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It has been proposed that blue‐green egg coloration is a condition‐dependent female sexual trait that may modify paternal care in a post‐mating sexual selection scenario. This pattern may arise because the pigment responsible for eggshell colouration (biliverdin) may be a costly and limited resource, whose availability is linked to female health state. Thus, it can be predicted that females whose condition is compromised should be constrained in their capacity to deposit biliverdin in the eggshell, thus producing paler clutches. To test this hypothesis, we performed a handicapping experiment by clipping some feathers of female spotless starlings before egg laying and measuring the colour of their clutches. We expected the handicapping treatment to increase flying costs, impairing female overall condition and resulting in paler clutches. Our experiment was successful in lowering the weight gain of handicapped with respect to control females. However, in contrast to our expectations, we found no effect of the treatment on eggshell colouration. Eggshell colour varied along the laying order, with initial eggs of the laying sequence being relatively paler than the rest of the clutch, but this pattern was not different between experimental groups. Despite a very similar methodology, our results differ from a previous study on the same species and offer no support to the post mating sexual selected hypothesis, questioning the general applicability of the sexual selection role of eggshell coloration. [ABSTRACT FROM AUTHOR]

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]

Inter‐ and intraspecific variation in eggshell colouration has long fascinated evolutionary biologists. Among species, such variation may accomplish different functions, the most obvious of which is camouflage and background matching. Within species, it has been proposed that inter‐female variation in eggshell pigmentation patterns can reflect egg, maternal or paternal traits and hence may provide cues to conspecifics about egg, maternal or paternal phenotypic quality. However, the relationship between protoporphyrin‐based eggshell pigmentation and egg or maternal/paternal traits appears to be highly variable among species. We investigated patterns of intraspecific variation in Eurasian barn swallow Hirundo r. rustica protoporphyrin‐based eggshell pigmentation, and analysed its association with egg and clutch characteristics, maternal/paternal phenotypic traits and parental feeding effort. Eggshell pigmentation pattern significantly varied between breeding colonies, was significantly repeatable in first clutches laid by the same females in different years (intraclass correlation coefficient ranging between 0.56 and 0.63), but it was not significantly associated with egg traits, such as position in the laying sequence, egg mass, yolk testosterone concentration and antioxidant capacity. It was weakly or non‐significantly associated with female and male traits (sexual ornaments), but females laying darker (higher pigment intensity) first clutches had higher hatching success, suggesting that eggshell pigment intensity may predict fitness. Male nestling feeding effort was not predicted by eggshell pigmentation. In addition, females with darker breast plumage colouration (a melanin‐based trait related to fitness) laid highly protoporphyrin‐covered eggs, suggesting the presence of a previously unappreciated link between protoporphyrin biosynthesis and plumage melanisation. Moreover, the proportion of male offspring increased in clutches originating from highly protoporphyrin‐covered eggs, suggesting that parents could acquire visual cues about their future brood sex composition before egg hatching. Our results support the idea that intraspecific signalling via eggshell pigmentation is a species‐specific rather than a general feature of avian taxa. [ABSTRACT FROM AUTHOR]

Incubation prior to clutch completion may be adaptive if it maintains egg viability by inhibiting eggshell microbial growth, thus reducing the likelihood that the embryo becomes infected. To test this hypothesis, we examined the effect of partial incubation during egg laying on eggshell microbial loads in eastern bluebirds Sialia sialis breeding at a temperate‐zone site. We sampled eggshell microbes prior to and following four days of exposure to either partial incubation during the laying period or ambient environmental conditions without incubation (experimental eggs). Microbial colony counts declined significantly for eggs left in the nest during the laying period but did not vary significantly for eggs exposed to ambient conditions. Initial microbial loads were more similar to those previously reported from tropical than temperate environments, and microbes from potentially pathogenic groups were detected on 88% of first‐laid eggs on the day of laying. Egg viability was maintained when eggs were held indoors for four days without incubation but declined sharply thereafter. Our results suggest that partial incubation during egg laying may enhance egg viability in eastern bluebirds by reducing eggshell microbial loads; these effects appear stronger than those usually reported from the temperate zone. [ABSTRACT FROM AUTHOR]

