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.     

Eggshell colour does not predict measures of maternal investment in eggs of <Emphasis Type="Italic">Turdus</Emphasis> thrushes

The striking diversity of avian eggshell colour has long fascinated biologists. Recently, it has been proposed that the blue-green colour of some eggs may function as a post-mating sexually selected signal of female phenotypic quality to their mates to induce higher allocation of paternal care. It has been suggested that maternally deposited yolk carotenoids may be the specific aspect of reproductive quality that the female is signalling via eggshell colour. We use the known properties of the thrush visual system (Turdus sp.) to calculate photon capture for the four single cone photoreceptors, and the principal member of the double cone class for eggs in clutches of two introduced European thrush species (Turdus merula and Turdus philomelos) in New Zealand. We show that differences in the avian-perceived colours of individual eggs are not consistently correlated with different measures of maternal investment in the egg. Given the growing extent of the knowledge between maternal quality, parental investment and eggshell pigmentation across avian taxa, we encourage the use of avian perceptual modelling for testing alternative non-signalling explanations for the structural and physiological basis of these relationships.     

Does contrast between eggshell ground and spot coloration affect egg rejection?

Obligate avian brood parasitic species impose the costs of incubating foreign eggs and raising young upon their unrelated hosts. The most common host defence is the rejection of parasitic eggs from the nest. Both egg colours and spot patterns influence egg rejection decisions in many host species, yet no studies have explicitly examined the role of variation in spot coloration. We studied the American robin Turdus migratorius, a blue-green unspotted egg-laying host of the brown-headed cowbird Molothrus ater, a brood parasite that lays non-mimetic spotted eggs. We examined host responses to model eggs with variable spot coloration against a constant robin-mimetic ground colour to identify patterns of rejection associated with perceived contrast between spot and ground colours. By using avian visual modelling, we found that robins were more likely to reject eggs whose spots had greater chromatic (hue) but not achromatic (brightness) contrast. Therefore, egg rejection decision rules in the American robin may depend on the colour contrast between parasite eggshell spot and host ground coloration. Our study also suggests that egg recognition in relation to spot coloration, like ground colour recognition, is tuned to the natural variation of avian eggshell spot colours but not to unnatural spot colours.     

Hoopoes color their eggs with antimicrobial uropygial secretions

Uropygial gland secretions are used as cosmetics by some species of birds to color and enhance properties of feathers and teguments, which may signal individual quality. Uropygial secretions also reach eggshells during incubation and, therefore, may influence the coloration of birds’ eggs, a trait that has attracted the attention of evolutionary biologists for more than one century. The color of hoopoe eggs typically changes along incubation, from bluish-gray to greenish-brown. Here, we test experimentally the hypothesis that dark uropygial secretion of females is responsible for such drastic color change. Moreover, since uropygial secretion of hoopoes has antimicrobial properties, we also explore the association between color and antimicrobial activity of the uropygial secretion of females. We found that eggs stayed bluish-gray in nests where female access to the uropygial secretion was experimentally blocked. Furthermore, experimental eggs that were maintained in incubators and manually smeared with uropygial secretion experienced similar color changes that naturally incubated eggs did, while control eggs that were not in contact with the secretions did not experience such color changes. All these results strongly support the hypothesis that female hoopoes use their uropygial gland secretion to color the eggs. Moreover, saturation of the uropygial secretion was associated with antimicrobial activity against Bacillus licheniformis. Given the known antimicrobial potential of uropygial secretions of birds, this finding opens the possibility that in scenarios of sexual selection, hoopoes in particular and birds in general signal antimicrobial properties of their uropygial secretion by mean of changes in egg coloration along incubation.     

Irregular brood patterns and worker reproduction in social wasps

The potential for reproductive conflict among colony members exists in all social insect societies. For example, queens and workers may be in conflict over the production of males within colonies. Kin selection theory predicts that in a colony headed by a multiply-mated queen, worker reproduction is prevented by worker policing in the form of differential oophagy. However, few studies have demonstrated that workers actually lay eggs within queenright colonies. The purpose of this study was to determine if workers laid male eggs within unmanipulated queen-right colonies of the polyandrous social wasps Vespula maculifrons and V. squamosa. We focused our analysis on an unusual brood pattern within colonies, multiple egg cells. We were primarily interested in determining if individuals reared in these irregular circumstances were queen or worker offspring. To address this question, we genotyped 318 eggs from eight V. maculifrons and two V. squamosa colonies. No worker‑reproduction was detected in any of the queenright colonies; all of the eggs found in multiple egg cells were consistent with being queen‑produced. However, the frequency of multiple egg cells differed among colonies, suggesting that queens vary in the frequency of errors they make when laying eggs within cells. Finally, we suggest that workers may not be laying eggs within queenright colonies and that worker reproduction may be controlled through mechanisms other than differential oophagy in polyandrous Vespula wasps.     

