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The adaptive value of routinely laying more eggs than can be successfully fledged has intrigued evolutionary biologists for decades. Extra eggs could, for instance, be adaptive as insurance against hatching failures. Moreover, because recent literature demonstrates that sibling cannibalism is frequent in the Eurasian hoopoe (Upupa epops), producing extra offspring that may be cannibalized by older siblings might also be adaptive in birds. Here, directed to explore this possibility in hoopoes, we performed a food supplementation experiment during the laying period and a clutch size manipulation during the hatching stage. We found that females with the food supplement laid on average one more egg than control females and that the addition of a close-to-hatch egg at the end of the hatching period increased the intensity of sibling cannibalism and enhanced fledging success in hoopoe nests. Because none of the extra nestlings from the experimental extra eggs survived until fledging, these results strongly suggest that hoopoes obtain fitness advantages by using temporarily abundant resources to produce additional nestlings that will be cannibalized. These results therefore suppose the first experimental demonstration of the nutritive adaptive function of laying extra eggs in vertebrates with parental care.

Parents might use signals of need or of quality to decide food provisioning among their offspring, while the use of one or another signal might depend on food availability. Begging success of nestlings of different quality (i.e., body size) would also depend on food availability, and we here explore the effect of experimental food supply in begging success of nestlings and in provisioning of female hoopoes (Upupa epops), a species with extreme hatching asynchrony and nestlings size hierarchy. We video-recorded food allocation of females, begging success of nestlings of different size, and the social context (i.e., the size category of the other nestlings that were begging for food) during periods when experimental food supply was or was not available in the same nests. We found that when experimental food supplementation was present, begging success of the intermediate, but not that of large or small-sized nestlings, increased. The experiment, however, did not affect the feeding preferences of females toward nestlings of different size. Moreover, when small nestlings were the only ones that were begging for food, their begging success decreased in the experimental period, and females used supplemented prey to feed themselves. Those results, on one hand, confirm the importance of food availability for the begging success of nestlings of particular sizes and, on the other, indicate that females prefer to use extra food for their own rather than for the smallest nestlings. We discuss possible mechanisms explaining the detected experimental effects and the adaptive and nonadaptive explanations of mothers ignoring the small nestlings.

Data file includes information about identity of each sampled individual, the species id, nest type, age, biometric measurements, the mean of the antagonistic halos shown against each 9 indicator bacteria, the antagonistic activity (average values of the width of antagonistic halos (mm) when tested against each of the nine indicator bacteria), the antagonistic range (Shannon index of the antagonistic activity) and the total density of bacteria on the gland.

Animal coloration results from pigments, nanostructures, or the cosmetic use of natural products, and plays a central role in social communication. The role of cosmetic coloration has traditionally been focused in scenarios of sexual selection, but it could also take place in other contexts. Here, by using spotless starlings (Sturnus unicolor) as a model system, we explore the possibility that nestlings cosmetically use their intensely yellow-colored uropygial secretion to signal their genetic and/or phenotypic quality. In agreement with the hypothetical cosmetic use of the uropygial secretion, (i) video recorded nestlings collected secretion with the bill at the age of feathering, (ii) cotton swabs turned to the color of secretion after rubbing with them nestlings’ gape, and (iii) gape and skin colorations correlated positively with that of secretion. Furthermore, we found that (iv) secretion coloration has a genetic component, and (v) associated positively with Vitamin E supplementation and (vi) with plasma carotenoid concentration, which highlights the informative value of nestling secretion. Finally, (vii) coloration of begging-related traits and of secretion of nestlings predicted parental feeding preferences. Consequently, all these results strongly suggest that the cosmetic use of colored uropygial secretion might also play a role in parent-offspring communication, complementing or amplifying information provided by the flamboyant colored gapes and skin of nestlings. The use of makeups by offspring for communication with relatives has been scarcely explored and we hope that these results will encourage further investigations in birds and other taxa with parental care.

Sibling cannibalism is relatively common in nature, but its evolution in birds and certain other vertebrates with extended parental care had been discarded. Here, however, we demonstrate its regular occurrence in two European populations of the Eurasian hoopoe (Upupa epops) and explore possible adaptive and non-adaptive explanations. Results showed that sibling cannibalism was more frequently detected in Spain (51.7%) than in Austria (5.9%). In these two populations, the hoopoes laid similar clutch sizes, resulting in similar fledging production, but hatching failures were more frequent in the northern population. Consequently, having more nestlings condemned to die in the southern population may explain the higher incidence of sibling cannibalism. In accordance with this interpretation, hatching span and failure, but not breeding date, explained the probability of sibling cannibalism in the Spanish hoopoes, while all three variables predicted brood reduction intensity. Furthermore, experimental food supply reduced the probability of sibling cannibalism, but not the intensity of brood reduction. Finally, females allocated fewer resources to the smallest nestlings when they were going to starve, but not necessarily when they were going to be used as food for their siblings. These results suggest that hoopoes produce extra eggs that, in the case of reduced hatching failure and food scarcity, produce nestlings that are used to feed older siblings. These findings provide the first evidence that sibling cannibalism occurs regularly in a bird species, thus expanding our evolutionary understanding of clutch size, hatching asynchrony, parent-offspring conflict, infanticide, and sibling cannibalism in the animal kingdom.