Variation in blue-green and brown coloration of avian eggshells could be affected by several factors, including environmental nutritional constraints. Better availability of nutrients could influence the synthesis and deposition of pigments into the eggshell, so we may expect a link between the food availability during egg formation, the body condition of the female and intensity of eggshell coloration. This hypothesis has received mixed support so far: in some bird species a positive correlation between female body condition and eggshell blue-green coloration could be demonstrated, but other studies failed to find a significant link. In this study, we experimentally limited the food availability for domestic canaries Serinus canaria prior to and during egg laying, and examined its effects on the biliverdin- and protoporphyrin-based eggshell coloration. Treatment had no significant effect on eggshell blue-green chroma and biliverdin concentration. However, we found a significant positive relationship between female body condition and eggshell background blue-green chroma in the control group, but not in the food restricted group. Females possibly experiencing a decline in antioxidant capacity due to food limitation may not be able to produce a blue-green eggshell colour intensity reliably indicating their body condition. Furthermore, foodrestricted canary females laid eggs with significantly higher eggshell brown chroma, spot intensity, and protoporphyrin concentration. Therefore, our results suggest that limitation in actual nutrient availability increases deposition of protoporphyrin into the eggshell, and it may also modify the association between female body condition and intensity of blue-green eggshell coloration. [ABSTRACT FROM AUTHOR]

Avian eggshell coloration may have arisen due to selection on the biological, chemical, or physical properties of the pigments embedded within the eggshell, or due to selection on the coloration that emerges due to pigment deposition. Within both hypothetical frameworks, pigment-based eggshell coloration would be related to metrics of egg quality; however, no one has evaluated the relative strength of coloration and pigment concentration in predicting egg quality. Here, we examined 66 European starling Sturnus vulgaris eggs and quantified eggshell biliverdin concentration (an antioxidant that produces the eggshell's blue coloration) and used 28 different coloration metrics derived from both photographic and spectrophotometric data. We also measured egg size, eggshell thickness, concentration of carotenoids in the yolk, and concentration of lysozyme in the albumen to capture variation in egg quality. We found that throughout the laying sequence, biliverdin concentration increased while eggshell thickness, yolk carotenoid concentration, and lysozyme concentration in the albumen all decreased, but this variation was not captured by any eggshell coloration metric. Both eggshell coloration and biliverdin concentration were negatively associated with yolk carotenoid concentration, but only eggshell biliverdin concentration was negatively associated with yolk mass. Lastly, biliverdin concentration explained, at most, only 46% of the variation for all eggshell coloration metrics. Our results suggest biliverdin concentration is a better predictor of egg quality than egg coloration in European starlings, supporting the hypothesis that eggshell pigment concentration per se may be the target of selection. [ABSTRACT FROM AUTHOR]

The striking variation in colour and maculation of bird eggs has fascinated biologists since centuries, and many hypotheses based on mechanical, physiological or signalling functions have been proposed for its evolution by natural and sexual selection. Protoporphyrin is the main eggshell pigment found in brown maculated eggs, and is assumed to function as a mediator of these selection processes. It is a precursor of heme with pro-oxidant properties, and hence a link between brown maculation and female condition has been proposed and tested in a number of studies, albeit with contrasting results. A variety of different visual methods have been used to quantify outer eggshell pigmentation, which has been assumed to correspond to overall quantity of protoporphyrin in the shell. Yet, this relationship has rarely been tested. The aim of this study was to apply four commonly used methods to assess pigmentation in great tit eggs with protoporphyrin as the predominant eggshell pigment, and to compare the results of these methods. Specifically, we 1) ranked eggshell pigmentation by human naked eye, 2) applied a granularity approach and 3) measured spectrophotometric reflectance of eggshell pigments. Second, we estimated the relationship between outer eggshell pigmentation (i.e. estimated by three different methods above) and true protoporphyrin concentration deposited in the entire shell measured by HPLC. Among-method estimates were significantly correlated for the traits describing pigment ' darkness ' only. While the model including scores based on human naked eyes explained 16% of the variance of pigment concentration in the entire shell, spectrometry explained 27%, and the granularity approach explained 40%. Thus, the estimation of true pigment concentration in the entire shell from the visible outer side of the shell is most reliable with the granularity approach. It is relevant for studies where the maintenance of the integrity of the eggs is essential. [ABSTRACT FROM AUTHOR]

