Sexual dimorphism of egg size in the European Blackbird Turdus merula

Recently, a number of studies have shown that female birds are able to control the sex of their progeny at the stage of the gamete. There is also some evidence that females adjust their investment in offspring depending on the sex of the embryo during egg formation. Differential maternal investment to the eggs depending on their sex is usually interpreted as an adaptive strategy, by which females can increase competitive abilities of the smaller sex, or preferentially invest towards the sex with the potentially higher fitness returns. Here, we studied variation in egg size in relation to embryo sex and laying order in the European Blackbird Turdus merula. We found male and female eggs to differ in size, with larger eggs containing male embryos, as well as a significant interaction between embryo sex and laying order. This interaction resulted from the fact that egg size increased with the laying sequence among eggs bearing females but did not change with laying order among eggs bearing males. There was no relationship between offspring sex and the laying sequence within a clutch. We suggest that sexual dimorphism in egg size recorded in the European Blackbird may reflect favouritism of the sex which may give higher fitness returns.     

Maternal investment during egg laying and offspring sex: an experimental study of zebra finches

To study whether offspring sex is related to the amount of resources invested in eggs we performed an experiment on zebra finches, Taeniopygia guttata. By manipulating their food supply, we forced two groups of females either to increase or to decrease investment in subsequent eggs. Since zebra finches are sexually dimorphic and the reproductive value of the sexes may vary with maternal nutritional status, we predicted that females would adjust the sex of their offspring to egg quality. Females that received poor-quality food for 7 weeks before egg laying, then food supplemented with proteins after they laid the first egg, significantly increased the mass of subsequent eggs. An increase of egg mass with laying sequence was less pronounced in females that received high-quality food before laying and experienced food deterioration after starting to lay. The proportion of sons in subsequent eggs tended to increase in the latter group (although this was marginally significant) but was not related to laying sequence in the other group: these patterns differed significantly between the groups. Offspring sex was not related to egg mass, but newly hatched male chicks were heavier than female chicks. Furthermore, the hatching success of male eggs was lower than that of female eggs. We suggest that differential hatching success of the sexes and sex differences in mass at hatching may constitute important factors shaping brood sex ratios. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Sex allocation in haplodiploids is mediated by egg size: evidence in the spider mite Tetranychus urticae Koch

Haplodiploid species display extraordinary sex ratios. However, a differential investment in male and female offspring might also be achieved by a differential provisioning of eggs, as observed in birds and lizards. We investigated this hypothesis in the haplodiploid spider mite Tetranychus urticae, which displays highly female-biased sex ratios. We show that egg size significantly determines not only larval size, juvenile survival and adult size, but also fertilization probability, as in marine invertebrates with external fertilization, so that female (fertilized) eggs are significantly larger than male (unfertilized) eggs. Moreover, females with on average larger eggs before fertilization produce a more female-biased sex ratio afterwards. Egg size thus mediates sex-specific egg provisioning, sex and offspring sex ratio. Finally, sex-specific egg provisioning has another major consequence: male eggs produced by mated mothers are smaller than male eggs produced by virgins, and this size difference persists in adults. Virgin females might thus have a (male) fitness advantage over mated females.     

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.     

Egg quality and parental ornamentation in the blue tit Parus caeruleus

Components of avian egg quality often show remarkable variation between females and yet the causes of variation in this fitness-related trait remain poorly understood. We investigated the egg size and yolk carotenoid investment of blue tit Parus caeruleus females in relation to multiple parental traits. We show that females produce eggs with more colourful yolks when mated to ultraviolet (UV) attractive males. Since yolk carotenoids may enhance offspring fitness, but may be available to females in limiting amounts, the correlation we found suggests adaptive egg quality adjustment in response to mate attractiveness, but an experimental approach is required to rule out alternative explanations. Yolk colour was also correlated with laying date and yolk mass. Though the effect of laying date could not be explained by ambient temperature during egg formation, it suggests a proximate constraint of general carotenoid availability on yolk composition. Egg size was not affected by male attractiveness; however, females with a brighter crown produced larger eggs, suggesting that crown coloration might indicate individual quality in females too.     

