Differential effects of egg albumen content on barn swallow nestlings in relation to hatch order

In diverse animal taxa, egg mass variation mediates maternal effects with long-term consequences for offspring ontogeny and fitness. Patterns of egg mass variation with laying order differ considerably among birds, but no study has experimentally investigated the function of variation in albumen or yolk egg content in the wild. In barn swallows (Hirundo rustica), absolute and relative albumen mass increased with egg laying order. Experimental albumen removal delayed hatching, had larger negative effects on growth of late-hatched nestlings, and reduced nestling survival. Laying order positively predicted hatch order. Because nestling competitive ability depends on size, and albumen egg content influences hatchling size, present results suggest that by increasing albumen content of late eggs mothers reduce hatching asynchrony and enhance growth particularly of late-hatched nestlings. Thus, variation in albumen mass with laying order may function to mitigate the negative phenotypic consequences of hatching late in species that adopt a 'brood-survival' strategy.     

Experimental manipulation of food availability leads to short‐term intra‐clutch adjustment in egg mass but not in yolk androgen or thyroid hormones

In birds, mothers can affect their offspring's phenotype and thereby survival via egg composition. It is not well known to what extent and time‐scales environmental variation in resource availability, either via resource constrains or adaptive adjustment to predicted rearing conditions, influences maternal effects. We experimentally studied whether egg and yolk mass and yolk hormone levels respond to short‐term changes in food availability during laying in wild great tits Parus major. Our treatment groups were: 1) food supplementation (mealworms) from the 1st until the last egg; 2) food supplementation from the 1st until the 5th egg, where the effect of cessation of the supplementary food treatment could also be studied; 3) no food supplementation (controls). We analysed both nutritional resources (egg, yolk and albumen mass), and the important developmental signals, yolk androgens (testosterone and androstenedione), and for the first time in a wild population, yolk thyroid hormones (thyroxine and 3,5,3′‐triiodothyronine). Egg mass is a costly resource for females, androgens most likely non‐costly signals, whereas thyroid hormones may be costly signals, requiring environmental iodine. In the food supplemented group egg, yolk and albumen mass increased rapidly relative to controls and when food supplementation was halted, egg and albumen mass decreased, indicating rapid responses to resource availability. Yolk androgen and thyroid hormone levels were not affected by food supplementation during laying. Thyroxine showed an increase over the laying sequence and its biological meaning needs further study. The rapid changes in egg mass to variation in within‐clutch food availability suggest energetic/protein/nutrient constrains on egg formation. The lack of a response in yolk hormones suggest that perhaps in this species the short‐term changes in resource availability during egg laying do not predict offspring rearing conditions, or (for thyroid hormones) do not cause systemic changes in circulating hormones, and hence do not affect maternal signaling.     

Yolk testosterone and estradiol variation relative to clutch size, laying order and hatching asynchrony in Common Grackles

Maternal effects are typically thought to enhance rather than reduce offspring performance, but asynchronous hatching (ASH) in birds typically produces a size hierarchy within a clutch that frequently reduces the growth and survival of nestlings from eggs that hatch later. Given that yolk steroids can significantly affect offspring phenotype and that in many species the levels of yolk steroids have been found to increase with laying order, the maternal transfer of steroids to egg yolk has been proposed as a mechanism for females to offset the deleterious effects of ASH. To test this hypothesis, we determined whether yolk steroids varied with laying order or clutch size in Common Grackles (Quiscalus quiscula). Because ASH varies with clutch size (hatching span averages 48 h in five-egg clutches, 24 h in four-egg clutches) and regularly results in the starvation of later hatched nestlings, we predicted: (1) testosterone and 17?-estradiol levels should increase with laying order in both clutch sizes to mitigate the negative effects of ASH on last-hatched nestlings, and (2) the increase should be greater in five-egg clutches due to more pronounced hatching asynchrony. Using a competitive-binding steroid radioimmunoassay, we found no systematic variation in either testosterone or estradiol levels relative to laying order or clutch size. In the absence of evidence that yolk steroids interact adaptively with ASH, research must look elsewhere for potential benefits that might compensate for the costs these steroids impose on nestlings.     