Eggshell maculation of most passerines is due to the deposition of the pigment protoporphyrin which is produced during biosynthesis of blood haem. Its functional significance has only received empirical attention in recent years. This interest has generated a number of hypotheses of which some remain untested partly because the quantification of protoporphyrin is analytically challenging and can be prohibitively expensive. Many studies have therefore used the extent of eggshell spotting as a proxy for total eggshell protoporphyrin concentration, although this has not been formally tested. Pigment scoring involves recording visible eggshell pigment attributes, such as spot intensity, distribution and size. Since even immaculate eggs can contain some protoporphyrin, there remains doubt over the degree to which visible pigment correlates with total pigment content of the shell. In this study, we test whether visible pigment scoring can be used as a proxy for protoporphyrin concentration of an eggshell. We use pigmented eggshells of two common British passerine species to compare eggshell spot intensity, distribution and spot size (as used by the visual pigment scoring method) with direct measures of eggshell protoporphyrin concentration. In addition, we compared an alternative method of pigment scoring, the pixel pigment scoring method, using a computer programme to quantify the number of pixels exceeding a specified colour threshold. We demonstrate that although results from both scoring methods were positively correlated with eggshell protoporphyrin concentrations, the correlations were not sufficiently strong to be used as surrogates in studies where actual pigment concentrations are required. [ABSTRACT FROM AUTHOR]

Diet strongly influences the chemistry of vertebrate soft and hard tissues. Bird bone and eggshell mineral preserve reliable records of prey consumption, even beyond the life of the predator, and analyses of hard tissues have usefully reconstructed avian diet. Here, we assess the feasibility of a non-destructive method for distinguishing krill-poor from krill-rich diets in penguins. Krill (Euphausiaceae) are fluoride-rich, and penguins that consume krill produce fluoride-rich bones. The chemistry of bone mineral may be elucidated using Raman spectroscopy without recourse to specialised sample preparation. Published data from the diet of six penguin species were compared to a fluoride-informative spectral band (phosphate symmetric stretch, ν1-PO43−) in the Raman spectra of penguin humeri. Penguins that consume abundant krill (e.g. Adélie and emperor) have ν1-PO43−-band positions higher than 963 cm−1, whereas penguins that primarily eat teleost fish or cephalopods (e.g. Fiordland crested, Humboldt, little blue and yellow-eyed) have ν1-PO43−-band positions lower than 963 cm−1. A krill-rich diet can therefore be determined from the Raman spectra of penguin bones. Raman spectroscopy could be a useful supplement to existing diet analysis techniques. [ABSTRACT FROM AUTHOR]

The expression and impact of maternal effects may vary greatly between populations and environments. However, little is known about large-scale geographical patterns of variation in maternal deposition to eggs. In birds, as in other oviparous animals, the outermost maternal component of an egg is the shell, which protects the embryo, provides essential mineral resources and allows its interaction with the environment in the form of gas exchange. In this study, we explored variation of eggshell traits (mass, thickness, pore density and pigmentation) across 15 pied flycatcher populations at a large geographic scale. We found significant between-population variation in all eggshell traits, except in pore density, suggesting spatial variation in their adaptive benefits or in the females' physiological limitations during egg laying. Between- population variation in shell structure was not due to geographic location (latitude and longitude) or habitat type. However, eggshells were thicker in populations that experienced higher ambient temperature during egg laying. This could be a result of maternal resource allocation to the shell being constrained under low temperatures or of an adaptation to reduce egg water loss under high temperatures. We also found that eggshell colour intensity was positively associated with biliverdin pigment concentration, shell thickness and pore density. To conclude, our findings reveal large- scale between-population variation of eggshell traits, although we found little environmental dependency in their expression. Our findings call for further studies that explore other environmental factors (e.g. calcium availability and pollution levels) and social factors like sexual selection intensity that may account for differences in shell structure between populations. [ABSTRACT FROM AUTHOR]