Allocation of offspring size and sex by female black ratsnakes

How females allocate resources to each offspring and how they allocate the sex of their offspring are two powerful potential avenues by which mothers can affect offspring fitness. Previous research has focussed extensively on mean offspring size, with much less attention given to variance in offspring size. Here we focussed on variation in offspring size in black ratsnakes, Elaphe obsoleta . We collected and hatched 105 clutches (1283 eggs) over 9 years. We predicted that females should lay larger eggs, or more variable eggs, when the environment is less predictable. We also predicted that females laying early or laying larger eggs should produce mostly sons because adult males are larger than adult female ratsnakes. The largest hatchling was more than twice the length and almost four times the mass of the smallest hatchling. Variation in offspring size was itself highly variable, with CVs in offspring mass among clutches ranging from 1% to 25%. With one exception, the variables we expected should influence variation in offspring size had little effect. We found that clutch size increased with maternal size and that egg size decreased with clutch size, but we found no evidence that variance in egg size among clutches increased as the season progressed or that females increased the mean size of their offspring the later in the season they laid their eggs. Females in better condition after they finish laying their eggs did produce larger eggs. There was no relationship between within-clutch variation in egg size and laying date or mean egg size. Finally, sex ratio did not vary with mean egg size or hatching date. Given evidence that offspring size in snakes affects survival, selection should reduce variation in offspring size unless that variance enhances maternal fitness and yet we found little support for hypothesized advantages of varying offspring size.     

Maternal effects mediated by egg quality in the Yellow-legged Gull Larus michahellis in relation to laying order and embryo sex

Background

Maternal effects mediated by egg size and quality may profoundly affect offspring development and performance, and mothers may adjust egg traits according to environmental or social influences. In avian species, context-dependency of maternal effects may result in variation in egg composition, as well as in differential patterns of covariation among selected egg components, according to, for example, position in the laying sequence or offspring sex. We investigated variation in major classes of egg yolk components (carotenoids, vitamins and steroid hormones) in relation to egg size, position in the laying sequence and embryo sex in clutches of the Yellow-legged Gull (Larus michahellis). We also investigated their covariation, to highlight mutual adjustments, maternal constraints or trade-offs in egg allocation.

Results

Laying sequence-specific patterns of allocation emerged: concentration of carotenoids and vitamin E decreased, while concentrations of androgens increased. Vitamin A, estradiol and corticosterone did not show any change. There was no evidence of sex-specific allocation or covariation of yolk components. Concentrations of carotenoids and vitamins were positively correlated. Egg mass decreased along the laying sequence, and this decrease was negatively correlated with the mean concentrations of carotenoids in clutches, suggesting that nutritionally constrained females lay low quality clutches in terms of carotenoid content. Finally, clutches with smaller decline in antioxidants between first- and last-laid eggs had a larger increase in yolk corticosterone, suggesting that a smaller antioxidant depletion along the laying sequence may entail a cost for laying females in terms of increased stress levels.

Conclusions

Since some of the analyzed yolk components (e.g. testosterone and lutein) are known to exert sex-specific phenotypic effects on the progeny in this species, the lack of sex-specific egg allocation by mothers may either result from trade-offs between contrasting effects of different egg components on male and female offspring, or indicate that sex-specific traits are controlled primarily by mechanisms of sexual differentiation, including endogenous hormone production or metabolism of exogenous antioxidants, during embryonic development.     

Great tits provided with ad libitum food lay larger eggs when exposed to colder temperatures

The amount of nutrients deposited into a bird egg varies both between and within clutches of the same female. Larger eggs enhance offspring traits, but as a tradeoff, laying large eggs also infers energetic costs to the female. Income breeders usually lay larger eggs later in the season, when temperatures and food availability are higher. Egg size is thus affected by the daily amount of energy available to produce an egg under cold conditions, but it is less well known in how far temperature exerts direct effects on egg size. We show that great tit females Parus major with access to ad libitum food and breeding in climate‐controlled aviaries varied their egg investments. The size of an individual egg was best predicted by mean temperatures one week pre‐laying, with females laying larger, rather than smaller, eggs under colder conditions. Eggs increased in size over the season, but not significantly over the laying sequence. The degree of daily temperature fluctuation did not influence egg size. In addition to a substantial between‐female variation, sisters were more similar to each other than unrelated females, showing that egg size does also reflect heritable intrinsic female properties. Natural variation in egg size is thus not only determined by energy‐limitation, but also due to females allocating more resources to eggs laid in colder environments, thus increasing early survival of the chicks. That the positive correlation between temperature and egg investments that is found in a natural population is reversed under ad libitum food conditions demonstrates that wild great tits tradeoff own condition with survival prospects of their chicks as a function of available food, not ambient temperature.     