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 (β = ?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.     

Maternal androgens in the pied flycatcher: timing of breeding and within-female consistency

Maternal hormones can have substantial phenotypic effects in the progeny of many vertebrates. It has been proposed that mothers adaptively adjust hormone levels experienced by particular young to optimize their reproductive output. In birds, systematic variation in egg hormone levels has been related to different female reproductive strategies. Because in many bird species prospects of the offspring change seasonally and with brood number, strategic adjustment of yolk androgen levels would be expected. To test this idea, we induced pied flycatcher (Ficedula hypoleuca) females to nest twice during the same season by removing their first clutches shortly after clutch completion. We collected eggs of first and replacement clutches to measure yolk concentrations of androstenedione (A4) and testosterone (T) and captured the females that laid these clutches for phenotypic measurements. Although average egg androgen levels were remarkably consistent within females, hormone patterns differed considerably between first and replacement clutches. Eggs of replacement clutches were heavier with larger yolks compared to first clutches, but they contained on average lower levels of androgens. Within clutches, androgen concentration increased over the laying sequence in the first clutch, but decreased or remained more constant over the laying sequence in the replacement clutch. Mean yolk T, but not A4 levels, were negatively associated with laying date for both breeding attempts. Moreover, females in good body condition produced eggs containing lower levels of androgens than females in poor condition. Our results are consistent with the idea that differences in yolk androgen levels may be one mechanism underlying seasonal variation in reproductive success and it is possible that changes in egg androgen patterns may reflect a change in female reproductive strategy. High within-female consistency also highlights the possibility that there may be some underlying genetic variation in yolk androgen levels.     

Discrepancy between factors affecting nestling growth and survival and maternal success in Common Grackles

ABSTRACT Multiple factors potentially affect nestling survival and maternal reproductive success. However, little is known about the relative importance of different factors when operating simultaneously or whether the same factors are important for nestlings and their mothers. We determined the effect of hatching asynchrony, individual egg size, mean egg size, nestling sex, and clutch initiation date on the survival of individual nestlings and on maternal reproductive success in Common Grackles (Quiscalus quiscula) from 2004 to 2006 in central Illinois. Factors most important to maternal success differed from those important for individual nestling growth and survival. Hatching asynchrony had the greatest within‐nest influence on the fate of nestlings; the earlier a nestling hatched relative to siblings, the greater its mass and likelihood of fledging. Clutch size had the greatest influence on maternal reproductive success, with females with larger clutches fledging more young. Thus, both nestling survival and maternal success were largely determined by a single, albeit different, factor. A possible explanation for the apparent unimportance of most factors we measured in determining maternal success is that we did not consider variation among females. Individual variation in maternal attributes such as condition, size, age, experience, or mate quality may result in females tailoring clutch attributes (i.e., egg size, sex, and degree of hatching asynchrony) in ways that allow them to maximize their reproductive success. The discordance between factors that benefited mothers versus their offspring illustrates the importance of considering the maternal consequences of any factor that appears to affect offspring survival. Factors that increase the mass and survival of some offspring may not result in increased maternal reproductive success.     

Brood Reduction via Intra-clutch Variation in Testosterone - An Experimental Test in the Great Tit

In birds, yolk androgen concentrations in eggs can increase or decrease over the laying sequence and common hypotheses hold that this serves to favour the competitive ability of either first- or last-hatched chicks depending on the prevailing conditions, and thus promote brood reduction or maintenance of original brood size respectively. Intra-clutch variation of testosterone can shift relative competitive ability of siblings and hence competitive dynamics. In a natural population of great tits, we experimentally investigated the effects and function of maternal testosterone on offspring phenotype in relation to the laying position of the egg in a context of hatching asynchrony. To this end, we created three types of clutches where either the first three or the last three eggs of a clutch were injected with testosterone (T) dissolved in sesame oil, and the remaining eggs with sesame oil only, or where all eggs of a clutch were injected with sesame oil. Increased levels of yolk T in the last-laid eggs resulted in the last-hatched chicks being significantly lighter and smaller than their siblings, while increased levels of T in the first-laid eggs had no direct effect on the first-hatched chicks, but an indirect negative effect on their siblings. Our results suggest that females can potentially adjust offspring phenotype by modulating, over the laying sequence, the amounts of T deposited in the eggs. These results are in contradiction, however, with current hypotheses and previous findings, which suggest that under good conditions higher levels of maternally derived T in the last-laid eggs should mitigate the negative effects of hatching asynchrony.     