The colourful appearance of avian eggshells is a prominent aspect of maternal reproductive effort in birds. Some differences in eggshell coloration have been reported to co-vary with various measures of maternal condition and these patterns support the hypothesis that, in some bird species, several aspects of eggshell colour (i.e. primary chroma and brightness) function as a signal of maternal and offspring quality to induce greater paternal investment. We directly quantified eggshell pigment concentrations of blackbird Turdus merula and song thrush T. philomelos eggshells and tested how the two key pigments (protoporphyrin IX and biliverdin) co-varied with other eggshell traits and egg constituents as measures of maternal reproductive investment, including total yolk carotenoid concentration, total lipid concentration, yolk mass, and shell thickness. Contrary to predictions, we detected few statistical patterns overall. We found that protoporphyrin IX concentration was negatively associated with blue-green chroma in blackbirds but not in song thrush. The concentration of protoporphyrin IX was significantly greater in blackbirds and also showed different patterns of association with total yolk lipids and yolk carotenoid concentrations between these two species (significant species interaction terms). Our results reveal that it is not appropriate to simply assume in these two avian species that reflectance-based eggshell colour measures are a suitable proxy for eggshell pigment concentrations or can be used as consistent predictors of maternal reproductive investment. These results highlight the need to assess and validate the strength and direction of the statistical relationships between eggshell colour measures, pigment concentrations, and maternal resource deposition in the egg for other species of birds. [ABSTRACT FROM AUTHOR]

The brown-headed cowbird Molothrus ater is a brood-parasite that lays eggs in nests of a wide range of host species, including the closely-related red-winged blackbird Agelaius phoeniceus and the dickcissel Spiza americana. Although cowbird eggs have accelerated development and hatch sooner than similar-sized host eggs, this development takes place within a thickened eggshell that could impede gas flux to the developing embryo. We tested the hypothesis that the accelerated development of the cowbird embryo relative to hosts is enabled by an increase in eggshell porosity that allows increased fluxes of respiratory gases to and from the developing embryo. We found cowbird eggshell thickness was significantly greater than the eggshells of these two common hosts. Although the number of pores per egg was similar among all three species, the total pore area per egg in cowbirds was significantly greater than that of either host, despite having a smaller eggshell surface area than the red-winged blackbird. Cowbird egg pore area was 1.9×larger than that of the red-winged blackbird. Cowbird eggshells had a significantly greater gas flux than those of the red-winged blackbird and the dickcissel. When conductance was normalized to published values of egg mass, cowbird eggs had a higher mass-specific conductance than red-winged blackbird or dickcissel eggs. These results are consistent with the hypothesis that the rapid development of brown-headed cowbird embryos is facilitated by increased eggshell porosity, and that changes in eggshell porosity represent an adaptation that enables cowbird eggs to hatch earlier than equivalently-sized host eggs. [ABSTRACT FROM AUTHOR]

While avian eggshell colouration has attracted biologists for decades, little is known about its variation within individuals. The main goal of this study was to explore within- and between-season repeatability of eggshell appearance in the great reed warbler Acrocephalus arundinaceus. To achieve this, we measured eggshell reflectance of first and replacement clutches of individual females within one breeding season, and the reflectance of their first clutches in two following breeding seasons. As environmental conditions may affect egg colouration, repeatability was estimated from linear mixed-effects models where we initially included temperature and rainfall during egg laying, as well as year, clutch order and egg-laying date as fixed effects, and clutch identity nested in females as random effects. Eggshell appearance within clutches showed moderate repeatability, while both within- and between-season repeatability in clutch colouration for individual females was low. Our findings indicate that low intra-clutch variation may have a function within the context of brood parasitism, i.e. in facilitating host recognition of alien eggs. With variable eggs between successive clutches, however, host females may need to relearn the appearance of their eggs with every clutch they lay. This could represent a significant constraint for host egg-discrimination abilities. Yet, whether environmental or intrinsic physiological factors are responsible for the variation in eggshell colouration between successive clutches of the same females still remain to be discovered. [ABSTRACT FROM AUTHOR]