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

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

The evolution of sexual size dimorphism in the house finch. V. Maternal effects

The phenotype of a mother and the environment that she provides might differentially affect the phenotypes of her sons and daughters, leading to change in sexual size dimorphism. Whereas these maternal effects should evolve to accommodate the adaptations of both the maternal and offspring generations, the mechanisms by which this is accomplished are rarely known. In birds, females adjust the onset of incubation (coincident with the first egg or after all eggs are laid) in response to the environment during breeding, and thus, indirectly, determine the duration of offspring growth. In the two house finch (Carpodacus mexicanus) populations that breed at the extremes of the species' distribution (Montana and Alabama), females experience highly distinct climatic conditions during nesting. We show that in close association with these conditions, females adjusted jointly the onset of incubation and the sequence in which they produced male and female eggs and consequently modified the growth of sons and daughters. The onset of incubation in newly breeding females closely tracked ambient temperature in a pattern consistent with the maintenance of egg viability. Because of the very different climates in Montana and Alabama, females in these populations showed the opposite patterns of seasonal change in incubation onset and the opposite sex bias in egg-laying order. In females with breeding experience, incubation onset and sex bias in laying order were closely linked regardless of the climatic variation. In nests in which incubation began with the onset of egg laying, the first-laid eggs were mostly females in Montana, but mostly males in Alabama. Because in both populations, male, but not female, embryos grew faster when exposed to longer incubation, the sex-bias produced highly divergent sizes of male and female juveniles between the populations. Overall, the compensatory interaction between the onset of incubation and the sex-biased laying order achieved a compromise between maternal and offspring adaptations and contributed to rapid morphological divergence in sexual dimorphism between populations of the house finch breeding at the climatic extremes of the species range.     

Egg-size variation in the bluethroat (Luscinia s. svecica): constraints and adaptation

We examined inter- and intra-clutch egg-size variation in the bluethroat (Luscinia s. svecica), an open-nesting passerine breeding in the sub-alpine region in southern Norway. By removing first clutches shortly after egg-laying, we induced laying of a repeat clutch. Females significantly reduced the number of eggs from the first to the second nesting attempt, but increased mean egg size. Females in good condition laid significantly larger eggs than those in poor condition. Consistent with predictions of the brood survival hypothesis, assuming an adaptive investment in last eggs to ensure survival of all eggs in the clutch, we found that the size of the last eggs in first clutches was generally larger than the mean egg size of the clutch, and that the relative size of the last egg increased with clutch size. However, a large last egg reflected a general increase in egg size throughout the laying sequence rather than a specific investment in the last egg only. Egg size was not significantly influenced by sex or paternity (within-pair versus extra-pair) of the embryo. In repeat clutches the last egg was not consistently larger than the mean for the clutch. We conclude that female bluethroats face resource limitations during egg formation early in the season, and that the patterns of increase in egg size with laying order for first clutches, and from first to repeat clutches, can largely be explained by proximate constraints on egg formation.     

Against the odds? Nestling sex ratio variation in green-rumped parrotlets

We investigated nestling sex ratio variation in the green-rumpedparrotlet (Forpus passerinus), a small neotropical parrot breedingin central Venezuela. There are strong theoretical reasons topredict a female-biased sex ratio in this system according tothe local resource hypothesis; juvenile males are philopatricand there are high levels of competition between male siblingsfor access to breeding females. Data were collected from twobreeding sites over a 14-year period incorporating 564 broodswith a total of 2728 nestlings. The mean percentage of malenestlings across years was 51%. Despite extreme hatching asynchronyin this system and increased survival of earlier hatched offspring,there was no bias in sex allocation associated with egg sequence.Patterns in sex allocation were not associated with clutch size,age, or size of the breeding female or breeding site. The potentialfor selective resorption of eggs was considered; however, nosignificant relationship was found between extended laying intervalsand the sex of subsequent eggs. Together, these results suggestthat female parrotlets are unable to regulate the sex ratioof their clutch at laying or that facultative manipulation ofnestling sex ratio may not confer a fitness benefit to breedersin these populations.     

