Intraclutch Variation Research Articles

Evaluating the potential importance of individual identity, maternal traits, and environment as predictors of egg characteristics in walleye Sander vitreus

Individual and environmental factors may influence gamete characteristics and contributions to recruitment in fishes. We tested for the influence of maternal, abiotic, and biotic factors on egg diameter and quality (i.e., oil droplet diameter) for walleye Sander vitreus in Escanaba Lake, Wisconsin, during 2018–2023 (omitting 2020). Analyses were conducted on fish captured once and for the same individuals captured multiple times during our study period. In the single-capture analysis, increasing maternal length was weakly related to larger egg diameter and oil droplet size. Increased yellow perch abundance was somewhat related to increased intra-clutch variation, whereas a later ice-off date was related to reduced intra-clutch variation. Fish that spawned later had more variable egg and oil droplet diameters. Among fish that were sampled over multiple years, individual identity was a strong predictor of egg and oil diameter. Our results suggest that regulations that preserve among individual variation in egg traits could increase the chances that environmental conditions will be favorable for spawning and recruitment for at least some fish.

Female red-winged blackbirds (Agelaius phoeniceus) do not alter nest site selection, maternal programming, or hormone-mediated maternal effects in response to perceived nest predation or brood parasitism risk

Predation or brood parasitism risks can change the behaviors and reproductive decisions in many parental animals. For oviparous species, mothers can mitigate their reproductive success in at least three ways: (1) by avoiding nest sites with high predation or parasitism risks, (2) through hormonal maternal effects that developmentally prime offspring for survival in risky environments, or (3) by investing less in reproduction when predation or parasitism risks are high. Here, we tested if perceived predation and parasitism risks can induce any of these behavioral or physiological responses by exposing female red-winged blackbirds (Agelaius phoeniceus) to playbacks of two major nest threats, a predator (Cooper’s hawk, Accipiter cooperii) and an obligate brood parasite (brown-headed cowbird; Molothrus ater), as well as two controls (harmless Eastern meadowlark, Sturnella magna; and silence). We found that female blackbirds did not avoid nesting at sites treated with predator or brood parasite playbacks, nor were females more likely to abandon nesting attempts at these sites. Egg size and yolk hormone profiles, which are common proxies for maternal investment in oviparous species, were statistically similar across treatment sites. Instead, we found intraclutch variation in yolk steroid hormone profiles: concentrations of three progestogens (pregnanedione, 17α-hydroxypregnenolone, and deoxycorticosterone) and two androgens (testosterone and androstenedione) were higher in third-laid than first-laid eggs. Our study largely confirms previous findings of consistent intraclutch yolk hormone variation in this species, in birds in general, and in other oviparous lineages, but uniquely reports on several yolk steroid hormones largely overlooked in the literature on hormone-mediated maternal effects.

Empirical evidence of coevolution between the channel-billed cuckoo and its host, the pied currawong

Abstract Avian obligate brood parasites and their hosts exhibit coevolution, where parasites respond to the evolution of host defences with counter-adaptations, such as egg mimicry, which can then lead to the evolution of more efficient host defences. Using objective measurements and avian visual modelling, we quantified the degree of similarity between the eggs of the channel-billed cuckoo (Scythrops novaehollandiae) and the eggs of three host species from the hosts’ visual perspective. We found no evidence that this cuckoo species has host-specific egg types, but it does appear to mimic one of its primary hosts, the pied currawong (Strepera graculina), in both colour and pattern. Currawongs showed significantly higher interclutch variation in colour and luminance than the other two hosts and had significantly lower intraclutch variation in pattern than interclutch variation. High interclutch and low intraclutch variation in a host could make it more difficult for a cuckoo to match closely any particular host clutch, potentially allowing a host to detect a parasitic egg more easily. This is the first empirical study using objective measurements to quantify the degree of egg mimicry in the channel-billed cuckoo and the results suggest a potential host defence to egg mimicry that merits further investigation.