The hypothesis that eggshell colouration is a sexually selected trait of female birds is based on the fact that biliverdin, the pigment responsible for blue-green colours of the eggshell, is a potent antioxidant and that only females with high antioxidant capacity can deposit higher concentrations of biliverdin as eggshell pigment. Antioxidants (e.g. carotenoids, vitamins) are also abundant in the egg yolk, which serve as nutrient reserves for the developing embryo, and eggshell colour intensity may reflect maternal investment in yolk antioxidants. Here, we test the relationship between blue-green eggshell colour intensity and concentration and amount of carotenoids, vitamin A, and vitamin E in the egg yolk of spotless starling Sturnus unicolor, a species for which we have previously shown good evidence of sexual selection driving egg coloration. As could be extrapolated from the hypothesis of sexual selection driving the evolution of blue-green eggshells, we found that eggshell colour intensity was positively related to the concentration and amount of carotenoids and vitamin E in the yolk. Thus, mothers may use egg colour intensity to signal to fathers the antioxidant status of their offspring. Moreover, we provide evidence suggesting that maternal yolk investment in more coloured eggs can also explain the detected association between feeding decisions of males and egg colour intensity that we have found previously in this species. [ABSTRACT FROM AUTHOR]

Current research into the evolution and adaptive function of avian eggshell pigmentation, including maculation, has focused mostly on signalling-based and structural function hypotheses but ignored the potential consequences of shell pigmentation for the developing avian embryo, especially in moderating the embryo's interaction with its light environment. The exposure of the eggs to sunlight that frequently accompanies avian incubation behaviour is one of the major evolutionary steps setting apart birds and reptiles, and coincides with the appearance of eggshell pigmentation. This suggests that shell pigments could play a major role in ensuring the successful development of the avian embryo. We propose that the effects of shell pigments on the egg contents should be considered in addition to established hypotheses of shell pigmentation such as crypsis, egg recognition or a possible structural function. This approach has the potential to identify trade-offs between different pigment functions and to resolve some of the long standing paradoxa in the evolution of eggshell colour, such as the occurrence of conspicuous blue eggs in passerines or the secondary evolution of white eggshells in cavity nesters. In particular, we identify seven hypotheses, which address how the interaction of eggshell pigments and the light environment may influence embryonic development. These hypotheses are the: thermo-regulation; UV-B protection; photo-acceleration; lateralization; circadian rhythm; photo-reactivation; and antimicrobial defence. We believe that the understanding of eggshell pigmentation will greatly benefit from taking these hypotheses into consideration when studying the functional significance of eggshell pigmentation and suggest a number of promising directions for future experimental and comparative research. [ABSTRACT FROM AUTHOR]

The rejection of eggs of brood parasites in several species of hosts is based on cues only at the blunt pole (BP) and not at the sharp pole (SP) of the foreign and own eggshell. We investigated whether intraclutch variation is confined to a specific egg pole in species where the extent of intraclutch variation in the overall egg appearance is known to positively covary with either egg rejection rate or the probability of being parasitized. For the two poles separately, we analysed intraclutch variation of eggshell brightness and blue chroma. We quantified intraclutch variation as the standard deviations of these colour metrics, instead of their coefficients of variation which would represent a statistically flawed approach. Pooling measurements of brightness across the whole egg surface led to statistically non-significant results and masked positive correlations of BP brightness with egg rejection or parasitism risk, respectively. In contrast, patterns of blue chroma were important across the whole egg. Thus, the traditional whole egg 'averaging' approach may mask biologically important effects of intraclutch variation when the variation and potential signalling functions of egg appearance are confined to a specific egg part (brightness). However, analyses based on only BP and SP eggshell region specific data may also lack the power to detect effects of phenotypic traits that do not vary between egg poles (blue chroma). We advocate the use of a combination of region-specific and whole-eggshell based colour metrics and manipulations in cognitive, perceptual, and ecological studies of foreign egg rejection. [ABSTRACT FROM AUTHOR]