Moorhens have an internal representation of their own eggs

How do birds recognize their own eggs? Do they have a stored template for their own egg characteristics, or do they use another mechanism? Intraspecific brood parasitism is considered to be an additional reproductive tactic where females can increase their own reproductive success. Because of the costs involved in rearing young that are not their own, it will pay females to detect and reject the eggs of a parasite, although it is not known how they do this. Here, we show experimentally that moorhens will cease laying in a nest when their first egg is replaced with another hen’s egg but not when it is replaced with their own egg taken from an earlier clutch. This provides good evidence that birds have an internal representation of their own eggs and use this in decisions about whether to reject foreign eggs.     

Minute theropod eggs and embryo from the Lower Cretaceous of Thailand and the dinosaur-bird transition

We report on very small fossil eggs from the Lower Cretaceous of Thailand, one of them containing a theropod embryo, which display a remarkable mosaic of characters. While the surficial ornamentation is typical of non-avian saurischian dinosaurs, the three-layered prismatic structure of the eggshell is currently known only in extant and fossil eggs associated with birds. These eggs, about the size of a goldfinch's, mirror at the reproductive level the retention of small body size that was paramount in the transition from non-avian theropods to birds. The egg-layer may have been a small feathered theropod similar to those recently found in China.     

A drowned Mesozoic bird breeding colony from the Late Cretaceous of Transylvania

Despite a rapidly improving fossil record, the reproductive biology of Mesozoic birds remains poorly known: only a handful of undisputed, isolated Cretaceous eggs (some containing embryonic remains) are known. We report here the first fossil evidence for a breeding colony of Mesozoic birds, preserved at the Late Cretaceous (Maastrichtian) Oarda de Jos (Od) site in the Sebe? area of Transylvania, Romania. A lens of calcareous mudstone with minimum dimensions of 80?cm length, 50?cm width and 20?cm depth contains thousands of tightly packed, morphologically homogenous eggshell fragments, seven near-complete eggs and neonatal and adult avialan skeletal elements. Eggshell forms 70–80?% of the matrix, and other fossils are entirely absent. The bones exhibit clear characters of the Cretaceous avialan clade Enantiornithes, and the eggshell morphology is also consistent with this identification. Both taphonomy and lithology show that the components of this lens were deposited in a single flood event, and we conclude that it represents the drowned remains of a larger enantiornithine breeding colony, swamped by rising water, washed a short distance and deposited in a shallow, low-energy pond. The same fate often befalls modern bird colonies. Such a large concentration of breeding birds suggests aquatic feeding in this species, augments our understanding of enantiornithine biology and shows that colonial nesting was not unique to crown birds.     

Costs and benefits of subordinate queens in colonies of the ant,<Emphasis Type="Italic"> Leptothorax gredleri</Emphasis>

Queens of some species of social insects form dominance hierarchies in which only the top-ranking individual reproduces. Such unequal partitioning of reproduction can be stable when subordinate queens increase their inclusive fitness indirectly by helping to rear a related dominant queen's offspring. We investigated whether subordinate queens of the ant, Leptothorax gredleri, affect the reproductive success of the dominant queen. In laboratory colonies with two queens, first eggs were laid approximately 20 days later than in colonies with a single queen, presumably due to aggressive competition among nestmate queens. Nevertheless, two-queen colonies produced on average more brood than one-queen colonies, although egg laying was completely or almost completely monopolized by one single queen. The presence of a subordinate queen therefore appears to increase the productivity of the dominant.     

Sex-biased investment in yolk androgens depends on female quality and laying order in zebra finches (Taeniopygia guttata)

The Trivers–Willard hypothesis predicts sex biases in parental investment according to parental condition. In addition, parents may need to sex bias their investment if there is an asymmetry between the sexes in offspring fitness under different conditions. For studying maternal differential investment, egg resources are ideal subjects because they are self contained and allocated unequivocally by the female. Recent studies show that yolk androgens can be beneficial to offspring, so here we test for sex-biased investment with maternal investment of yolk testosterone (T) in zebra finch (Taeniopygia guttata) eggs. From the Trivers–Willard hypothesis, we predicted females to invest more in male eggs in optimum circumstances (e.g. good-condition mother, early-laid egg), and more in female eggs under suboptimal conditions (e.g. poor-condition mother, late-laid egg). This latter prediction is also because in this species there is a female nestling disadvantage in poor conditions and we expected mothers to help compensate for this in female eggs. Indeed, we found more yolk T in female than male eggs. Moreover, in accordance with our predictions, yolk T in male eggs increased with maternal quality relative to female eggs, and decreased with laying order relative to female eggs. This supports our predictions for the different needs and value of male and female offspring in zebra finches. Our results support the idea that females may use yolk androgens as a tool to adaptively manipulate the inequalities between different nestlings.     