High within-clutch repeatability of eggshell phenotype in Barn Swallows despite less maculated last-laid eggs

Abstract Ecological and life-history variation and both interspecific and intraspecific brood parasitism contribute to diversity in egg phenotype within the same species. In this study, Barn Swallows (Hirundo rustica erythrogaster) laid eggs with high intraclutch repeatability in egg size, shape, and maculation. Despite this high intraclutch repeatability, last-laid eggs had consistently less of the eggshell covered in spots and fewer spots than earlier-laid eggs in the clutch. We examined sources of interclutch and intraclutch variation using both direct measurements and custom software (SpotEgg, NaturePatternMatch) that provide detailed information on egg characteristics, especially maculation measures. In addition to our main findings, maculation on different sides of the egg was highly repeatable; however, only shape, proportion of the eggshell maculated, and average spot size were repeatable between first and replacement clutches. Low intraclutch variation in maculation could allow females to recognize their clutch and this may be adaptive for colonial nesting species, such as the Barn Swallow. Characterizing intraspecific variation in egg size, shape, and maculation is the first step in understanding whether intraclutch variation is low enough—and interclutch variation high enough—such that eggs could serve as identity signals.

Rejection of Conspecific Eggs Associated with the Appearance of Eggs, in the Azure-Winged Magpie Cyanopica cyanus in Korea

Avian brood parasitism is a reproductive strategy in which a parasitic species lays its eggs in the nests of a host. In response to parasitism, hosts have evolved various defence strategies. One of the host defences against intraspecific and interspecific brood parasitism is to distinguish parasitic eggs based on intraclutch variation in egg appearance. Hosts can improve their ability to discriminate parasitic eggs by lowering the degree of their intraclutch variation in egg appearance. Therefore, intraclutch variation may affect the egg rejection behaviour of hosts. In this study, we conducted experiments to study egg recognition and rejection behavior for mimetic eggs in Azure-winged Magpies Cyanopica cyanus in Korea. We also investigated the level of variation in egg appearance (egg color, spottiness and volume) in order to explore whether the degree of this variation is associated with the rejection decision. We found that there were no differences in egg coloration or spottiness between the rejecter and accepter. However, we found that intraclutch egg volume variation in rejecters was significantly lower than that in accepters, indicating that the Azure-winged Magpie is likely to recognize its own eggs by lowering its intraclutch variation in egg volume as a defensive strategy against brood parasitism. This study suggests that the Azure-winged Magpie use not the differences in egg coloration but egg volume to discriminate and reject closely mimetic conspecific eggs.

The Role of Intra-Clutch Variation of Magpie Clutches in Foreign Egg Rejection Depends on the Egg Trait Considered

The existence of a coevolutionary process between avian brood parasites and their hosts predicts a lower intra-clutch variation in egg appearance of host eggs among rejecters as this would favor egg discrimination of parasite eggs by hosts once parasitic egg mimicry had evolved. So far empirical tests of this prediction have ignored the fact that different aspects of host egg phenotypes may differ in the relative role of environmental vs. genetic determination, and hence that the role of intra-clutch variation in egg rejection within a population cannot be invariant. Here, we estimated whether the intra-clutch variation in several aspects of host eggshell features is consistently associated to rejection of parasitic foreign eggs across years in a magpie host population parasitized by great spotted cuckoos. We innovatively estimated spottiness by means of the fractal dimension of eggs, which considers the homogeneity of spot pattern complexity in eggshells. Our results show that low intra-clutch variation in the blue-green coloration at the middle area of the eggs associated with a high chance of rejection, but only in one of the 3 years we conducted the study. In addition, females that rejected foreign eggs presented more homogenous spot patterns in their clutches as estimated by their fractal dimension than females that accepted experimental eggs, independently of the year of study. Finally, intra-clutch variation in egg volume of host eggs was not associated to rejection. Analyses at the individual level revealed that the relative role of genetic vs. environmental factors that determine egg phenotype would be feature-specific in magpies, females having a characteristic spottiness, but not color or volume, pattern. Our work stresses the importance of considering a holistic approach including several aspects of variation in host egg phenotype (size, color, and homogeneity of spot pattern), as some aspects might be more susceptible to selection through egg rejection than others, presumably because they are less influenced by variation in the environmental conditions. Moreover, our study highlights the importance of replication in studies on the adaptive value of host traits in egg rejection.

