Blue-green Eggs Research Articles

Transcriptome analyses of shell color and egg production traits between the uteruses of blue-green eggshell chickens and Hy-Line brown layers

Blue-green eggs exhibit unique shell color; however, compared to commercial layers, blue-green eggshell chickens have lower egg production and lack uniform shell colors. Aiming to confirm the molecular mechanisms that affect shell color and egg production, this study collected the uteruses of 12 blue-green eggshell chickens (BG group) and six Hy-Line layers (Brown group), which had significantly different shell color indexes (SCI) and egg numbers at 300 days of age (EN300). Transcriptome sequencing and comparative analyses were subsequently performed. BG hens were divided into two groups for comparative analysis (BGblue vs. BGgreen and BGlow vs. BGhigh, respectively), based on the differences in SCI and EN300, respectively. The result of weighted gene co-expression network (WGCNA) analysis showed that the sequenced and mapped 16,785 genes were clustered into 18 modules, among which six modules with a total of 4270 genes were highly correlated with SCI and EN300 traits. Five hundred and eleven differentially expressed genes (DEGs) belonged to the six key modules. Through KEGG mapping, GO enrichment, and Cytoscape network analysis, nine Hub genes were tightly associated with SCI and EN300. The up-regulated genes were CCR2, CCR8, CD40LG, IL18RAP, INHBA, and P2RY13, while the down-regulated genes were ABCA13, ADCY2, and GRM1. Co-analyses with the results of comparisons between the BG subgroups revealed that the expression of solute carrier (SLC) proteins and ABC transporters were highly related to eggshell color, while cytokine-cytokine receptor interactions and neuroactive ligand-receptor interactions were key pathways affecting egg production. The expression of extracellular cytokines and membrane receptors were significantly up-regulated in low-yield chickens. The candidate DEGs and pathways found in the study will assist in clarifying the molecular mechanisms affecting shell color and egg production, and improve the breeding of blue-green eggshell chickens.

Low evolutionary potential for blue-green egg colouration in a wild bird population.

Exploring the evolutionary architecture of female sexual traits and their potential evolvability is important to understand their possible role as post-mating sexual signals. Egg colouration has been proposed to be one of these post-mating sexual signals, honestly advertising female quality in birds, especially in blue-green laying species. In this study, we used an animal model in a Bayesian framework to estimate the evolvability of multiple descriptors of blue-green egg colouration and egg size in a wild long-term monitored population of spotless starlings (Sturnus unicolor). Our results show low to moderate heritability (h2 = 0.31 - 0.44) for three egg colour descriptors (blue-green chroma, chroma and lightness) and egg size. Using the coefficient of additive genetic variance (CVA) and the evolvability (IA) as proxies of evolutionary potential of all components of this trait, we found low values of CVA for all these variables, suggesting a small evolutionary potential of these phenotypic traits, contrasting to previous results reported in another blue-green egg laying species. Our results indicate a modest raw genetic material of this trait on which sexual selection can act upon and, therefore, a small probability for these traits to respond easily to selection.

First Documented Report of Complete Shell Pigment Loss in the Blue-Green Eggs of the Pied Flycatcher

First Documented Report of Complete Shell Pigment Loss in the Blue-Green Eggs of the Pied Flycatcher

Biogeographic history, egg colouration, and habitat selection in Turdus thrushes (Aves: Turdidae)