The developmental rate of cuckoo embryos and their hatching size is greater than that of host species, which may require more nutrient resources in the egg and more intensive gas exchange during development. In the present study, we compared various egg characteristics of a brood parasite, the common cuckoo Cuculus canorus, and its frequent host, the great reed warbler Acrocephalus arundinaceus. As maternally-derived testosterone is known to enhance growth rate of embryos and hatchlings, cuckoo eggs are expected to contain higher concentration of testosterone than host eggs. In addition, we expected higher concentration of antioxidants in cuckoo eggs to protect embryos from oxidative stress associated with accelerated growth. Our results showed that cuckoo eggs had thicker shells and higher pore density than great reed warbler eggs. Yolk was significantly heavier in cuckoo eggs and contained higher concentrations of carotenoids and vitamin E, however, yolk androgen and immunoglobulin concentrations were lower in cuckoo eggs as compared to great reed warbler eggs. We also examined whether eggshell colour was associated to egg quality, and detected a positive association between blue-green chroma and yolk antioxidant concentration in both species, suggesting that eggshell colour reflects the antioxidant investment of the female into the eggs. Our results suggest that cuckoo females increase the size, growth rate and competitive ability of their young by providing them with more nutrients and more dietary antioxidants for embryonic development, and not through elevated yolk testosterone or antibody levels. In addition, increased porosity of cuckoo eggshells may allow embryos to develop more rapidly because of a greater capacity of gas exchange. [ABSTRACT FROM AUTHOR]

Eggshell colour patterns play a crucial role in avian host–parasite coevolution. In contrast to many experiments investigating general host egg discrimination abilities, studies testing where specific recognition cues are located on the eggshells (on blunt, sharp or both egg poles) are lacking. Previous studies suggested that discrimination cues might be located at the blunt egg pole, where the shell patterning is typically concentrated. We tested this hypothesis experimentally in species subject to interspecific (great reed warblers Acrocephalus arundinaceus, reed warblers A. scirpaceus), and also intraspecific parasitism (song thrushes Turdus philomelos, blackbirds T. merula). We examined host responses towards two types of intraspecific eggs painted non-mimetic immaculate blue either at blunt or sharp poles. All four species rejected eggs manipulated at the blunt pole at significantly higher rates, indicating that they perceive the critical recognition cues in the same egg part. Thus, the presence of egg recognition cues at the blunt egg pole may be a general phenomenon in birds parasitized by both intraspecific and interspecific parasites. [ABSTRACT FROM AUTHOR]

The blue-green eggshell pigment biliverdin is an efficient antioxidant, the deposition of which may be costly and act as a handicap to the female's antioxidant capacity. Thus, it is hypothesized that there exists a positive correlation between the degree of eggshell coloration and the condition and health state of the mother. In this study, we measured the blue-green chroma of collared flycatcher Ficedula albicollis eggs, and investigated the associations between egg colour and parental characteristics, laying date, clutch size and laying order. We also analysed the possible correlations between egg size, yolk carotenoid and antibody levels and eggshell pigmentation. Our results revealed no significant relationships between egg colour and female age, morphological and serological variables. Blue-green colour intensity of eggs increased throughout the breeding season, which could be the result of either a proximate constraint or an adaptive strategy. We found a positive association between egg colour saturation and yolk lutein concentration. This relationship could be the result of the seasonal improvement of caterpillar supply, which could influence the deposition of both components into the egg. Furthermore, we found that females mated to lower-quality males produced eggs with higher biliverdin pigmentation. It is possible that deposition of biliverdin is correlated to some unmeasured aspect of egg quality that could enhance the viability of offspring of lower-quality fathers. We found that intensity of blue-green coloration increased with laying order between the first and fifth eggs. [ABSTRACT FROM AUTHOR]

We explore the relationship between general eggshell colour (background and spots) and proportion of surface covered by spots on the eggs of blue tits Cyanistes caeruleus and several variables reflecting female health and condition. Females laying more spotted eggs showed a poorer body condition, higher cellular concentration of the stress protein HSP70 and marginally lower total immunoglobulin levels in blood. In addition, these females were paired with males with higher levels of HSP70 and lower concentrations of immunoglobulins. We interpret these results based on the properties of the eggshell pigment protoporphyrin, which is known as an inductor of oxidative stress in females. Although observational, this study presents, for the first time, clear evidence of eggshell spottiness and general colour as indicators of general condition and stress in avian populations. [ABSTRACT FROM AUTHOR]