Persistent pollutants in fresh and abandoned eggs of Common Tern (Sterna hirundo) and Arctic Tern (Sterna paradisaea) in Ireland

Higher levels of persistent pollutants (Σ16PCB, Σ6PBDE, ΣHCH, ΣDDT, ΣCHL) were detected in fresh eggs of Common Terns Sterna hirundo from Rockabill Island near Dublin (Ireland's industrialised capital city) compared to Common and Arctic Terns S. paradisaea from Ireland's west coast. Intra-clutch variation of pollutant levels in Common Terns was shown to be low, providing further evidence that random sampling of one egg may be an appropriate sampling strategy. Significant differences in pollutant concentrations were detected between fresh and abandoned eggs on Rockabill. However, abandoned eggs can still provide a useful approximation of pollutants in bird eggs if non-destructive sampling is preferred. Levels of p,p’ –DDE in tern eggs have decreased over time according to this study, in concurrence with worldwide trends. Results in this study fall below toxicological thresholds for birds and OSPARs EcoQO thresholds set for Common Tern eggs, except for mercury and HCH in the west coast.

Condition dependent strategies of egg size variation in the Common Eider Somateria mollissima.

We analysed intraclutch egg-size variation over the laying sequence in relation to clutch size, and the relation between clutch size and female body condition, in the Common Eider Somateria mollissima during an 8-year period. The aim was to assess if eiders adjusted egg size within the laying sequence depending on clutch sizes in response to body condition, as such an adjustment could have adaptive implications on reproductive success through a size advantage for the hatchlings. The analyses were performed on a population level; and then at the individual level using data from recaptured females that changed clutch size between years. Based on 1,099 clutches from 812 individual females, population clutch size averaged 4.13 eggs (range: 1-6), with 4- and 5-egg clutchesconstituting c.70% of all clutches, taking turns in being the most represented clutch size. Clutch size was positively related to female pre-laying body condition at both the population and individual levels. Egg size varied significantly within and between clutch sizes and changes were significantly related to the laying sequence. First eggs were significantly larger in 4-egg clutches and second eggs marginally smaller than in 5-egg clutches, a pattern also found among individual females changing clutch size between years. The relationship between female pre-laying body condition and clutch size, and the intraclutch egg-size pattern indicate that both clutch size and egg size are actively adapted to the pre-breeding body condition of the female. As egg size potentially optimise reproductive success through a size advantage in hatchlings, the observed pattern of intraclutch egg-size variation suggests that female eiders possesses a finely tuned conditional dependent mechanism that may optimize reproductive output in years were females are in suboptimal body condition for breeding.

Egg investment strategies adopted by a desertic passerine, the Saxaul Sparrow (Passer ammodendri)

BackgroundAs one of the reproductive strategies adopted by bird species, variation in investment in egg production and its influencing factors are important and well-studied subjects. Intraclutch changes in egg size associated with laying order may reflect a strategy of “brood survival” or “brood reduction” adopted by female birds in different situations.MethodsWe conducted field studies on the breeding parameters of the Saxaul Sparrow (Passer ammodendri) in Gansu Province, China from 2010 to 2017, to clarify the factors affecting the egg investment and reproductive performance of this passerine species.ResultsOur results revealed significant differences in clutch size, egg size and the fledging rate between the first and second brood of Saxaul Sparrows and suggested that this typical desert species allocates more breeding resources to the more favourable second brood period, leading to greater reproductive output. Female body size presented a positive relationship with egg size, and male body size presented positive relationships with clutch size and hatchability. The females that started their clutches later laid more eggs, and hatchability and the fledging rate also increased with a later laying date in the first brood period. With successive eggs laid within the 5-egg clutches (the most frequent clutch size), egg size increased for the first three eggs and then significantly decreased.ConclusionsOur results indicate that female Saxaul Sparrows increased egg investment because of good quality of paired males and good environmental conditions. The intraclutch variation of egg size suggests that this species inhabiting an arid environment adopts a “brood reduction” strategy.