Symbiotic bacteria on animal hosts can prevent pathogenic bacterial infections by several mechanisms. Among them, symbiotic bacteria can indirectly enhance host's immune responses or, directly, produce antimicrobial substances against pathogens. Due to differences in life-style, different host species are under different risks of microbial infections. Consequently, if symbiotic bacteria are somewhat selected by genetically determined host characteristics, we would expect the antimicrobial properties of bacterial symbionts to vary among host species and to be distributed according to risk of infection. Here we have tested this hypothesis by measuring the antimicrobial ability of the bacterial strains isolated from the uropygial-gland skin of 19 bird species differing in nesting habits, and, therefore, in risk of microbial infection. In accordance with our predictions, intensity and range of antimicrobial effects against the indicator strains assayed varied among bird species, with hole-and open-nesters showing the highest and the lowest values, respectively. Since it is broadly accepted that hole-nesters have higher risks of microbial infection than open nesters, our results suggest that the risk of infection is a strong driver of natural selection to enhance immunocompetence of animals through selecting for antibiotic-producing symbionts. Future research should focus on characterizing symbiotic bacterial communities and detecting coevolutionary processes with particular antibiotic-producing bacteria within-host species.

Feathers are essential for avian life, and factors affecting their integrity are important to understand their evolution. These factors should depend on, among other traits, species‐specific bacterial environments and life‐history characteristics. However, interspecific variation in feather deterioration, feather susceptibility to degradation by keratinolytic bacteria (degradability), and bacterial environment, have rarely been quantified. Here, we did so by measuring deterioration and degradability of wing feathers of fledglings in 16 bird species, and characterizing the bacterial environment where they developed. We found statistically significant interspecific variation for all considered variables. On average, non‐melanised were more deteriorated than melanised feathers, but differences depended on the species. Moreover, nest bacterial loads were related to feathers wear, but the sign of the association depended on the bacterial group considered and on feather pigmentation. We also found a positive association of feather degradability with wear of non‐melanised feathers, and with bacterial loads. These results suggest that bacterial environments determine the integrity of fledgling feathers as well as their resistance to bacterial degradation, which implies a preponderant role of bacteria in driving the evolution of avian feathers.

[Simple Summary] Microorganisms may have direct negative effects on their animal hosts and cause diseases, but some others provide animals with protection against infections, parasites, and even predators. However, parasites or predators might take their cue from odors produced by bacteria, even those from protecting microorganisms, which could turn net benefits of bacteria that are a priori considered beneficial into a neutral or even negative outcome. This possibility has scarcely been studied in wildlife populations; we manipulated the bacterial community of nest-material of hoopoes and detected a negative effect in terms of the intensity of parasitism by blood-sucking flies that nestlings suffer. We also detected a positive link between the bacterial density of the nest-materials and the intensity of ectoparasitism, which further points at the importance of the bacteria determining the level of parasitism. Blood-sucking ectoparasites are also disease vectors, affecting both humans and livestock, and considering the role of the bacterial environment might help to establish new transmission control protocols., Nest bacterial environment influences avian reproduction directly because it might include pathogenic- or antibiotic-producing bacteria or indirectly because predators or ectoparasites can use volatile compounds from nest bacterial metabolism to detect nests of their avian hosts. Hoopoes (Upupa epops) do not build nests. They rather reuse holes or nest-boxes that contain remains of nest-materials from previous breeding seasons. Interestingly, it has been recently described that the nest’s bacterial environment partly affects the uropygial gland microbiota of hoopoe females and eggshells. Blood-sucking ectoparasites use chemical cues to find host nests, so we experimentally tested the hypothetical effects of microorganisms inhabiting nest-material remains before reproduction regarding the intensity of ectoparasitism suffered by 8-day-old nestling hoopoes. In accordance with the hypothesis, nestlings hatched in nest-boxes with autoclaved nest-material remains from the previous reproductive seasons suffered less from ectoparasites than those hatched in the control nest-boxes with nonautoclaved nest-material. Moreover, we found a positive association between the bacterial density of nest-material during the nestling phase and ectoparasitism intensity that was only apparent in nest-boxes with autoclaved nest-material. However, contrary to our expectations, nest bacterial load was positively associated with fledgling success. These results suggest a link between the community of microorganisms of nest-material remains and the intensity of ectoparasitism, and, on the other hand, that the nest bacterial environment during reproduction is related to fledging success. Here, we discuss possible mechanisms explaining the experimental and correlative results, including the possibility that the experimental autoclaving of nest material affected the microbiota of females and nestlings’ secretion and/or nest volatiles that attracted ectoparasites, therefore indirectly affecting bot