Despite broad interest and recent experimentation, there is no single ecological model accounting for the adaptive significance of the diversity of avian eggshell colouration. The often blue-green eggs of Turdus thrushes are a charismatic example of this, having long captured cultural and scientific attention. Although the biology and evolutionary history of “true” thrushes is well understood, little is known about correlated evolution between shifts in habitat and eggshell pigmentation, and how these shifts map with Turdus biogeography. We applied phylogenetic comparative methods to assess the evolutionary timing of divergence and variation of life history traits and eggshell colouration and maculation presence in the genus. We found that eggshell colour diversified independently on several occasions in the past 11 million years, with much of the variation occurring within the last 4 million years. The majority of Turdus species lay blue-green eggs and also tend to be sedentary and forest-dwelling. Diet generalist species and species which have transitioned to a forest habitat are more likely to lay white eggs (10% of studied species). In turn, lineages in any habitat were more likely to transition to blue-green eggs. We found that variation in egg colour is increased in some clades, of which two lineages radiated in South America and the East Palearctic, in the past 2–4 million years. These findings provide support for the hypothesis that white eggs are more conspicuous to predators in open environments and that multiple, non-mutually exclusive constraints operate on the adaptive function of avian eggshell colour.

Analysis of Egg Variation and Foreign Egg Rejection in Rüppell’s Weaver (Ploceus galbula)

Egg appearance is notable for its variation and as a source of recognition cues in bird species that are subject to egg-mimicking brood parasitism. Here I analyze the egg appearance of an East African weaverbird species that has variable eggs and is a host of brood parasitism by an egg-mimicking cuckoo, in order to (1) compare population variation to variation within a clutch as a measure of the distinctiveness of eggs; (2) assess modularity versus correlation among egg appearance traits as an indication of the complexity of egg signatures; and (3) address whether the eggs are discretely polymorphic or continuously variable in appearance. I also compare three methods of assessing egg coloration: reduction of spectral data to orthogonal components, targeted spectral shape variables, and avian visual modeling. Then I report the results of egg replacement experiments that assess the relationship between egg rejection behavior and the difference in appearance between own and foreign eggs. Rüppell’s weaver (Ploceus galbula) eggs are variable in appearance between individuals and consistent within a clutch, but vary widely in the distinctiveness of particular traits. Most aspects of color and spotting are decoupled from each other, including coloration likely to derive from different pigments. Egg ground color is bimodal, with a broad continuous class of off-white/UV eggs and another broad class of blue-green eggs. Variation in all other traits is unimodal and usually normal in distribution. Females reject foreign eggs on the basis of the difference in brightness of the ground color and spotting of foreign eggs relative to their own, and the difference in degree to which spots are aggregated at the broad end of the egg. This aggregation is among the most distinctive features of their eggs, but the brightness of the ground color and spotting brightness are not; the birds’ use of brightness rather than the more distinctive chromatic variation to recognize eggs might reflect the salience of achromatic contrast in a dim enclosed nest.

When are egg-rejection cues perceived? A test using thermochromic eggs in an avian brood parasite host.

At the core of recognition systems research are questions regarding how and when fitness-relevant decisions made. Studying egg-rejection behavior by hosts to reduce the costs of avian brood parasitism has become a productive model to assess cognitive algorithms underlying fitness-relevant decisions. Most of these studies focus on how cues and contexts affect hosts' behavioral responses to foreign eggs; however, the timing of when the cues are perceived for egg-rejection decisions is less understood. Here, we focused the responses of American robins Turdus migratorius to model eggs painted with a thermochromic paint. This technique modified an egg's color with predictably varying temperatures across incubation: at the onset of incubation, the thermochromic model egg was cold and perceptually similar to a static blue model egg (mimicking the robin's own blue-green egg color), but by the end of an incubation bout, it was warm and similar to a static beige egg (mimicking the ground color of the egg of the robin's brood parasite, the brown-headed cowbird Molothrus ater). Thermochromic eggs were rejected at statistically intermediate rates between those of the static blue (mostly accepted) and static beige (mostly rejected) model eggs. This implies that at the population level, egg-rejection relevant cues are not perceived solely when arriving to or solely when departing from the nest. We also found that robins rejected their own eggs more often when exposed to color-changing model eggs relative to static eggs, suggesting that recognizing variable foreign eggs entails costly rejection errors for this host species.