Set a thief to catch a thief: brown-necked raven (Corvus ruficollis) cooperatively kleptoparasitize Egyptian vulture (Neophron percnopterus)

Our study describes how brown-necked ravens (Corvus ruficollis) are able to take advantage of an ordinarily inaccessible, high-quality food source by relying upon their innovative and manipulative thinking capabilities to exploit methods used by Egyptian vulture (Neophron percnopterus) to overcome the problem. In five observed interactions, the ravens were first seen in the vicinity of an abandoned clutch of ostrich eggs (Struthio camelus). The area was frequented by a pair of Egyptian vultures that bred on the cliffs across the road from the nature reserve. The Egyptian vulture exhibits tool use in birds, and is able to crack the hard shells of ostrich eggs by lifting a rock in the beak and pounding at the egg till it breaks open or cracks. If the egg is only cracked, the vulture inserts its narrow bill into the fissure and widens it by opening the mandibles. Pieces of eggshell are removed from around the crack in order to further open the egg. This is the point at which the pair of ravens attacked the vulture and harassed it till it abandoned the egg and left the area. The ravens then jointly enjoyed the contents of the egg which was otherwise inaccessible to them because of the strong egg shell.     

Queens and major workers of Acanthomyrmex ferox redistribute nutrients with trophic eggs

 In Acanthomyrmex ferox, two distinct egg-types are produced: reproductive eggs that give rise to offspring and trophic eggs that serve to distribute nutrients. Queens lay both reproductive and trophic eggs, while major and minor workers lay only trophic eggs in the presence of the queen. The queen lays on average 17% of the trophic eggs in a colony, while majors and minors produce 42% and 41%, respectively. The large proportion of trophic egg production by the queen and soldiers is quite remarkable, since ant queens are expected to focus entirely on reproduction and majors, which have a defensive function in many species, to be sterile. Received: 25 April 2000 / Accepted in revised form: 16 June 2000     

Sexually dichromatic coloration reflects size and immunocompetence in female Spanish terrapins, Mauremys leprosa

Many studies have shown the importance of colorful ornamentation in mate choosiness or intrasexual conflict. However, research on color ornaments has focused mainly on birds, lizards or fish, but remains practically unknown in other animal groups such as turtles. In addition, female ornaments and their relation with sexual selection also remain almost unknown. Here, we measured the coloration of the shell and the limb stripes of male and female Spanish terrapins Mauremys leprosa and explored the existence of sexual dichromatism and the relation of color characteristics with body size and health state estimated from the immune response to the injection of an antigen (phytohaemagglutinin test). Our results showed that shell coloration, which could be constrained by natural selection to be cryptic, changed with body size, but did not differ between sexes. In contrast, females had brighter and less ultraviolet-saturated and more orange-saturated limb stripes than males. In females, interindividual variation in limb stripe coloration was related with body size and immune response suggesting that this coloration may inform honestly about multiple traits that could be important in sexual selection. In contrast, coloration of limb stripes of males was duller than in females, and was not related with any trait suggesting that coloration is not important in sexual selection for males.     

Melanin-based coloration covaries with ovary size in an age-specific manner in the barn owl

While the adaptive function of black eumelanin-based coloration is relatively well known, the function of reddish-brown pheomelanin-based coloration is still unclear. Only a few studies have shown or suggested that the degree of reddish-brownness is associated with predator–prey relationships, reproductive parameters, growth rate and immunity. To gain insight into the physiological correlates of melanin-based coloration, I collected barn owl (Tyto alba) cadavers and examined the covariation between this colour trait and ovary size, an organ that increases in size before reproduction. A relationship is expected because melanin-based coloration often covaries with sexual activity. The results showed that reddish-brown juveniles had larger ovaries than whiter juveniles particularly in individuals in poor condition and outside the breeding season, while in birds older than 2 years lightly coloured females had larger ovaries than reddish-brown conspecifics. As barn owls become less reddish-brown between the first and second year of age, the present study suggests that reddish-brown pheomelanic and whitish colorations are associated with juvenile- and adult-specific adaptations, respectively.     