Hatching asynchrony can have long‐term consequences for offspring fitness in zebra finches under captive conditions

Hatching asynchrony can have profound short‐term consequences for offspring, although the long‐term consequences are less well understood. The purpose of this study was to examine the long‐term consequences of hatching asynchrony for offspring fitness in birds. Specifically, we aimed to test the hypothesis that hatching asynchrony increases the sexual attractiveness and fecundity, respectively, of early‐hatched male and female zebra finch, Taeniopygia guttata (Vieillot, 1817) offspring. Mate‐choice trials comparing male nestlings with the same parents, but that were reared in asynchronous or experimentally synchronous broods, revealed no female preference in relation to hatching regime. We did however find strong evidence that, as adults, late‐hatched males were more attractive to females than siblings that had hatched earlier. Meanwhile, we found a weak trend towards early‐hatched females depositing more carotenoids and retinol in the egg yolk than late‐hatched or synchronously hatched females, although there were no differences in terms of clutch characteristics or the deposition of α‐tocopherol or γ‐tocopherol in the egg yolk. Therefore, we found that the beneficial long‐term consequences of hatching asynchrony were sex specific, being accrued by late‐hatched male nestlings and by early‐hatched female nestlings. Consequently, we conclude that the long‐term consequences of hatching asynchrony are more complex than previously realised. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 430–438.     

Egg temperature and initial brood patch area determine hatching asynchrony in Magellanic penguin Spheniscus magellanicus

In birds, the adaptive significance of hatching asynchrony has been under debate for many years and the parental effects on hatching asynchrony have been largely assumed but not often tested. Some authors suggest that hatching asynchrony depends on the incubation onset and many factors have been shown to influence hatching asynchrony in different species. Our objective was to analyze the exact timing of the onset of incubation and if this affects hatching asynchrony; and, in addition, which other factors (brood patch development, incubation position, adult body condition, intra‐clutch egg dimorphism, laying date and year) affect hatching asynchrony in Magellanic penguins Spheniscus magellanicus. We first estimated the eggshell temperature at which embryo development starts, with a non‐destructive and novel method. We then recorded individual egg temperatures in 61 nests during incubation, and related them, and other breeding parameters, to hatching asynchrony. We also observed incubation positions in 307 nests. We found a significant positive relationship between hatching asynchrony and the temperature that the first‐laid egg experienced during egg laying and between hatching asynchrony and the initial brood patch area. We also found a negative relationship between hatching asynchrony and the difference in temperature between second and first‐laid eggs within a clutch, measured after the egg‐laying period was finished. We ruled out position of the eggs during incubation, adult body condition, egg volume, laying date, and study year as factors influencing hatching asynchrony. The egg temperature during laying and the difference in temperature between eggs of a clutch are determinants of hatching asynchrony in Magellanic penguins.     

Maternal androgens in black-headed gull (Larus ridibundus) eggs: consequences for chick development