Recent studies have demonstrated that wild bird breeding in acidified areas may have difficulties in obtaining sufficient calcium for their eggshells, and that a shortage of land snails may be the cause. Here we present experimental evidence that calcium deficiency may affect reproductive traits also in birds breeding in non-acidified but naturally base-poor forest habitats. The study was conducted in Estonia in 1995 and 1996. We compared egg characteristics and other life-history traits of the Pied Flycatcher Firedula hypoleuca and the Great Tit Parus major in two habitats differing in natural availability of calcium (base-poor pine forest and base-rich deciduous forests). In both species, egg volume was smaller in the poor than the rich habitat. Eggshell thickness also tended to be less in the poor habitat in 199, but the difference was significant only in Pied Flycatcher. An experimental supplementation of calcium was performed in the habitats to determine to what extent differences between rich and poor habitats might be caused by differences in calcium availability. Egg volume and shell thickness were positively affected by calcium supplementation in the Pied Flycatcher. The same tendency was observed in the Great Tit in 1996, although not significant. However, there was no effect of calcium provisioning on hatching success, nest desertion rate and number or quality of fledglings. Our results show that calcium limitation may occur in wild birds on naturally calcium-poor soils. Birds breeding in such habitats may be vulnerable to possible increases in acid deposition. [ABSTRACT FROM AUTHOR]

Obligate avian brood parasites lay stronger eggs than their hosts or non‐parasitic relatives because they are rounder and have a thicker eggshell. Additionally, some other characteristics of the brood parasitic eggshells related to their microstructure such as size and orientation of calcite crystal units could also contribute to generating even stronger shells. An eggshell microstructure formed by small randomly oriented calcite crystal units can increase the robustness of the eggshells of birds. Here, the eggshell microstructure of avian brood parasites as well as their hosts have been characterized in detail, using X‐ray diffraction analyses to estimate the size and degree of orientation of calcite crystal units making the eggshell. Specifically, the brood parasitic great spotted cuckoo Clamator glandarius and two hosts (jackdaws Corvus monedula and magpie Pica pica) and one non‐host species (the pigeon, Columba livia domestica) were considered. Calcite crystal of the eggshell of the brood parasitic species was smaller and more randomly oriented than those of the eggshells of non‐parasitic species, which suggest that eggshell microstructure would contribute to explain why parasitic eggs are more resistant to breakage than those of their hosts. [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]

Eggshell pigmentation may signal the quality of the egg, that of the female and the environment, and thus nestling development may be related to this egg trait. However, so far, few studies have investigated the relationship between eggshell pigmentation and nestling development. Our aim was to study in a partial cross‐fostering experiment whether the protoporphyrin‐based eggshell pigmentation (spot intensity and spot distribution) showed any significant associations with several traits related to the development of nestling great tits Parus major. We found that nestlings from clutches with darker spotted eggshells hatched more synchronously, had higher hematocrit value and shorter wings. Females under adverse environmental circumstances were previously shown to lay darker‐spotted eggs, and these females probably delayed the start of incubation due to energetic constraints, resulting in a more synchronous hatching. The shorter wings of fledglings originating from clutches with darker‐spotted eggs may be caused by the lower general quality of their mothers, mediated through either genetic factors or early maternal effects. Darker spotted eggs may contain different levels of some biomolecules than paler spotted eggs, which could have an effect on nestling metabolism and activity, resulting in an elevated hematocrit value. More aggregated eggshell spotting was related to faster bone growth and slightly longer fledging tarsus length. Eggs with aggregated spotting may have thinner eggshells that could lead to hatchlings with smaller bones due to the lower availability of calcium during development. Nestlings may increase bone growth rate to compensate for their smaller initial size. Overall, our results indirectly suggest lower fitness for nestlings hatched from darker spotted eggs and for nestlings hatched from eggs with more aggregated spotting distribution due to the possible long‐term costs of synchronous hatching, shorter wings, and accelerated tarsus growth rate. [ABSTRACT FROM AUTHOR]