Anti-parasitic egg rejection by great reed warblers (Acrocephalus arundinaceus) tracks differences along an eggshell color gradient

One of the most effective defenses against avian brood parasitism is the rejection of the foreign egg from the host’s nest. Until recently, most studies have tested whether hosts discriminate between own and foreign eggs based on the absolute differences in avian-perceivable eggshell coloration and maculation. However, recent studies suggest that hosts may instead contrast egg appearances across a directional eggshell color gradient. We assessed which discrimination rule best explained egg rejection by great reed warblers Acrocephalus arundinaceus, a frequent host to an egg-mimetic race of common cuckoos Cuculus canorus. We deployed 3D-printed model eggs varying in blue-green to brown coloration and in the presence of maculation. Using visual modeling, we calculated the absolute chromatic and achromatic just-noticeable differences (JNDs), as well as directional JNDs across a blue-green to brown egg color gradient, between host and model eggs. While most model eggs were rejected by great reed warblers, browner eggs were rejected with higher probability than more blue-green eggs, and the rejection probability did not depend on maculation. Directional egg color discrimination shown here and in a suite of recent studies on other host species may shape the cognitive decision rules that hosts use to recognize foreign eggs and affect the course of evolution in parasitic egg mimicry.

Patterns of variation in eggshell colour of Mountain Bluebirds (Sialia currucoides) provide mixed support for the sexually selected eggshell colour hypothesis

Determining the functional significance of ornamental traits and whether they are shaped by sexual selection requires understanding how they vary among individuals, and their relationship with other measures of quality. Conspicuous eggshell coloration is a trait that may have evolved through sexual selection as a signal of female quality. This hypothesis is based on the possibility that the pigment used to colour blue-green eggs is limited and potentially costly to allocate to eggshells. We investigated the signalling potential of blue-green eggshell colour by quantifying patterns of within- and among-clutch colour variation in Mountain Bluebirds (Sialia currucoides) to determine if pigment was limited and if egg colour traits are intrinsic to individual females. We also explored relationships over two breeding seasons between eggshell colour and other female traits and investment in eggs. Eggshell colour was highly repeatable within clutches for most measures, and relatively consistent between breeding events in different years, indicating that eggshell colour is an intrinsic trait. Eggs in second clutches within a season had less saturated blue-green colour than in first clutches, and within clutches, later-laid eggs were brighter but generally less blue-green in colour than earlier laid eggs, providing evidence of pigment limitation. However, associations between eggshell colour and other female traits and investment in eggs were less clear, and inconsistent among study years. Our findings suggest that there is potential for blue-green egg colour to function as a signal of female quality, but further study is needed to determine if blue-green eggshell colour is a truly informative signal.

Dinosaur origin of egg color: oviraptors laid blue-green eggs.

Protoporphyrin (PP) and biliverdin (BV) give rise to the enormous diversity in avian egg coloration. Egg color serves several ecological purposes, including post-mating signaling and camouflage. Egg camouflage represents a major character of open-nesting birds which accomplish protection of their unhatched offspring against visually oriented predators by cryptic egg coloration. Cryptic coloration evolved to match the predominant shades of color found in the nesting environment. Such a selection pressure for the evolution of colored or cryptic eggs should be present in all open nesting birds and relatives. Many birds are open-nesting, but protect their eggs by continuous brooding, and thus exhibit no or minimal eggshell pigmentation. Their closest extant relatives, crocodiles, protect their eggs by burial and have unpigmented eggs. This phylogenetic pattern led to the assumption that colored eggs evolved within crown birds. The mosaic evolution of supposedly avian traits in non-avian theropod dinosaurs, however, such as the supposed evolution of partially open nesting behavior in oviraptorids, argues against this long-established theory. Using a double-checking liquid chromatography ESI-Q-TOF mass spectrometry routine, we traced the origin of colored eggs to their non-avian dinosaur ancestors by providing the first record of the avian eggshell pigments protoporphyrin and biliverdin in the eggshells of Late Cretaceous oviraptorid dinosaurs. The eggshell parataxon Macroolithus yaotunensis can be assigned to the oviraptor Heyuannia huangi based on exceptionally preserved, late developmental stage embryo remains. The analyzed eggshells are from three Late Cretaceous fluvial deposits ranging from eastern to southernmost China. Reevaluation of these taphonomic settings, and a consideration of patterns in the porosity of completely preserved eggs support an at least partially open nesting behavior for oviraptorosaurs. Such a nest arrangement corresponds with our reconstruction of blue-green eggs for oviraptors. According to the sexual signaling hypothesis, the reconstructed blue-green eggs support the origin of previously hypothesized avian paternal care in oviraptorid dinosaurs. Preserved dinosaur egg color not only pushes the current limits of the vertebrate molecular and associated soft tissue fossil record, but also provides a perspective on the potential application of this unexplored paleontological resource.

