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Avian eggshell colouration might function as a post‐mating sexually selected signal of female quality, influencing male parental investment and, hence, reproductive success. This hypothesis has been tested for intrinsic eggshell pigments as biliverdin (blue‐green colouration) and/or protoporphyrin (brown coloured spots), but never for colourations applied post‐laying. Post‐laying staining colouration due to, for instance, uropygial secretion of the female could reflect its phenotypic properties and, thus, might be a cue for male investment in reproduction. In hoopoes, the uropygial gland of incubating females hosts symbiotic bacteria that are responsible for the brown colour of their uropygial secretion and of the eggshells, as they cover their bluish‐grey eggshells with gland secretion after laying. The secretion protects embryos from pathogenic trans‐shell infections and, thus, egg colouration may function as a cue or even as a post‐mating sexually selected signal of antimicrobial potential in hoopoes. In a wild hoopoe population breeding in nest boxes in Spain, we test this hypothesis by exploring whether egg colour predicts male parental investment. In accordance with the hypothesis, we found that the amount of food provided by males to incubating females was higher in nests with less saturated eggshell colours. This relationship was affected by female body condition. High quality females in terms of body condition and/or in secretion colour obtained better males in terms of provisioning effort during incubation. Given that eggshell saturation is negatively related to density of bacterial symbionts in uropygial secretions, one possibility is that males may regulate their parental investment in accordance to the expected characteristics of mutualistic bacteria hosted in uropygial glands and deposited on eggshells. We discuss alternative explanations for our results, concluding that the post‐mating sexual selection hypothesis is the most likely but experimental modification of egg colour is needed to test it further. [ABSTRACT FROM AUTHOR]

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 feather 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. [ABSTRACT FROM AUTHOR]

Animals can engage in visual displays, which may target conspecifics, heterospecifics or both. Here we studied the function of the flamboyant crest-raising display of hoopoes Upupa epops in experiments performed with males in captivity. Males were exposed to sounds of a conspecific (male hoopoe song), a potential predator (human voice), and two controls (the song of a blackbird Turdus merula, and background noise). These stimuli were presented to males in the presence and absence of females. Males raised the crest with a significantly higher probability when confronted with stimuli indicating potential threats (rival mate or predator) than with controls. The crest display was frequent when confronted with both kinds of threats independently of the presence of a female, suggesting that it was directed to the predator and the rival male. The probability of raising the crest was not related to body condition, and there was a marginally but not significant negative relationship between probability of raising the crest and the number of black spots on the crest feathers, which may suggest that crest display could be informing about male quality. Therefore, male hoopoes display the crest in a heterospecific context in response to detection of potential threats, which could be a deceptive or pursuit-deterrent signal. The results also support a role of the crest in sexual selection, suggesting that crest display in male hoopoes may serve multiple functions. [ABSTRACT FROM AUTHOR]

Animals live in a bacterial world, and detecting and exploring adaptations favouring mutualistic relationships with antibiotic-producing bacteria as a strategy to fight pathogens are of prime importance for evolutionary ecologists., Uropygial secretion of European hoopoes ( Upupa epops, Linnaeus) contains antimicrobials from mutualistic bacteria that may be used to prevent embryo infection. Here, we investigated the microscopic structure of hoopoe eggshells looking for special features favouring the adhesion of antimicrobial uropygial secretions., We impeded female access to the uropygial gland and compared microscopic characteristics of eggshells, bacterial loads of eggs and of uropygial secretion, and hatching success of experimental and control females. Then, we explored the link between microbiological characteristics of uropygial secretion and these of eggs of hoopoes, as well as possible fitness benefits., The microscopic study revealed special structures in hoopoes' eggshells (crypts). The experimental prevention of females' gland access demonstrated that crypts are filled with uropygial secretion and that symbiotic enterococci bacteria on the eggshells come, at least partially, from those in the female's uropygial gland. Moreover, the experiment resulted in a higher permeability of eggshells by several groups of bacteria and in elimination of the positive relationships detected for control nests between hatching success and density of symbiotic bacteria, either in the uropygial secretion of females or on the eggshell., The findings of specialized crypts on the eggshells of hoopoes, and of video-recorded females smearing secretion containing symbiotic bacteria at a high density onto the eggshells strongly support a link between secretion and bacteria on eggs. Moreover, the detected associations between bacteria and hatching success suggest that crypts enhancing the adhesion of symbiont-carrying uropygial secretion likely protect embryos against infections. [ABSTRACT FROM AUTHOR]

