longer wings and bills with increasing temperature in first broods, whereas the effect was the opposite in second broods. Ambient temperature was not related to nestling body mass or tarsus‐length in first broods, but in second broods, nestlings were lighter and had smaller tarsi with higher ambient temperatures. The exposure of nestboxes to heat influenced nestling morphology: heat exposure index was negatively related to nestling body mass and wing‐length in second broods, but not in first broods. Furthermore, there was a positive relationship between nest heat exposure and nestling dehydration. Our results suggest that optimal nest choice is constrained by varying environmental conditions in birds breeding over prolonged periods, and that there should be selection for parents to switch from sun‐exposed to sun‐protected nest‐sites as the season progresses. However, nest‐site availability and competition for sites are likely to impose constraints on this choice.

Sibling competition has been shown to affect overall growth rates in birds. However, growth consists on the coordinated development of a multitude of structures, and there is ample scope for developmental plasticity and trade-offs among these structures. We would expect that the growth of structures that are used in sibling competition, such as the gape of altricial nestlings, should be prioritized under intense competition. We conducted an experiment in the spotless starling (Sturnus unicolor), cross-fostering nestlings to nests with different levels of sibling competition. We predicted that nestlings subjected to higher levels of sibling competition should develop larger gapes than control birds. We found that, halfway through the nestling period, overall size (a composite index of mass, wing, tarsus and bill) was reduced in nests with intense sibling competition, whereas gape width remained unaffected. At the end of the nestling period, experimental nestlings had wider gapes than controls. Additionally, a correlative study showed that nestling gape width increased when feeding conditions worsened and overall size decreased. These patterns could either be due to increased growth of gape flanges or to delayed reabsorption of this structure. Our results show that birds can invest differentially in the development of organs during growth, and that the growth of organs used in sibling competition is prioritized over structural growth.

Summary1. Social and ecological conditions experienced by individuals during early life can strongly influence their development and survival. Nestlings of many species present important variations in plasma androgens that can be associated with begging and sibling competition and may translate into fitness effects, since broods with higher testosterone (T) production may have better body condition and higher fledging success. However, the positive effects of androgens may be counterbalanced by a reduction of immune defences and a greater susceptibility to diseases. 2. In this study we examined the potential relationships between natural variation in plasma T, immunity and post-fledging survival rate in nestlings of the spotless starling (Sturnus unicolor). 3. We found that nestlings with higher cellular-mediated immune responses (CMI; measured as a swelling response to phytohaemagglutinin injection) were more likely to be recruited in the population than nestlings with lower CMI responses. Males presented higher CMI response than females, possibly due to differences in competitive advantage over food. We also found that CMI response was negatively related to T levels, as predicted by the immunocompetence handicap hypothesis. However, despite this reduction in CMI response, we failed to find an association between nestling T levels and survival prospects. Our results add to the evidence of the role played by immune defences in determining survival prospects in natural populations. 4. In conclusion, our study reveals that CMI response can be considered as a good predictor of post-fledging recruitment. As far as we know, this is the first study attempting to evaluate the relationship between nestling T and post-fledging survival. Our results suggest that the potential benefits accrued by high levels of T in sibling competition during the nestling stage do not translate into increased survival.

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The sexually selected egg colour hypothesis (SSECH) proposes that egg colouration is as a post‐mating sexually selected signal of female phenotypic quality, maintained by a higher allocation of paternal care. Similarly, some female traits can reflect genetic quality or condition and males could use this information in mate choice or in modulating parental investment. In our study, we examined the correlation of individual variation in egg colouration with female expression of a male ornament and how male feeding covaried with these two female traits in the spotless starling, in which egg colour varies widely between clutches and where both sexes possess showy throat feathers that are age dependent and that may signal individual quality. According to the SSECH, high‐quality females (females with longer throat feathers) are expected to lay more colourful eggs than low‐quality females and males should modify their feeding behaviour accordingly. By means of a principal component analysis, we found that most of the variation in egg colouration was due to brightness differences, and in a lower proportion to chromatic variation. Chromatic variation reflected a ultraviolet (UV) vs. greenness trade‐off and was positively associated with throat feather length: females with larger throat feathers laid eggs with higher UV and lower green reflectance. However, egg brightness was not related to female feather length, as the SSECH would predict. Male feedings were positively related to female throat feather length and negatively related to chromatic variation, meaning that males contributed more to nests of females with long throat feathers who laid eggs with higher UV and lower green reflectance. In conclusion, our data provide mixed support for the SSECH: although egg chromatic variation was related to female expression of a male ornament and male parental care, we found no evidence that egg brightness was involved in these processes.

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Spotless and European starlings (Sturnus unicolor and Sturnus vulgaris) have attracted attention from researchers interested in sexual selection, evolution of parental care and reproductive strategies. Both species show high levels of intraspecific nest parasitism and extra‐pair paternity, but research in this area is hampered by a lack of molecular markers specific for these species. Here, we describe a set of primers for nine microsatellite loci in spotless starlings, eight of which are highly polymorphic (7.37 alleles in average). These microsatellites are also polymorphic in European starlings (6.75 alleles in average).

Variation in avian reproductive strategies is often studied from a comparative perspective, since even closely-related taxa differ greatly in the degree of polygyny, extra-pair paternity (EPP) or intra-specific brood-parasitism. However, substantial variation at the species level suggests that ecological factors are important in shaping these patterns. In this study, we examined the temporal plasticity of these strategies, following a population from the year of colony formation to 2 years after this. Parentage data from these years shows that polygyny decreased with time, likely as a consequence of increased competition for nesting sites and mates by new recruits, and immigrants of higher quality arriving to the colony as time passed. In parallel to this temporal change, we found an increase in intra-specific brood-parasitism and quasi-parasitism (QP). We interpret these patterns as a consequence of an increase of floaters with time; these birds pursue a mixture of alternative mating strategies to succeed in the population. We also found evidence of conspecific brood parasitism (CBP), by nesting females that laid part of the clutch in another nest or that after losing a partially laid clutch resorted to lay the last eggs in another nest. Analyses of the distance between the main nest and nests containing the secondary polygynous brood or extra-pair or parasitic young showed an avoidance of contiguous nests for conducting these alternative reproductive tactics. At the same time, these secondary nests were closer to the main nest than random distances within the colony, suggesting that access to public information was restricted to a narrow area around the main nest. Our study emphasizes how behavioral patterns are plastic traits that vary not only with individual circumstances, but also with time, tracking changes in density and social structure.