Population consequences of maternal effects: sex‐bias in egg‐laying order facilitates divergence in sexual dimorphism between bird populations

Abstract When costs and benefits of raising sons and daughters differ between environments, parents may be selected to modify their investment into male and female offspring. In two recently colonized environments, breeding female house finches (Carpodacus mexicanus) modified the sex and growth of their offspring in relation to the order in which eggs were laid in a clutch. Here we show that, in both populations, these maternal effects strongly biased frequency distribution of tarsus size of fully grown males and females and ultimately produced population divergence in this trait. Although in each population, male and female offspring show a wide range of growth patterns, maternal modifications of sex‐ratio in relation to egg‐laying order resulted in under‐representation of the morphologies that were selected against and over‐representation of morphologies that were favoured by the local selection on juveniles. The result of these maternal adjustments was fast phenotypic change in sexual size dimorphism within and between populations. Maternal manipulations of offspring morphologies may be especially important at the initial stages of population establishment in the novel environments and may have facilitated recent colonization of much of North America by the house finch.     

Temporal variation in reproductive allocation in a shield bug Elasmostethus interstinctus

We investigated changes in the reproductive output and the effect of female phenotype on reproductive parameters in a shield bug Elasmostethus interstinctus (L.) (Heteroptera: Acanthosomatidae) over the whole reproductive period. At the beginning and the middle of the reproductive period eggs were smaller than at the end of the period. Clutch mass and number of eggs per clutch decreased in laying sequence, first clutches being much larger than any of the later ones. Lifetime fecundity correlated positively with female size: large females produced more eggs and lived longer than small ones. Egg size did not vary with female size. Offspring survival until adulthood increased with egg weight. Individuals overwinter before reproduction, and because the nymphs from later-laid eggs have the least time to gather resources before overwintering, it may be important for later-laid eggs to be of high quality. Reproductive allocation varies during the reproductive period; females allocate resources relatively more to offspring number at the beginning of the reproductive period and more to offspring quality at the end of their life.     

Egg size, offspring sex and hatching asynchrony in zebra finches Taeniopygia guttata

Experimental synchronization of onset of incubation was employed in laboratory held zebra finches Taeniopygia guttata to study whether differential resource allocation and possible bias of offspring sex in subsequent eggs in the laying order could mitigate the effects of hatching asynchrony. We found that egg mass increased with laying order, but offspring sex was not related to laying order. Among synchronized clutches, eggs hatched more synchronously than eggs from control nests. Survival probability was related to egg mass, and as expected, this effect differed between experimental groups: it was positive among synchronized broods and not significantly related among asynchronous broods. This suggests that increase in egg mass with the laying order might reduce disparities between early and late hatching chicks. Female nestlings survived better than male nestlings. However their growth was impaired in synchronized broods, whilst growth of males was not affected by hatching synchronization.     

浙江丽水中国石龙子的食性、两性异形和雌性繁殖

丽水分布的中国石龙子（Ｅｕｍｅｃｅｓ ｃｈｉｎｅｎｓｉｓ）摄入的食物均为无脊椎动物,分别属于环节、软体和节肢动物,涉及３０余科,成体和幼体的食物生态位宽度分别为７．２６和６．６９,成体的幼体的食物生态们重叠度为０．５９。性成熟雄性个体大于雌体。成雄和幼体的头长和头宽随体长ＳＶＬ的增长速率大于成雌,成雄头长随ＳＶＬ的增长速度显著大于幼体,成雌和幼体的头长随ＳＶＬ的增长速率无显著差异。成雄头部大于成雌     

Social polyandry, parental investment, sexual selection, and evolution of reduced female gamete size

Abstract Sexual selection in the form of sperm competition is a major explanation for small size of male gametes. Can sexual selection in polyandrous species with reversed sex roles also lead to reduced female gamete size? Comparative studies show that egg size in birds tends to decrease as a lineage evolves social polyandry. Here, a quantitative genetic model predicts that female scrambles over mates lead to evolution of reduced female gamete size. Increased female mating success drives the evolution of smaller eggs, which take less time to produce, until balanced by lowered offspring survival. Mean egg size is usually reduced and polyandry increased by increasing sex ratio (male bias) and maximum possible number of mates. Polyandry also increases with the asynchrony (variance) in female breeding start. Opportunity for sexual selection increases with the maximum number of mates but decreases with increasing sex ratio. It is well known that parental investment can affect sexual selection. The model suggests that the influence is mutual: owing to a coevolutionary feedback loop, sexual selection in females also shapes initial parental investment by reducing egg size. Feedback between sexual selection and parental investment may be common.     