Research on the mechanisms involved in host location by parasites is of paramount importance and may aid in developing protective measures against them. This topic attains far-reaching repercussions for human and animal welfare regarding parasites transmitting vector-borne pathogens, such as blood-feeding flies. Very few studies have evaluated the effect of bird-derived cues on attraction of vectors in field conditions. We here explored the attraction of different groups of blood-feeding flies (mosquitoes, blackflies and biting midges) to auditory cues produced by begging hoopoe (Upupa epops) nestlings, and to three chemical cues derived from hoopoe nestlings or nests (uropygial secretion, symbiotic bacteria isolated from the secretion, and nest material) in the field. We deployed insect traps baited with the different stimuli at the beginning and at the end of the hoopoe breeding-season in four different habitats. Abundance of blood-feeding flies varied depending on habitat and sampling period. Begging auditory cues of nestling hoopoes did not affect abundance of flies. However, chemical stimuli affected abundance of mosquitoes, which were less abundant in traps baited with bacteria or with nest material than in control traps. Abundance of biting midges in traps also depended on the chemical stimulus but in interaction with sampling period or habitat. Fewer biting midges were collected in traps baited with bacteria and with secretion in the habitats where abundance of biting midges is higher. Our results suggest that uropygial secretion of hoopoes, and symbiotic bacteria living in this secretion, may repel blood-feeding flies from their nests.

It contains data collected in Guadix, Granada (southeast Spain) in 2019, in the framework of a research to evaluate the effects of chemical and auditory cues of hoopoes (Upupa epops) in repellence and attraction of blood-feeding flies. Dataset includes number of mosquitoes, blackflies and biting midges collected in insect traps in the field in four habitat types and two sampling periods, in relation to different chemical and auditory stimuli derived from nestlings of this bird species.

Sexually selected signals reliably reflect individual phenotypic or genetic quality and, thus, survival prospects of holders. Telomere length is considered a good predictor of life expectancy and, consequently, exploring the links between telomere length and sexually selected traits is much needed to better understand the mechanisms that maintain the honesty of sexual signals. We manipulated the length of throat feathers in spotless starling (Sturnus unicolor) males (a sexually selected signal) before reproduction and explored its effects on telomere shortening and breeding performance in subsequent reproductive events. We did not detect an effect of the feathers clipping manipulation, but males with longer throat feathers before the experiment had shorter telomeres that also shortened more slowly than those of males with shorter throat feathers did. Moreover, length of throat feathers of males before manipulation was positively related to hatching success of second clutches. Thus, correlative but not experimental results support the expected associations between sexually selected signals, telomeres and reproduction. We discuss such results in scenarios of sexual selection where feather length reflects, but does not directly cause, telomere attrition, and enhanced reproductive success. Males with longer throat feathers might be older, more experienced males (i.e., with shorter telomeres), able to buffer telomere shortening between reproductive events. Because of the absence of experimental effects, differential incubation effort of females cannot explain the detected association with hatching success, but other sexually selected traits that covary with throat-feather length could be responsible. Exploring those physiological and/or morphological characteristics related to throat-feather length should therefore be the matter of future research.

Within the context of complex sexual signalling, most research has focussed on exploring the associations between several signals and/or their relationships with different proxies of individual quality. However, very few studies have focused on checking whether the expression of one signal is conditioned by the expression of the others. Here, by experimentally shortening the throat feathers of male spotless starlings (Sturnus unicolor), we evaluated the influence of this trait on the colour expression of the bea19 base. In addition, we tested the relationship between these two sexually dimorphic characters with traits indicating individual quality such as body condition and colour reflectance at the wavelength related to carotenes in the tip of the beak. Our results show that the colouration of the beak base in males, but not in females, is positively related to body condition and to the length of ornamental throat feathers. Moreover, the experimental shortening of throat feathers in males had a negative effect on the blue chroma intensity of their beak base one year after manipulation. These results support for the first time a causal link between the expression of two sexually dimorphic characters, which is essential to understand their functionality in a multiple signalling framework

Sexually selected signals reliably reflect individual phenotypic or genetic quality and, thus, survival prospects of holders. Telomere length is considered a good predictor of life expectancy and, consequently, exploring the links between telomere length and sexually selected traits is much needed to better understand the mechanisms that maintain the honesty of sexual signals. We manipulated the length of throat feathers in spotless starling (Sturnus unicolor) males (a sexually selected signal) before reproduction and explored its effects on telomere shortening and breeding performance in subsequent reproductive events. We did not detect an effect of the feathers clipping manipulation, but males with longer throat feathers before the experiment had shorter telomeres that also shortened more slowly than those of males with shorter throat feathers did. Moreover, length of throat feathers of males before manipulation was positively related to hatching success of second clutches. Thus, correlative but not experimental results support the expected associations between sexually selected signals, telomeres and reproduction. We discuss such results in scenarios of sexual selection where feather length reflects, but does not directly cause, telomere attrition and enhanced reproductive success. Males with longer throat feathers might be older, more experienced males (i.e., with shorter telomeres), able to buffer telomere shortening between reproductive events. Because of the absence of experimental effects, differential incubation effort of females cannot explain the detected association with hatching success, but other sexually selected traits that covary with throat-feather length could be responsible. Exploring those physiological and/or morphological characteristics related to throat-feather length should therefore be the matter of future research