The intensity of selection exerted by brood parasites on their hosts depends on the proportion of nests that are parasitized and the fitness costs of parasitism. Nest detection by brood parasites influences the probability of parasitism, and we propose that the difficulty faced by brood parasites of finding nests on the ground may make ground-nesting species subject to lower levels of parasitism, causing a reduction in levels of defence compared with species breeding in shrubs, trees and elsewhere above the ground. We tested the prediction that the rejection rate of Common Cuckoo C uculus canorus eggs by hosts is inversely related to the frequency with which they build nests on the ground, both at local and at continental scales. First, we used estimates of the rejection rate of non-mimetic model eggs experimentally introduced into the nests of 26 potential host species breeding in the Sierra Nevada Mountains of southern Spain. Most species tested in the Sierra Nevada showed high rejection rates of both mimetic and non-mimetic eggs, whereas the European Robin E rithacus rubecula, with a low rejection rate, was the only species that was regularly parasitized. At the continental scale we used all available published information on rejection rates of non-mimetic models by European hosts of the Common Cuckoo. The frequency of ground-nesting explained interspecific variation in rejection rate of non-mimetic model eggs both for the species tested in the Sierra Nevada and for all European hosts after controlling for all other life-history variables known to affect rejection rates. An effect of the abundance of trees in a particular habitat, previously shown to affect parasitism by the Common Cuckoo, was only apparent from analyses of continental-scale data and not from the Sierra Nevada mountains, suggesting that particular properties of mountainous areas affect Common Cuckoo parasitism. Ground-nesting species showed lower rejection rates than species breeding in bushes or trees. These results suggest that species nesting on the ground may have suffered lower parasitism pressures in their historical coevolutionary interactions with the Common Cuckoo. [ABSTRACT FROM AUTHOR]

Selection pressures due to parasitism play an important role in driving the evolution of life history traits of birds in general and of behaviour at the nest in particular. Eggshell bacterial load has been shown to predict hatching failure (i.e. the probability of embryo infection) but the relationships between the bacterial environment of the nest and life history characteristics of birds remain poorly investigated. We explored interspecific variation in eggshell bacterial load of mesophilic bacteria, Enterococcus spp., Staphylococcus spp. and Enterobacteriaceae groups across 24 bird species and assessed whether bacterial load is associated with breeding traits. Interspecific variation was much higher than intraspecific variation for all measures of bacterial load even after controlling for annual variation. Thus, we were able to assess the correlation between bacterial community characteristics and life history traits. After correcting for phylogenetic effects, we found that nest type, the use of feathers or plants as lining material, and incubation behaviour explained a significant proportion of the variance in bacterial communities on eggshells. The strength of these associations depended on study year, suggesting an important role of environmental conditions for eggshell bacterial load or community. Overall, these results suggest that bacteria on eggshells are associated with bird species traits, probably because birds are mediating the deleterious effect of eggshell microbes through behavioural traits that modify bacterial load. [ABSTRACT FROM AUTHOR]

The common cuckoo, Cuculus canorus, is an evictor brood parasite that generally reduces host fitness to zero, exerting strong selective pressure on hosts to evolve egg recognition and rejection. However, a great deal of variation in egg-rejection efficiency exists, egg-rejection behaviour being considered a flexible conditional response against parasitic eggs. Recently, it has been shown that some of the individuals that recognized (pecked) the parasitic egg finally accepted it. Here, we present the results of two egg-recognition experiments made in two populations of rufous-tailed scrub robins, Cercotrichas galactotes, in which one plaster model or a real house sparrow's egg, respectively, was experimentally introduced into their nests. The hosts' response was video recorded, allowing us to quantify the number of pecks, pecking strength, and ejection behaviour. We have found that rufous-tailed scrub robin females ejected the model egg easily by grasping it after weakly pecking it; however, sometimes (55%) females pecked the experimental egg but did not eject it. This acceptance after pecking is not the consequence of a failure at puncture ejection, given that when confronted with real eggs (soft and easily punctured) in the second experiment, only 20% of pecked eggs were ejected, signifying that females that pecked at eggs without ejecting them had a low motivation to eject. We also discuss the effect of clutch inspection and the function of pecking strength. Finally, based on our own and previous research demonstrating pecking not followed by rejection, we propose a stepwise discrimination process in which accumulating motivation plays a key role in determining behavioural pathways shaping host response to parasitic eggs. [ABSTRACT FROM AUTHOR]