Egg discrimination along a gradient of natural variation in eggshell coloration.

Accurate recognition of salient cues is critical for adaptive responses, but the underlying sensory and cognitive processes are often poorly understood. For example, hosts of avian brood parasites have long been assumed to reject foreign eggs from their nests based on the total degree of dissimilarity in colour to their own eggs, regardless of the foreign eggs' colours. We tested hosts' responses to gradients of natural (blue-green to brown) and artificial (green to purple) egg colours, and demonstrate that hosts base rejection decisions on both the direction and degree of colour dissimilarity along the natural, but not artificial, gradient of egg colours. Hosts rejected brown eggs and accepted blue-green eggs along the natural egg colour gradient, irrespective of the total perceived dissimilarity from their own egg's colour. By contrast, their responses did not vary along the artificial colour gradient. Our results demonstrate that egg recognition is specifically tuned to the natural gradient of avian eggshell colour and suggest a novel decision rule. These results highlight the importance of considering sensory reception and decision rules when studying perception, and illustrate that our understanding of recognition processes benefits from examining natural variation in phenotypes.

Biliverdin- and protoporphyrin-based eggshell pigmentation in relation to antioxidant supplementation, female characteristics and egg traits in the canary (Serinus canaria)

Avian eggs exhibit a large variability in coloration and patterns, which are produced by blue-green biliverdin and red-brown protoporphyrin pigments. Several hypotheses have been proposed to explain the function of eggshell coloration. In this experimental study, we tested two hypotheses (signalling-function hypothesis and structural-function hypothesis) on both eggshell pigment types in an open-nesting songbird, the canary (Serinus canaria). Also, we aimed to examine whether deposition of pigments into the eggshell has any cost in terms of the plasma oxidative status of the female. We found that eggshell average blue-green chroma was increased by antioxidant supplementation, although we note that there had already been a pre-existing bias in plasma antioxidant capacity between the supplemented and control groups. Eggshell average blue-green chroma was positively related to female body condition during egg laying. However, blue-green eggshell colour was not related to female oxidative status during or after the laying period, and blue-green chroma increased with laying order. Accordingly, we found some support for that eggshell blue-green colour could reflect maternal antioxidant availability and body condition, but did not find evidence that it has a cost for the female’s oxidative status. By contrast, eggshell spot brightness was positively related to body condition, suggesting that darker spotting reflected poorer nutritional condition in the canary. Eggshell blue-green pigmentation was not significantly connected to the egg volume or yolk antioxidant level, but we found that eggs with lower yolk antioxidant concentration had higher average eggshell brown chroma. In sum, our results suggest that eggshell colour reflected female antioxidant and nutrient availability. Finally, we found that eggs with thinner eggshells had a more aggregated spot distribution, supporting the view that aggregated spots may help to strengthen eggshells. Avian eggs have a large variability in colours and patterns, which are due to two pigments: the blue-green biliverdin and the red-brown protoporphyrin. In this study, we tested whether eggshell pigmentation, measured by a spectrophotometer, could reflect female and egg quality in the canary, an open nesting songbird. We found that females supplemented with antioxidants before and during egg-laying laid more intense blue-green eggs. Females in better body condition laid eggs that had a more intense blue-green coloration, but lower intensity of brown coloration. Egg yolk antioxidant level was lower in eggs with more intense brown eggshell coloration. These results suggest that eggshell coloration could reflect the antioxidant and nutrient availability of females. Moreover, we found that eggs with thinner eggshells had a more aggregated spot distribution on the eggshell, supporting the view that aggregated brown pigment spots may help to strengthen eggshells.