Color polymorphism in a land snail Cepaea nemoralis (Pulmonata: Helicidae) as viewed by potential avian predators

Avian predation is one of the most probable factors maintaining polymorphism of shell coloration in Cepaea nemoralis. This assumption is justified by the fact that birds frequently forage on snails and their prey choice varies with morph coloration. However, in all preceding studies, the conspicuousness of morphs was determined only by using human vision which is significantly different from birds’ visual perception. In this study, we assessed how birds perceive colors of four Cepaea nemoralis morphs using physiological models of avian color vision. We calculated combined chromatic and achromatic contrast between shells and three habitat background types as a measure of shell conspicuousness. The degree of background color matching in Cepaea nemoralis depended on both shell morph and habitat type. On average, banded morphs were more conspicuous than unbanded morphs. Morphs were the most cryptic against dry vegetation and the most conspicuous on bare ground. We also found a significant interaction between habitat type and color morph. The relative conspicuousness of shell morphs depended on habitat and was the most variable against green vegetation. Our study provides the first insight into how potential avian predators view Cepaea nemoralis morphs. The results are discussed in light of multiple hypotheses explaining selective predation on Cepaea nemoralis morphs.     

Increase in membrane thickness during development compensates for eggshell thinning due to calcium uptake by the embryo in falcons

We compared membrane thickness of fully developed eggs with those of non-developed eggs in different endangered falcon taxa. To our knowledge, membrane thickness variation during development has never been examined before in falcons or any other wild bird. Yet, the egg membrane constitutes an important protective barrier for the developing embryo. Because eggshell thinning is a general process that occurs during bird development, caused by calcium uptake by the embryo, eggs are expected to be less protected and vulnerable to breakage near the end of development. Thus, egg membranes could play an important protective role in the later stages of development by getting relatively thicker. We used linear mixed models to explore the variation in membrane thickness (n = 378 eggs) in relation to developmental stage, taxon, female age, mass and identity (73 females), egg-laying sequence (105 clutches) and the study zone. Our results are consistent with the prediction that egg membranes are thicker in fully developed eggs than in non-developed eggs, suggesting that the increase in membrane thickness during development may compensate for eggshell thinning. In addition, our data shown that thicker membranes are associated with larger, heavier and relatively wider eggs, as well as with eggs that had thinner eggshells. Egg-laying sequence, female age and the study zone did not explain the observed variation of membrane thickness in the falcon taxa studied. As we provide quantitative data on membrane thickness variation during development in falcons not subjected to contamination or food limitation (i.e. bred under captive conditions), our data may be used as a reference for studies on eggs from natural populations. Considering the large variation in membrane thickness and the multiple factors affecting on it and its importance in the protection of the embryo, we encourage other researchers to include measurements on membranes in studies exploring eggshell thickness variation.     

Body size phenotypes are heritable and mediate fecundity but not fitness in the lepidopteran frugivore Cydia pomonella

The inheritance and functional roles of quantitative traits are central concerns of evolutionary ecology. We report two sets of experiments that investigated the heritability and reproductive consequences of body size phenotypes in a globally distributed lepidopteran frugivore, Cydia pomonella (L.). In our first set of experiments, we tested the hypotheses that (1) body size is heritable and (2) parental body size mediates egg production and offspring survival. Midparent–offspring regression analyses revealed that body mass is highly heritable for females and moderately heritable for males. The contribution of fathers to estimates of additive genetic variance was slightly greater than for mothers. Egg production increased with mean parental size, but offspring survival rates were equivalent. Based on this result, we tested two additional hypotheses in a second set of experiments: (3) male size moderates female egg production and egg fertility and (4) egg production, egg fertility, and offspring survival rate are influenced by female mating opportunities. Females paired with large males produced more eggs and a higher proportion of fertile eggs than females paired with small males. Females with multiple mating opportunities produced more fertile eggs than females paired with a single male. However, egg production and offspring survival rates were unaffected by the number of mating opportunities. Our experiments demonstrate that body mass is heritable in C. pomonella and that size phenotypes may mediate fecundity but not fitness. We conclude that male size can influence egg production and fertility, but female mate choice also plays a role in determining egg fertility.     

No evidence of volatile chemicals regulating reproduction in a multiple queen ant

Efficient cooperation in eusocial insect colonies requires effective communication, and there is abundant evidence of non-volatile chemicals playing a role in regulating reproduction within colonies. In contrast, there have been fewer studies investigating the role of volatile chemicals. This study investigated the potential role of volatile chemicals in regulating queen reproduction either by directly inhibiting queen reproduction or by honestly signalling queen fecundity to workers. We tested this using multiple queen colonies of the ant (Leptothorax acervorum) from a functionally monogynous population where one queen monopolizes all reproduction. Nine colonies, each with an established laying queen, were split to produce two colony fragments—one containing the reproducing queen (group 1) and one containing only previously non-reproducing queens (group 2). Each group was separated by a fine wire mesh preventing physical contact, but allowing volatile chemical contact. In each group 2 fragment, we found that a single formerly non-reproductive queen commenced reproduction and that the rate of egg laying and maximum number of eggs recorded did not significantly differ between groups 1 and 2, results that do not support volatile chemicals as playing a role in regulating queen reproduction. Instead, our findings suggest that physical contact is necessary to maintain functional monogyny.