We tested the hypothesis that mother birds counterbalance the negative effects of hatching asynchrony for later-hatched chicks by increasing the yolk androgen concentrations in consecutive eggs of their clutch. In doing so, they may adaptively tune each offspring's competitive ability and, thus, growth and survival. However, evidence in support of this hypothesis is contradictory. The yolk concentrations of maternal androgens in the eggs of black-headed gulls increase significantly with the laying order of the eggs in a clutch. We experimentally tested the functional consequences of this increase on chick development under natural conditions by injecting eggs with either an oil or androgen solution. We created experimental clutches in which androgen levels either stayed constant or increased with laying order while controlling for differences in egg quality by using only first-laid eggs. We then compared development, growth and survival between these broods. Androgen treatment enhanced embryonic development because androgen-treated eggs hatched half a day earlier than controls, while their size at hatching was similar to oil-treated controls. Androgen treatment did not increase chick survival, but it enhanced growth. Androgen-treated, third-hatched chicks had a higher body mass and longer legs than third-hatched chicks that hatched from oil-treated eggs. At the same time, growth of first chicks (which were all oil treated) was reduced by the presence of two androgen-treated siblings, suggesting that yolk androgens enhance the competitive ability of later-hatched chicks. Our results support the hypothesis that transfer of different amounts of androgens to the eggs of a clutch is a mechanism by which mothers maximize their reproductive output.     

Do female spotless starlings Sturnus unicolor adjust maternal investment according to male attractiveness?

In birds, female egg allocation patterns have a strong influence in offspring development and differential investment in egg size and composition has been shown to respond to male attractiveness. In this study we experimentally manipulated the perceived attractiveness of male starlings Sturnus unicolor by increasing the amount of green material in some nests (a male courtship display in this species). We predicted that, if female investment before laying is related to male attractiveness, experimental females would increase their reproductive investment in response to the addition of plants in their nests when compared to control females. We found that our manipulation caused variations in female reproductive investment in a way that seems to influence offspring quantity but not offspring quality: Females laid larger clutch sizes but not larger eggs when green plant material was added. However, yolk androgens contents were not related to the experimental manipulation. Contrary to expectations, females breeding in experimental nests laid eggs with smaller amounts of eggshell pigments. Interestingly, we found that eggs laid later in the sequence had higher testosterone levels and showed more intense egg colouration than eggs laid earlier in the sequence. These differences at the within-clutch level suggest that selection has favoured compensatory strategies for hatching asynchrony. Alternatively, since nest sabotages by other females are most common at the beginning of laying, this could be seen as female strategy to minimise losses due to nest sabotages. As far as we know, this is the first study to show that an external egg characteristic such as blue-green colouration reflects yolk androgen concentration.     

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.     

Influence of mate preference and laying order on maternal allocation in a monogamous parrot species with extreme hatching asynchrony

It is well established that in many avian species, prenatal maternal resource allocation varies both between and within clutches and may affect offspring fitness. Differential allocation of maternal resources, in terms of egg weight and yolk composition, may therefore allow the female to adjust brood reduction and to fine-tune reproductive investment in accordance with the expected fitness returns. The adaptive value of such maternal resource allocation is thought to be context-dependent as well as species-specific. We investigated the effects of female preference for her mate on the allocation of prenatal maternal resources in the budgerigar, Melopsittacus undulatus, a monogamous species of parrot that shows an extreme hatching asynchrony. We assessed mate preferences in a two-way preference test and allowed females two breeding rounds: one with the preferred and one with the non-preferred partner. We found no effect of preference on either latency to lay or clutch size, but females mated with the preferred partner laid eggs that contained significantly more yolk. Their eggs also contained significantly more androstenedione but not testosterone. Our results suggest that in this species, female preference may influence maternal resource allocation, and that the functional roles of each androgen in the yolk should be considered separately. In addition, we found a significant effect of laying order on egg and yolk weight as well as on yolk testosterone and androstenedione levels. These measures, however, did not change linearly with the laying order and render it unlikely that female budgerigars compensate for the extreme hatching asynchrony by adjusting within-clutch allocation of prenatal maternal resources.     