Maternal provisioning by foam-nesting frogs of the genus Physalaemus (Anura, Leptodactylidae) in contrasting environments

In some environments prevailing conditions are unpredictable, thus anuran species use bet-hedging strategies and produce eggs of varied sizes. We investigated whether four species of Physalaemus (two from open, two from forest habitats) exhibit bet-hedging strategies, and if intra-clutch variation in egg size is greater for species that breed in the more unpredictable ponds of open habitats. All species studied adopted the bet-hedging strategy, with intra-clutch variation in egg size regardless of the environment; however, we found greater intra-clutch variation in the two species from open areas. The lower variation in egg-size found within forest habitats may be explained by the more stable environments that forest ponds provide for anuran eggs/embryos. Future studies in a phylogenetic context are needed to confirm the patterns detected here.

Variation in organochlorine and mercury levels in first and replacement eggs of a single-egg clutch breeder, the thick-billed murre, at a breeding colony in the Canadian Arctic

Contaminant concentrations may vary among sequentially-laid eggs in multi-egg clutches, and this variation has implications for the interpretation of contaminant concentrations in monitoring programs. The thick-billed murre (Uria lomvia) is a key species for monitoring contaminants in the Canadian Arctic and lays only a single egg per year. Therefore, the potential issue of intra-clutch variation in contaminant concentrations is avoided. However, if the egg is removed or lost early in the incubation stage, the adult female murre will relay. In this study, we examined contaminant concentrations and patterns in first-laid and replacement eggs of thick-billed murres breeding in northern Hudson Bay in order to determine whether or not these eggs could be sampled interchangeably. Concentrations of the major legacy organochlorines (e.g. PCBs, DDT, chlordanes) were generally higher, and total mercury concentrations lower, in the replacement eggs compared with the first-laid eggs. The organochlorine profile was comprised primarily of ΣDDT and Σ70PCB, and Σ70PCB was comprised primarily of hexa-hepta PCBs in both first-laid and replacement eggs. As both concentrations and organochlorine patterns showed differences between first-laid and replacement eggs, we recommend that randomly selected first-laid eggs of thick-billed murres be consistently sampled for contaminant monitoring in the Canadian Arctic.

The anti-predator role of within-nest emergence synchrony in sea turtle hatchlings.

Group formation is a common behaviour among prey species. In egg-laying animals, despite the various factors that promote intra-clutch variation leading to asynchronous hatching and emergence from nests, synchronous hatching and emergence occurs in many taxa. This synchrony may be adaptive by reducing predation risk, but few data are available in any natural system, even for iconic examples of the anti-predator function of group formation. Here, we show for the first time that increased group size (number of hatchlings emerging together from a nest) reduces green turtle (Chelonia mydas) hatchling predation. This effect was only observed earlier in the night when predation pressure was greatest, indicated by the greatest predator abundance and a small proportion of predators preoccupied with consuming captured prey. Further analysis revealed that the effect of time of day was due to the number of hatchlings already killed in an evening; this, along with the apparent lack of other anti-predatory mechanisms for grouping, suggests that synchronous emergence from a nest appears to swamp predators, resulting in an attack abatement effect. Using a system with relatively pristine conditions for turtle hatchlings and their predators provides a more realistic environmental context within which intra-nest synchronous emergence has evolved.