The uropygial glands of birds serve multiple functions, and there is great interspecific variability in the composition and properties of their secretions. A special case is the secretion in the hoopoes Upupa epops, and green woodhoopoes Phoeniculus purpureus, which, contrary to the commonly white and odourless secretions, are dark with pungent odour. Recently, bacteria have been isolated from glands of both woodhoopoes and hoopoes and here we test the hypothesis that bacteria are responsible of some of the special properties of glands and secretions of this group of birds. We explore natural seasonal changes and intersexual differences in the properties of hoopoe glands and secretions, check the natural occurrence of bacteria within secretions, and analyse the effect of experimental injection of antibiotics on uropygial gland properties. Male glands underwent no seasonal changes, and their secretions were invariably white and odourless, very similar to female glands outside the breeding season. However, in comparison to the uropygial gland of non-breeding females, those of incubating females showed a marked increase in size and volume of secretion produced, which became dark and pungent. All these parameters increased until the hatching date and returned to values similar to those in the prelaying phase towards the end of the nestling period. Nestling glands produced secretions similar to those of females in colour and odour. Gland size of both females and nestlings predicted the amount of secretion produced. Microscopic techniques confirmed the presence of bacteria at high density and in active division in all dark secretions examined. The antibiotic treatment significantly reduced the load of enterococci in nestling glands, did not affect size of glands, but diminished the volume of secretion, which was lighter in colour than that of control nestlings. In nesting females, the experimental injection of antibiotic affected some measurements of gland size and secretion colour. Because the experiment did not affect general health estimates (immunocompetence, body condition or growing) of nestlings, our results suggest that some of the special properties of hoopoe glands are mediated by the presence of symbiotic bacteria. [ABSTRACT FROM AUTHOR]

Animals should invest in the immune system to protect themselves from parasites, but the cost of immune responses may limit investment depending on resource availability. In birds’ broods, senior and junior chicks in size hierarchies face different rearing conditions, and thus we predicted that factors affecting immune response should differ between them. In asynchronously hatched hoopoe Upupa epops broods, we found that the immune response of senior nestlings was not related to their body condition, but positively related to risk of parasitism (which was indirectly estimated by laying date). This suggests that their immunocompetence is not limited by access to resources, and they can differentially invest in immune response with increasing risk of parasitism. On the other hand, immune response of junior nestlings was related to their body condition, but secondarily also to risk of parasitism. Our results agree with previous studies that have found significant influence of nutritional status and risk of parasitism on nestlings immune defence, but show that the effects of these environmental factors on nestling immunocompetence differ between nestlings occupying high and low rank positions in size hierarchies. The possible influence of maternal effects on the results found is also discussed. [ABSTRACT FROM AUTHOR]

Hoopoe ( Upupa epops, Coraciformes) males produce a very simple song during the breeding season in order to attract females and repel intruders. Strophes vary in length (i.e. number of elements) both within and between males, and previous studies have shown that this song cue is positively correlated with male condition and breeding success. In the present study we tested whether strophe length of males influences male behaviour during intra-sexual contests, in a colour-ringed population in southeast Spain. Paired males were presented with a recorded song with long strophes during the pre-laying period, while they were near their mates, in order to provoke male mate-defence behaviour. Most males responded to the playback, but the strategy of defence adopted depended on their own strophe length in spontaneous songs recorded before the experiments. While singing responses were common to most of the males, only those using long strophes adopted the most risky strategy of approaching the loudspeaker. However, the males that approached produced abnormal songs during playback, that were shorter and with fewer strophes than those of males that did not approach, and used shorter strophes in comparison with spontaneous songs before the experiment. These differences in quality of the song produced in response to the playback suggest that long-strophe males were basing their response mainly on attacking rather than singing, while short-strophe males tried to resolve the contest at a distance by means of their song. These results show that strophe length reflects some component of the competitive ability of males (either physical strength or aggressiveness) in the hoopoe, which together with previous results regarding its role for female choice, show that it is a sexual signal with dual function. [ABSTRACT FROM AUTHOR]

Abstract Egg rejection is the most common defence used by hosts against avian brood parasites and experimental studies have provided some of the best documented demonstrations of the coevolutionary process. However, the sex responsible for egg ejection and whether eggs are grasped or punctured are two essential questions that remain unanswered for most host species. In this paper, by filming the behaviour of individuals of three different species confronted with a foreign egg experimentally introduced into their nests, we first determine the relationship between recognition (when the birds aggressively pecked the experimental egg) and ejection. Secondly, we demonstrate that in the species where only the female incubates, only the female recognizes and ejects the model egg, whereas in the two species where both sexes incubate, both sexes eject the foreign egg. Finally, the large host species ejected the model egg by grasping it with the bill, whereas the two small species ejected it by puncturing it first. Furthermore, our data suggest that puncture ejection is more costly than grasping ejection considered both in terms of energetic and ejection costs. [ABSTRACT FROM AUTHOR]