Egg development time in the Zebra Finch Taeniopygia guttata varies with laying order and clutch size

As female birds are able to lay no more than a single egg each day, in those species producing larger clutches the first laid eggs may get a developmental head‐start over later eggs in the clutch. All other things being equal, the differential pattern of development across the clutch may contribute to hatching asynchrony and subsequent inequity in the competition between brood mates, and ultimately increase variance in the quality and fitness of first‐ and last‐laid offspring. It has been suggested that females might allocate resources differently across the laying sequence to moderate the developmental rate and hatching time of different embryos. We tested this theory in the Zebra Finch Taeniopygia guttata, a common model species for investigating maternal effects in birds. We removed 758 eggs from 160 nests shortly after they were laid and used artificial incubators to control for parental effects and monitor hatching times. Eggs from larger clutches consistently hatched sooner than those from average‐sized clutches, demonstrating that the intrinsic properties of an egg can alter the developmental time of embryos. There were also differences in the development time of eggs across the laying sequence, but these patterns were weaker, inconsistent and unrelated to sequential investment across the laying sequence in a straightforward way. This study indicates that maternal resource allocation to eggs across the laying sequence and across clutch sizes can influence development times and play a potentially important role in determining the competitive dynamics of broods.     

Differential effects of offspring and maternal inbreeding on egg laying and offspring performance in the burying beetle Nicrophorus vespilloides

We investigate the effect of offspring and maternal inbreeding on maternal and offspring traits associated with early offspring fitness in the burying beetle Nicrophorus vespilloides. We conducted two experiments. In the first experiment, we manipulated maternal inbreeding only (keeping offspring outbred) by generating mothers that were outbred, moderately inbred or highly inbred. Meanwhile, in the second experiment, we manipulated offspring inbreeding only (keeping females outbred) by generating offspring that were outbred, moderately inbred or highly inbred. In both experiments, we monitored subsequent effects on breeding success (number of larvae), maternal traits (clutch size, delay until laying, laying skew, laying spread and egg size) and offspring traits (hatching success, larval survival, duration of larval development and average larval mass). Maternal inbreeding reduced breeding success, and this effect was mediated through lower hatching success and greater larval mortality. Furthermore, inbred mothers produced clutches where egg laying was less skewed towards the early part of laying than outbred females. This reduction in the skew in egg laying is beneficial for larval survival, suggesting that inbred females adjusted their laying patterns facultatively, thereby partially compensating for the detrimental effects of maternal inbreeding on offspring. Finally, we found evidence of a nonlinear effect of offspring inbreeding coefficient on number of larvae dispersing. Offspring inbreeding affected larval survival and larval development time but also unexpectedly affected maternal traits (clutch size and delay until laying), suggesting that females adjust clutch size and the delay until laying in response to being related to their mate.     

Female variation in allocation of steroid hormones,antioxidants and fatty acids: a multilevel analysis in a wild passerine bird

The environment where an embryo develops can be influenced by components of maternal origin, which can shape offspring phenotypes and therefore maternal fitness. In birds that produce more than one egg per clutch, females differ in the concentration of components they allocate into the yolk along the laying sequence. However, identification of processes that shape female yolk allocation and thus offspring phenotype still remains a major challenge within evolutionary ecology. A way to increase our understanding is by acknowledging that allocation patterns can differ depending on the level of analysis, such as the population versus the among‐female (within‐population) level. We employed mixed models to analyze at both levels the variation in allocation along the laying sequence of four steroid hormones, three antioxidants, and four groups of fatty acids present in the egg yolks of wild great tits Parus major. We also quantified repeatabilities for each component to study female consistency. At a population level, the concentrations/proportions of five yolk components varied along the laying sequence, implying that the developmental environment is different for offspring developing in first versus last eggs. Females varied substantially in the mean allocation of components and in their plasticity along the laying sequence. For most components, these two parameters were negatively correlated. Females were also remarkably repeatable in their allocation. Overall, our data emphasize the need to account for female variation in yolk allocation along the laying sequence at multiple levels, as variation at a population level is underpinned by different individual patterns. Our findings also highlight the importance of considering both levels of analysis in future studies investigating the causes and fitness consequences of yolk compounds. Finally, our results on female repeatability confirm that analyzing one egg per nest is a suitable way to address the consequences of yolk resource deposition for the offspring.     