Egg-laying sequence influences egg mercury concentrations and egg size in three bird species: Implications for contaminant monitoring programs.

Bird eggs are commonly used in contaminant monitoring programs and toxicological risk assessments, but intraclutch variation and sampling methodology could influence interpretability. The authors examined the influence of egg-laying sequence on egg mercury concentrations and burdens in American avocets, black-necked stilts, and Forster's terns. The average decline in mercury concentrations between the first and last eggs laid was 33% for stilts, 22% for terns, and 11% for avocets, and most of this decline occurred between the first and second eggs laid (24% for stilts, 18% for terns, and 9% for avocets). Trends in egg size with egg-laying order were inconsistent among species, and overall differences in egg volume, mass, length, and width were <3%. The authors summarized the literature, and among 17 species studied, mercury concentrations generally declined by 16% between the first and second eggs laid. Despite the strong effect of egg-laying sequence, most of the variance in egg mercury concentrations still occurred among clutches (75-91%) rather than within clutches (9%-25%). Using simulations, the authors determined that accurate estimation of a population's mean egg mercury concentration using only a single random egg from a subset of nests would require sampling >60 nests to represent a large population (10% accuracy) or ≥14 nests to represent a small colony that contained <100 nests (20% accuracy). Environ Toxicol Chem 2016;35:1458-1469. Published 2015 Wiley Periodicals Inc. on behalf of SETAC. This article is a US Government work and, as such, is in the public domain in the United States of America.

Egg trait variation in a large hawk-cuckoo (Hierococcyx sparverioides) host population of Chinese babax (Babax lanceolatus).

Mutual interaction between brood parasites and their hosts is a well-known model system for studying host-parasite coevolution. Both parties have acted reciprocally, resembling an evolutionary arms race, in which adaptations and counter-adaptations have evolved as a result of host-parasite dynamics, such as the classical cuckoo-host system. Discrimination among parasite and cuckoo eggs and rejection of foreign eggs is regarded as an important anti-parasitism strategy. The Chinese babax (Babax lanceolatus) is a large hawk-cuckoo (Hierococcyx sparverioides) host distributed in southwest China. A previous study shows that the babax is an intermediate egg rejector, and most cuckoo eggs are accepted by the Chinese babax, although a small proportion of hosts reject cuckoo eggs. Interestingly, the large hawk-cuckoo lays non-mimetic eggs in contrast to the uniform blue eggs of babaxes. Because egg coloration is a critical cue used by host species in favor of the recognition of parasitic eggs by hosts, we used a spectrometer to quantify egg color variation to understand the differentiation in discrimination ability between the egg rejectors and acceptors. We found that the chroma of intra-clutch variation of babax eggs was more consistent in egg rejectors than in acceptors. However, no statistical significance was found in inter-clutch variation between these two types of hosts. Our results suggest that hosts lay eggs with a low level of intra-clutch variation without the necessity of a high level of inter-clutch variation simultaneously as predicted by the egg signature hypothesis. This study may further indicate that selection pressures from evolutionarily recent parasites can drive individual-based differences in an anti-parasitism strategy.

Experimental shifts in intraclutch egg color variation do not affect egg rejection in a host of a non-egg-mimetic avian brood parasite.