Previous studies of the Hoopoe Upupa epops have shown that the strophe length of male songs influences female mate choice, and is correlated with female reproductive rates and male production of fledglings in the male's own brood. However, frequent interactions between breeding pairs and non-pair males suggests that extrapair copulations could occur and could affect the real number of fledglings sired by males, and therefore the relationship between strophe length and breeding success. Here we analyse the incidence of interactions between breeding pairs and non-pair males, and of extrapair paternity, the interrelation of these parameters, the influence of male strophe length on them, and whether extrapair fertilizations affect the correlation between strophe length and breeding success of males, in a colour-ringed population of Hoopoes in south-eastern Spain. Multilocus DNA-fingerprinting revealed that 10% of the broods contained offspring sired by extrapair males, representing 7.7% of the chicks. However, the interactions between pairs and non-pair males were more frequent, with more than 25% of broods being visited by non-pair males, and about 10% being helped (fed or defended) by males other than the nest owner. Most of these relationships were apparently attempts by visitor males to obtain copulations with paired females, or to obtain access to such females or nests in future breeding attempts. However, there was no significant link between the detection of interactions with alien males in a nest and the occurrence of extrapair paternity in it, indeed extrapair paternity was found in only 30% of the nests with interactions, and therefore the detection of visits or helping by non-pair males cannot be considered evidence of extrapair paternity in visited or helped broods. Males that sang with long strophes never suffered losses of paternity within their broods, while 25% of males that sang with short strophes did. However, these differences were not significant. Nevertheless, strophe length of males was significantly positively correlated with per brood and seasonal production of fledglings after accounting for losses of paternity within their own broods. [ABSTRACT FROM AUTHOR]

Hoopoe Upupa epops males produce a very simple song, with a repertoire size of one, in which the main difference between different strophes of a male and between the songs of different males is the number of elements they include (strophe-length). In several passerine species it has been shown that strophe-length is a sexually selected trait that reflects male quality and correlates with reproductive success. Here we analyse whether in a non-passerine, the Hoopoe, strophe-length of males is correlated with several variables of their reproductive success. Females paired with males singing long strophes laid their first clutch earlier, produced larger first clutches and laid second clutches after a successful first one more frequently than those paired with males singing short strophes. Moreover, males with long strophes produced more fledglings in their first clutches and in the whole season, partly because they brought more food for the brood than males with short strophes. The relationships found are not mediated by age effects. Previously we have shown that Hoopoe females in the early spring are attracted preferentially to songs with long strophes. Here we show that males singing long strophes also obtain postpairing benefits in terms of reproductive success, and that females paired with these males obtain direct benefits because these males provide greater feeding effort in the second half of the nestling period. These findings support the hypothesis that in the Hoopoe, strophe-length is a sexually selected cue under direct selection. [ABSTRACT FROM AUTHOR]

Nest re‐use in birds has the potential cost of infection by parasites and pathogens but may also be a source of beneficial symbiotic bacteria transmitted horizontally. Eurasian hoopoes Upupa epops host antibiotic‐producing bacteria in their uropygial gland but only while breeding, which suggests that the nest‐hole may be a source of those symbionts. Interestingly, hoopoes do not build nests, thus might prefer for reproduction nest holes with soft materials from previous reproductions. Here, we tested experimentally this preference by installing in the field new nest boxes that were left empty or filled with either sawdust or a mixture of sawdust and hoopoe's nest material from the previous year. We explored the experimental effect on the composition of the uropygial secretion bacterial community, on eggshell bacterial loads, and on several proxies of reproductive success. Hoopoes bred significantly more often in nest boxes with nest material than in empty ones, but the type of nest material did not affect nest box occupancy. Eggs in nest boxes with old‐soft material harbored higher bacterial density on their shells, and the microbiota of the uropygial secretion of nestlings and females in these nest boxes differed from those in nest boxes without old‐soft material. Moreover, although the experiment did not affect breeding success or related proxies, several operational taxonomic units from female uropygial secretions were positively associated with hatching success. This is the first experimental evidence showing that re‐used nest material affects the bacterial community of the uropygial secretions of hoopoe females. This suggests that the nest material can be a source of strains for their incorporation to both the uropygial gland and eggshell communities, highlighting a possible advantage of nest re‐use previously unconsidered. [ABSTRACT FROM AUTHOR]