The dinosaur that laid blue-green eggs

Mark Hauber at Hunter College in New York and his colleagues discovered that the color of many birds' eggs comes from the way that two pigments in the shell--biliverdin and protoporphyrin--blend with each other and with the calcium carbonate that makes most of the shell. Martin Sander of Bonn University in Germany has an idea. In a separate study, he and his team looked at eggs from prehistoric sites in China where Oviraptor dinosaurs laid their eggs millions of years ago. Dinosaur eggs found elsewhere are typically deep brown or black as minerals have seeped in over time and stained them, obscuring pigment molecules in the shell. Team member Jasmina Wiemann found the Oviraptor eggs contained biliverdin and protoporphyrin in a ratio suggesting they were blue-green.

Light dulls and darkens bird eggs.

Although egg color is generally consistent within individual birds and robust to environmental variation, recent evidence suggests a degree of susceptibility to environmental perturbation or modulation of egg color. Most of this variation manifests via the physiology of the laying female, but some direct impacts of the environment on laid eggs have also been discovered. Here we test whether light changes bird egg color and we quantify its effect, by subjecting variable blue-green eggs of Rüppell's weaver (Ploceus galbula) to a broad-spectrum light source under laboratory conditions, and measuring egg reflectance every few hours. Eggshells gradually decreased in reflectance across the entire measured wavelength spectrum of 250–800 nm. Reflectance peaks were disproportionately affected, such that the height decreased of both the blue-green peak and the smaller UV peak typical of blue-green eggs. The reflectance of lighter eggs was affected slightly more than that of darker eggs. These changes are similar to previous results for changes over long periods of time in darkness, suggesting that light might hasten the same process of pigment degradation that proceeds even without light. Comparison between the experimental light source and both sunlight and typical artificial lighting situations raises the possibility that significant color change might occur during incubation in some birds, but indicates that eggshell illumination in museums for short periods of study is unlikely to affect their color to a detectable extent. Additional research should be performed on eggs of other species and in other light environments, with an eye to an eventual generalized model of the effect of light on eggshell color.

HSP70 Level in Blood is Associated with Eggshell Blue-green Colouration the Pied Flycatcher

Among the cellular pathways conferring protection against oxidative stress, a key role is played by heat-shock proteins, especially HSP70. It has been proposed that blue egg colouration in certain birds, which is related to the deposition of the pigment biliverdin, should be related to basic antioxidant processes in the laying female given the strong antioxidant role of biliverdin. If HSP70 levels express antioxidant defences a higher allocation of the pigment to the clutch could induce a higher response by other components of the antioxidant system like HSP70. In a field study of a breeding population of pied flycatchers Ficedula hypoleuca, a species laying blue-green eggs, we measured spectrometrically the colouration of all eggs and collected blood samples of incubating females one week after clutch completion. We estimated female condition, plasma antioxidant capacity and level of HSP70 in blood cells. HSP70 level was positively related to plasma antioxidant capacity indicating that HSP70 is involved in responses to oxidative stress. Variation in HSP70 level was in good part explained by mean egg blue-green chroma of the clutch when controlling for plasma antioxidant capacity. Results suggest that intensity of eggshell blue colouration in this population is related to a basic system involved in protection against oxidative stress.