Pre-breeding diet affects the allocation of yolk hormones in zebra finches Taeniopygia guttata

The ability of mothers to modify offspring phenotype to match prevailing environmental conditions is an important component of reproductive success, especially in variable environments. Pre-breeding conditions, such as food abundance, may have significant consequences for both the number and quality of offspring a female produces as well as her ability to rear the offspring. In an experiment where pre-breeding diet was manipulated, we investigated if allocation of yolk androgens (testosterone and 5α-dihydrotestosterone) was related to the quality of diet experienced prior to breeding. Female zebra finches Taeniopygia guttata produced larger clutches on high quality diet than on low quality diet but with no differences in egg mass. Yolk androgen levels were repeatable within subsequent clutches of the same female and females did not change mean androgen content in eggs in relation to diet quality. However, within-in clutch pattern of yolk testosterone and DHT changed with diet treatment. Testosterone and DHT decreased with laying order on the low quality diet but remained constant on the high quality diet. Differential yolk androgen allocation within the clutch may alter the competitive differences between chicks and provide females the possibility to adjust reproductive investment and offspring phenotype already at the egg stage.     

Maternal compensation for hatching asynchrony in the collared flycatcher Ficedula albicollis

Several hypotheses have been proposed to explain the adaptiveness of hatching asynchrony for the parents, but delayed hatching is generally detrimental for the late hatched young. These offspring often experience competitive disadvantage and delayed development. If hatching asynchrony has a reason other than producing competitive differences among offspring, it would be advantageous, not only for the offspring but even for the parents, to compensate for its detrimental effects. In some species, increasing investment into later laid eggs has been reported and discussed as a compensation mechanism, but its effect on nestling growth and fledging size has not been examined in details. In this study we investigated nestling growth and size at fledging in terms of body mass and length of primaries in relation to the accurate laying and hatching order in collared flycatcher Ficedula albicollis broods. We found that females laid larger eggs at the end of the laying sequence, and this helped to decrease the disadvantages for the last offspring. The last offspring had lower body mass growth rate and fledged with shorter feathers, but in both cases the larger the last egg was, the smaller the lag of the offspring was. We conclude, that even if females were not able to fully compensate for the detrimental effects of hatching asynchrony, larger eggs may improve the survival prospects of late hatched nestlings.     

Should males come first? The relationship between offspring hatching order and sex in the black‐headed gull Larus ridibundus

In birds with hatching asynchrony and sexual size dimorphism, chicks hatched earlier and later in the laying sequence usually suffer different mortalities due to uneven abilities to compete for food, especially in poor years. If sexes differ in vulnerability to environmental conditions, e.g., by having different food requirements due to differential growth rates, mothers can increase fitness by allocating sex according to the laying order, producing less vulnerable sex later rather than early in the clutch. By analysing variation in primary sex ratio using a PCR-based DNA technique, we tested this prediction in black-headed gull Larus ridibundus chicks where males may be the less viable sex under adverse conditions. The overall primary sex ratio of the population did not depart from parity. However, first hatched chicks were more likely to be males whereas last hatched chicks were more likely to be females. Both egg volume and hatchling body mass decreased with laying order irrespective of sex. Time of breeding had no effect on offspring sex or hatchling sex ratios.     

Maternal resource variation across the laying sequence in Canada geese Branta canadensis maxima

Variation in maternal investments to offspring presumably reflects an optimization of resource allocation such that a female's fitness is maximized. In birds, both egg size and yolk constituents are examples of resources that can vary among offspring within a clutch. Egg size and maternally-derived steroid hormone concentrations present in yolk have been characterized for many species that lay small clutches or have altricial young, but little information is available for species that lay moderate to large clutches of precocial young. In this study, we recorded laying position, measured fresh egg mass and determined maternally-derived testosterone and estradiol concentrations present in yolks for whole clutches of free-living Canada geese Branta canadensis maxima to assess variation in maternal resources within clutches. We found that egg size varied non-linearly across the laying sequence such that first laid eggs were small, the largest eggs in the clutch occurred in the second and third positions, and size declined in eggs laid in subsequent positions. Concentration of testosterone in the yolk followed a pattern in which the first and second laid eggs have the highest concentrations within a clutch and declining concentrations in subsequently laid eggs. In contrast, maternally-derived yolk estradiol concentrations (measured in a subset of clutches) did not change across the laying sequence.