Avian brood parasites lay their eggs in the nests of other birds, and impose the costs associated with rearing parasitic young onto these hosts. Many hosts of brood parasites defend against parasitism by removing foreign eggs from the nest. In systems where parasitic eggs mimic host eggs in coloration and patterning, extensive intraclutch variation in egg appearances may impair the host’s ability to recognize and reject parasitic eggs, but experimental investigation of this effect has produced conflicting results. The cognitive mechanism by which hosts recognize parasitic eggs may vary across brood parasite hosts, and this may explain variation in experimental outcome across studies investigating egg rejection in hosts of egg-mimicking brood parasites. In contrast, for hosts of non-egg-mimetic parasites, intraclutch egg color variation is not predicted to co-vary with foreign egg rejection, irrespective of cognitive mechanism. Here we tested for effects of intraclutch egg color variation in a host of nonmimetic brood parasite by manipulating egg color in American robins (Turdus migratorius), hosts of brown-headed cowbirds (Molothrus ater). We recorded robins’ behavioral responses to simulated cowbird parasitism in nests where color variation was artificially enhanced or reduced. We also quantified egg color variation within and between unmanipulated robin clutches as perceived by robins themselves using spectrophotometric measures and avian visual modeling. In unmanipulated nests, egg color varied more between than within robin clutches. As predicted, however, manipulation of color variation did not affect rejection rates. Overall, our results best support the scenario wherein egg rejection is the outcome of selective pressure by a nonmimetic brood parasite, because robins are efficient rejecters of foreign eggs, irrespective of the color variation within their own clutch.

Variation in the Size of Eggs of Kelp Gulls (Larus dominicarius) at Two Colonies in Patagonia, Argentina

Although several studies have reported basic information about the size of Kelp Gull (Larus dominicarius) eggs, no attention has been paid to the comparison of patterns of intra-clutch variation at different breeding sites. Here, the variation in egg size of Kelp Gulls at two colonies from Patagonia, Argentina, was analyzed. The objectives were to characterize egg size variation in Kelp Gulls breeding at a recently established colony in Punta Loma that primarily relies on natural prey; and to evaluate if patterns of intra-clutch variation in that colony differ from those at Vernaci Sudoeste Island, a larger and well-established colony with access to supplementary food in the form of fishery waste. In two-egg clutches, first-eggs were larger than second-eggs. Neither average egg size nor intra-clutch patterns differed between study sites. Also, in two-egg clutches, egg size decreased as date of laying increased, probably due to younger birds laying smaller eggs later in the season. The pattern of intra-clutch variation in three-egg clutches differed between sites. Three-egg clutches at Vernaci Sudoeste Island presented similar sizes for first- and second-eggs and these were larger than third-eggs. This pattern contrasted with the steady decrease in relation to laying order that was observed at Punta Loma. Also, second-and third-eggs from Vernaci Sudoeste Island were larger than corresponding eggs from Punta Loma. Our data suggest that differences in egg size between colonies could be the consequence of enhanced food availability at Vernaci Sudoeste Island in the form of discards generated by trawl fisheries. Future studies to evaluate age of birds and inter-annual environmental stochasticity are needed to elucidate the effect of these factors on egg size variation.

Effects of intraclutch variation in egg size and hatching asynchrony on nestling development and survival in semi-precocial Herring Gulls

Intraclutch egg size variation may non-adaptively result from nutritional/energetic constraints acting on laying females or may reflect adaptive differential investment in offspring in relation to laying/hatching order. This variation may contribute to size hierarchies among siblings already established due to hatching asynchrony, and resultant competitive asymmetries often lead to starvation of the weakest nestling within a brood. The costs in terms of chick mortality can be high. However, the extent to which this mortality is egg size-mediated remains unclear, especially in relation to hatching asynchrony which may operate concomitantly. I assessed effects of egg size and hatching asynchrony on nestling development and survival of Herring Gulls (Larus argentatus), where the smaller size and later hatching of c-eggs may represent a brood-reduction strategy. To analyze variation in egg size, I recorded the laying order and laying date of 870 eggs in 290 three-egg clutches over a 3-yr period (2010-2012). I measured hatchlings and monitored growth and survival of 130 chicks from enclosed nests in 2011 and 2012. The negative effect of laying date ( =− 0.18 ± SE 0.06, P = 0.002) on c-egg size possibly reflected the fact that late breeders were either low quality or inexperienced females. The mass, size, and condition of hatchling Herring Gulls were positively related to egg size (all P < 0.0001). C-chicks suffered from increased mortality risk during the first 12 d, identified as the brood-reduction period in my study population. Although intraclutch variation in egg size was not directly related to patterns of chick mortality, I found that smaller relative egg size interactively increased differences in relative body condition of nestlings, primarily brought about by the degree of hatching asynchrony during this brood-reduction period. Thus, the value of relatively small c-eggs in Herring Gulls may lie in reinforcing brood reduction through effects on nestling body condition. A reproductive strategy Herring Gulls might have adopted to maintain a three-egg clutch, but that also enables them to adjust the number of chicks they rear relative to the prevailing environmental conditions and to their own condition during the nestling stage.