Eggshell Conspicuousness is Related to Paternal Brood Patch Vascularisation in the American Thrashers

Conspicuous blue-green egg colouration has been of interest to scientists and naturalists for more than a hundred years; however, the function of this trait has never been fully understood and has spurred extensive debate. The American thrashers exhibit great variation in egg colouration and patterning, and also show variation in male brood patch development. A recent hypothesis suggests that conspicuous egg colouration has evolved as the product of sexual conflict, where females may create conspicuous eggs to obtain greater amounts of male care, either in the form of male incubation or incubation assistance to prevent their eggs from being vulnerable to visually orienting predators and brood parasites. The American thrashers, a genus comprising of 10 species, exhibits striking variation in egg colouration and patterning, and also show variation in male brood patch development. By using avian visual modelling, I show that blue-green chroma of American thrasher eggs is positively related to their degree of conspicuousness against a nest background, while brown chroma is negatively related. Interestingly, the degree of male brood patch vascularisation is directly related to eggshell conspicuousness in American thrashers. I suggest that in the American thrashers, male brood patch vascularisation has evolved in response to conspicuous eggshell colouration to mitigate visual risks. Here I show that the degree of male brood patch vascularisation is directly related to relative paternal incubation effort in more than 300 North American breeding birds. This suggests that American thrasher species that possess both conspicuous eggs and brood patches, have most likely evolved to keep these conspicuous nest contents concealed, thereby reducing risk of visual detection. I suggest that these patterns are not unique to thrashers, but may extend to several avian families.

Attractive blue-green egg coloration and cuckoo−host coevolution

Recent evidence suggests that blue-green coloration of bird eggshells may be related to female and/or egg phenotypic quality, and that such colour may affect parental effort and therefore the nutritional environment of developing nestlings. Here we suggest that these relationships and the signal function of eggshell coloration would affect the outcome of coevolution between avian brood parasites and their hosts in at least three different non-exclusive evolutionary pathways. First, by laying blue-green coloured eggs, cuckoo females may exploit possible sensory biases of their hosts, constraining the evolution of parasitic egg recognition, and thus avoid rejection. Second, because of the relatively high costs of laying blue eggs, cuckoo females may be limited in their ability to mimic costly blue-green eggs of their hosts because cuckoo females lay many more eggs than their hosts. Furthermore, costs associated with foreign egg recognition errors would be relatively higher for hosts laying blue eggs. Third, cuckoos may use coloration of host eggs for selecting individuals or specific hosts of appropriate phenotypic quality (i.e. parental abilities). We here explored some predictions emerging from the above scenarios and found partial support for two of them by studying egg coloration of European cuckoos (Cuculus canorus) and that of their 25 main hosts, as well as parasitism and rejection rate of hosts. Cuckoo hosts parasitized with more blue, green, and ultraviolet cuckoo eggs, or those laying more blue-green eggs, were more prone to accept experimental parasitism with artificial cuckoo eggs. In addition, coloration of cuckoo eggs is more variable when parasitizing hosts laying bluer-greener eggs, even after controlling for the effect of host egg coloration (i.e. degree of egg matching). Globally, our results are consistent with the proposed hypothesis that host egg traits that are related to phenotypic quality of hosts, such as egg coloration, may have important implications for the coevolutionary interaction between hosts and brood parasites. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106, 154–168.

Is UV and blue-green egg colouration a signal in cavity-nesting birds?

We analysed the eggs of the European starling Sturnus vulgaris for variability of UV and blue-green colouration. We noted the calcium deposition in eggshells, egg measurements, light conditions in nests and starling eye properties. Egg colouration shows high within-clutch similarity. Intensity of the blue-green egg colour neither depends on Ca deposition in the eggshell nor correlates with egg measurements, clutch size and laying date. This suggests that the intensity of the blue-green colour of starling eggs does not reflect female condition, which is reported to affect egg and clutch parameters. Low light intensity in cavities enables only mesopic vision, which is characterised by degraded visual acuity and colour sense, as well as Purkinje shift. As a consequence, the probability of precise assessment of the differences in eggs colour by starling males, crucial for the signalling hypothesis (Moreno & Osorno2003), is not high. On the other hand, irrespective of its intensity, blue-green is the last colour visible under decreased illumination. Thus such coloured eggs may attract higher parental attention in dark cavities. We also demonstrated a strong negative correlation between intensity of blue-green and UV colouration of starling eggs, as well as constrained sensitivity of the starling eye to the range of eggs' maximum UV reflectance. Our study indicates that it is blue-green, not the UV component of the egg's spectral characteristics, which seems to increase the perception of eggs by adults in dark cavities. We suggest that it might have been the most important pressure towards evolution of blue-green colouration of eggs in cavity-nesting birds.