Oviposition behavior and progeny allocation of the polyembryonic waspCopidosoma floridanum (Hymenoptera: Encyrtidae)

Oviposition behavior was used to determine the primary clutch size and sex ratio of the polyembryonic wasp Copidosoma floridanumAshmead (Hymenoptera: Encyrtidae) parasitizing Pseudoplusia includens(Walker) (Lepidoptera: Noctuidae). The laying of a female egg was associated with a pause in abdominal contractions during oviposition, while the laying of a male egg was associated with uninterrupted abdominal contractions. Although unmated females produced only male broods, they also displayed male and female egg oviposition movements. Wasps always laid a primary clutch of one or two eggs. For mated females if only one egg was laid, the emerging secondary clutch was all male or female, but if two eggs were laid a mixed brood of males and females was almost always produced. The secondary clutch of single sex broods was usually between 1000 and 1200 individuals, but the secondary clutch of mixed broods averaged 1143 females and 49 males. Thus, the primary sex ratio for mixed broods was 0.5 (frequency males), but the secondary sex ratio was 0.042. Manipulation of the sequence of male and female egg oviposition or of the primary clutch did not produce major alterations in the secondary clutch size or sex ratio.     

Maternal allocation strategies and differential effects of yolk carotenoids on the phenotype and viability of yellow-legged gull (Larus michahellis) chicks in relation to sex and laying order

Egg quality may mediate maternal allocation strategies according to progeny sex. In vertebrates, carotenoids have important physiological roles during embryonic and post-natal life, but the consequences of variation in yolk carotenoids for offspring phenotype in oviparous species are largely unknown. In yellow-legged gulls, yolk carotenoids did not vary with embryo sex in combination with egg laying date, order and mass. Yolk lutein supplementation enhanced the growth of sons from first eggs but depressed that of sons from last eggs, enhanced survival of daughters late in the season, and promoted immunity of male chicks and chicks from small eggs. Lack of variation in egg carotenoids in relation to sex and egg features, and the contrasting effects of lutein on sons and daughters, do not support the hypothesis of optimal sex-related egg carotenoid allocation. Carotenoids transferred to the eggs may rather result from a trade-off between opposing effects on sons or daughters.     

Patterns of variation in size of Boat-tailed Grackle Quiscalus major eggs

Boat-tailed Grackle Quiscalus major eggs averaged 8^.09 g wet weight. Mean egg weight represented 8 05 % of female body weight. Based on egg weight, Grackle eggs have a shorter incubation period than the general passerine pattern.
Egg weight varied significantly with sequence of laying and between clutches in both two-egg and three-egg clutches. Last laid eggs weighed less than first laid eggs. The mean weight of the first two eggs laid in three-egg clutches did not differ from the mean egg weight of two-egg clutches. The average clutch of two and three eggs weighed 16.33 g and 24.16 g, respectively.
Mean egg weight varied between study colonies and with season. Grackles at East Lake laid eggs that weighed less than grackles at either Alligator Lake or North Lake. Eggs laid during March averaged less than eggs laid later in the season. The locality variation reflects the different timing of nesting between sites rather than food abundance. As nutrient provisioning increases with an increase in egg weight, the seasonal change in egg weight probably reflects improved feeding conditions for the female. This suggests that selection favours beginning laying before reserves of the female are sufficient to lay the largest egg possible.
The size of the young at hatching was correlated with egg size. Newly hate had young averaged 79.6% of fresh egg weight. Egg weight did not determine the probability of hatching or starvation of the young. Females do not appear to adjust egg size facultatively depending on the sex of the young. Eggs producing males and females did not vary significantly in weight, whether comparisons were made within or between clutches.