Variation in concentrations of organochlorines and brominated flame retardants among eggs in abandoned clutches of a terrestrial raptor

Bird eggs are often used to measure various pollutants in monitoring programs. In protected species, however, it might only be possible to collect eggs abandoned by the parent birds. In order to assess the appropriateness of abandoned eggs for contaminant monitoring we examined intra-clutch variation in concentrations of 24 halogenated organic pollutants (16 organochlorines [OCs] and 8 brominated flame retardants) in six abandoned tawny owl (Strix aluco) clutches from central Norway. The variation among eggs within a clutch was low for nearly all compounds (intra-class correlation >0.9 both for lipid- and wet weight). This study suggests that abandoned eggs provide a relatively reliable measurement of lipophilic pollutants in clutches of terrestrial raptors such as the tawny owl.

Intraclutch variation in egg appearance of Brown-headed Cowbird hosts

ABSTRACT Several species of avian brood parasites have evolved egg mimicry, which can interfere with host egg rejection. Parasitic egg mimicry may select for decreased intraclutch variation in host egg appearance to facilitate the recognition and rejection of parasitic eggs. This hypothesis has received scant attention in hosts of the Brown-headed Cowbird (Molothrus ater) because the cowbird apparently has not evolved mimetic eggs. Nonetheless, hosts with eggs similar in appearance to cowbird eggs should minimize intraclutch variation to increase the likelihood of detecting parasitism. By contrast, there may be minimal selection pressure to reduce intraclutch variation in hosts with eggs that are divergent from cowbird eggs. Using reflectance spectrometry, we compared the intraclutch variation between accepters and rejecters of 2 groups of host species: those with eggs similar in appearance to cowbird eggs (white maculate eggs) and those with eggs that clearly diverge in appearance from cowbird eggs (blue...

Laying‐sequence variation in yolk carotenoids and egg characteristics in the red‐winged blackbird Agelaius phoeniceus

In many bird species with asynchronous hatching, smaller, later‐hatched nestlings are out‐competed for food by their larger, earlier‐hatched siblings and therefore suffer increased mortality via starvation. It is thought that female birds can either maintain or reduce the survival disadvantage of later‐hatched nestlings by differentially allocating maternal resources across the eggs of a clutch. Carotenoid pigments are an example of resources that female birds allocate differentially when producing a clutch, but laying sequence patterns for these pigments remain poorly studied in North American songbirds. We examined intraclutch variation in yolk carotenoids and egg metrics in 27 full clutches of red‐winged blackbird Agelaius phoeniceus eggs collected from eight wetlands in central Alberta, Canada. We predicted that carotenoids would decrease across the laying sequence, as in this species, later‐hatched, marginal nestlings suffer greater mortality than earlier‐hatched, core nestlings. We found nine carotenoid pigments in red‐winged blackbird egg yolks, including two that have never been described from avian yolks: α‐doradexanthin and adonirubin. As predicted, concentrations and amounts of most carotenoids decreased across the laying sequence, suggesting that female red‐winged blackbirds depleted their carotenoid resources as they laid more eggs. However, egg mass and yolk mass both increased across the laying sequence, suggesting that female red‐winged blackbirds may use other maternal resources to compensate for the size and survival disadvantage experienced by later‐hatched, marginal nestlings.