Blue-green eggshell coloration is not a sexually selected signal of female quality in an open-nesting polygynous passerine

It has been proposed that blue-green egg colours have evolved as a post-mating signal of female quality, selected by males allocating their parental effort in response to the strength of this signal. We tested two main assumptions of the sexually selected egg coloration hypothesis: (1) whether the intensity of eggshell blue-green chroma (BGC) reflects female quality; and (2) whether males make their decisions on the level of parental care that they provide according to the intensity of eggshell BGC. As a model species, we chose the facultatively polygynous great reed warbler (Acrocephalus arundinaceus). In this species, females simultaneously paired with the same male, compete for his nest attendance and could benefit from signalling their quality through egg coloration. However, we found no association between the variation in eggshell BGC and the measures of female quality (physical condition, mean egg volume and age). Moreover, great reed warbler males did not adjust their investment (as measured in terms of nest defence against a brood parasite) in relation to the eggshell BGC. We conclude that blue-green egg coloration in this open-nesting passerine is unlikely to have a signalling function. Rather, the large colour variation among clutches of individual females may depend on yearly fluctuations in environmental conditions.

Do Male Pied Flycatchers (Ficedula hypoleuca) Adjust Their Feeding Effort According to Egg Colour?

The colour of blue-green bird eggs has been hypothesized to signal female quality to attending males, who may adjust their level of investment in the brood accordingly. The hypothesis has gained support in studies of Spanish pied flycatchers Ficedula hypoleuca. We performed a cross-fostering experiment in a Norwegian population of pied flycatchers to provide an independent test of the sexually selected egg colour hypothesis in this species. Egg colour was not significantly correlated with estimates of female quality (clutch size, average egg volume, first egg laying date and feeding rate). There was a significant decrease in chroma and increase in brightness and egg volume during the laying sequence, with some marked differences between six-egg and seven-egg clutches that might reflect differences in female quality. However, we found no significant influence of egg colour (foster or original clutch) on male feeding rate or offspring viability (hatching success, average mass and fledging success) and no significant difference in feeding rate for males with six-egg and seven-egg foster or original clutches. We conclude that Norwegian male pied flycatchers do not use egg colour as a cue to female quality, thus questioning the generality of previous support for the sexually selected egg colour hypothesis from this species.

Robin’s egg blue: does egg color influence male parental care?

A recent hypothesis suggests that birds’ blue-green egg colors may be a sexually selected signal of female (and potentially nestling) quality that males use to make parental investment decisions. While there is some empirical support for this idea, both theory and observations question its validity. To test this hypothesis experimentally, we examined the influence of egg color on male American robin Turdus migratorius behavior by replacing natural clutches with four artificial eggs that were all either pale or vividly colored, close to the extremes in natural egg coloration. At the end of the incubation period, three unrelated nestlings were fostered into each experimental nest, and parental provisioning behavior was monitored when nestlings were 3, 6, and 9 days old. Male provisioning rate for 3-day-old nestlings was significantly higher in the vivid egg treatment compared to both the pale egg treatment and untreated controls, but there was no effect of egg color on paternal behavior at the older nestling stages. Male feeding rate at unmanipulated nests was only weakly positively related to natural egg color (chroma) when nestlings were 3 days old. These results suggest that blue-green egg color may act as a post-mating signal of female quality or investment in this species, but our findings do not exclude the possibility that egg color pigmentation also serves other adaptive functions.