Does the differential cost of sons and daughters lead to sex ratio adjustment in great frigatebirds Fregataminor?

Sex allocation theory predicts that if benefits of producing sons and daughters differ and outweigh the costs of sex ratio adjustment, parents should produce more of the offspring that provide them with greater fitness. Potential benefits may be more likely to outweigh costs where sexual size dimorphism and, in birds, single‐egg clutches exist. Great frigatebirds Fregataminor are seabirds in which females are larger than males and clutch size is one egg. In our study population, sexual size dimorphism develops primarily during the period of complete juvenile dependence on parental care, consistent with a higher cost of producing daughters than sons. Over the course of the 1998 breeding season there was a shift from early season prevalence of daughters to late‐season prevalence of sons. Variation in food availability at time of egg laying, as indexed by sea surface temperature (SST), was a strong predictor of offspring sex in 1998. In contrast, SST in 2003 was not a predictor of offspring sex, nor was there a seasonal shift in the hatching sex ratio, despite a seasonal shift in SST. Besides food availability, we tested two additional factors in 2003 that could explain sex ratio adjustment in relation to the cost of reproduction. Offspring sex in 2003 was not related to natural or experimentally induced variation in maternal body condition; pre‐laying food supplements raised the body condition of females at the time of egg laying but did not affect offspring sex or egg mass. In addition, offspring sex was not predicted by the length of maternal telomere restriction fragments (TRFs), an index of age and possibly of reproductive experience. Broad confidence intervals on effect size suggest that undetected effects of maternal condition on offspring sex ratio could easily exist, but confidence intervals were narrower on the non‐significant effects of SST and TRF length on offspring sex ratio. The cause of different seasonal patterns of hatching sex ratio and different SST effects in 1998 and 2003 is unclear.     

Seasonal change in offspring sex and size in Dawson's burrowing bees (Amegilla dawsoni) (Hymenoptera: Anthophorini)

Abstract.  1. Nesting females of Dawson's burrowing bees, Amegilla dawsoni , produce a large size class of offspring, which includes daughters and major sons, and a small size class, which consists entirely of minor sons averaging half the weight of their larger siblings. Female allocation patterns change over the flight season such that the initial pattern of producing daughters shifts toward the production of both daughters and major sons in the middle of the season, and then the production of primarily minor sons in the latter part of the nesting season.
2. In Dawson's burrowing bees, this pattern is correlated with declines in pollen and nectar availability as the nesting season progresses as well as a heightened risk of dying before the final brood cell is completed. Here, the relation between these factors and the provisioning tactics of nesting Dawson's burrowing bees is discussed.     

Seasonal variation in sex ratio and sexual egg dimorphism favouring daughters in first clutches of the spotless starling

We investigated possible pre‐hatching mechanisms of sex‐differential investment by females that may contribute to offspring sex‐ratio adjustment enhancing the fitness return from reproductive effort in the spotless starling (Sturnus unicolor). We found a seasonal shift in sex ratio from daughters to sons as the season advances. Furthermore, the probability of breeding at 1‐year old and recruitment into the breeding population in daughters is associated with laying date but not with mass at fledging. The reverse is true for males which rarely bred at 1‐year old. We also found that eggs containing female embryos are significantly heavier than those containing males in spite of the slight sexual dimorphism in favour of males. This suggests maternal control of provisioning, favouring daughters that may balance sibling mortality and competition with their brothers. Our results on seasonal variation in sex ratio and differential egg provisioning are consistent with an adaptive tactic in which mothers increase their reproductive return by enhancing the probability that daughters survive and breed in their first year of life.     

Maternal investment in relation to sex ratio and offspring number in a small mammal – a case for Trivers and Willard theory?

1.  Optimal parental sex allocation depends on the balance between the costs of investing into sons vs. daughters and the benefits calculated as fitness returns. The outcome of this equation varies with the life history of the species, as well as the state of the individual and the quality of the environment.
2.  We studied maternal allocation and subsequent fecundity costs of bank voles, Myodes glareolus , by manipulating both the postnatal sex ratio (all-male/all-female litters) and the quality of rearing environment (through manipulation of litter size by −2/+2 pups) of their offspring in a laboratory setting.
3.  We found that mothers clearly biased their allocation to female rather than male offspring regardless of their own body condition. Male pups had a significantly lower growth rate than female pups, so that at weaning, males from enlarged litters were the smallest. Mothers produced more milk for female litters and also defended them more intensively than male offspring.
4.  The results agree with the predictions based on the bank vole life history: there will be selection for greater investment in daughters rather than sons, as a larger size seems to be more influencial for female reproductive success in this species. Our finding could be a general rule in highly polygynous, but weakly dimorphic small mammals where females are territorial.
5.  The results disagree with the narrow sense Trivers & Willard hypothesis, which states that in polygynous mammals that show higher variation in male than in female reproductive success, high-quality mothers are expected to invest more in sons than in daughters.     

Adaptive sex allocation in relation to hatching synchrony and offspring quality in house wrens

Increased variance in the reproductive success of males relative to females favors mothers that optimally allocate sons and daughters to maximize their fitness return. In altricial songbirds, one influence on the fitness prospects of offspring arises through the order in which nestlings hatch from their eggs, which affects individual mass and size before nest leaving. In house wrens (Troglodytes aedon), the influence of hatching order depends on the degree of hatching synchrony, with greater variation in nestling mass and size within broods hatching asynchronously than in those hatching synchronously. Early-hatching nestlings in asynchronous broods were heavier and larger than their later-hatching siblings and nestlings in synchronous broods. The effect of hatching order was also sex specific, as the mass of males in asynchronous broods was more strongly influenced by hatching order than the mass of females, with increased variation in the mass of males relative to that of females. As predicted, mothers hatching their eggs asynchronously biased first-laid, first-hatching eggs toward sons and late-laid, late-hatching eggs toward daughters, whereas females hatching their eggs synchronously distributed the sexes randomly among the eggs of their clutch. We conclude that females allocate the sex of their offspring among the eggs of their clutch in a manner that maximizes their own fitness.     

Laying order and offspring sex in blue tits Parus caeruleus

According to sex allocation theory, females may benefit from a differential investment in sons and daughters, where fitness returns from male and female offspring vary. One way in which investment may be differentiated is by assigning male and female offspring to eggs of different sizes, and/or eggs laid early or late in the laying order. Here, we investigate whether such sex-specific adjustments occur in blue tits Parus caeruleus . We found that the proportion of males changed non-linearly with laying order showing the increase in early-laid eggs. Egg size increased with laying order, but this pattern differed between female and male eggs. Female eggs increased initially in size and became smaller later in the laying sequence, while male eggs exhibited constant increase in mass. However, egg size did not differ between genders. Egg size and laying order was found to interact in moulding sex of the embryo. Thus, females seem to simultaneously adjust the sex of the offspring in relation to their investments into eggs and the position of the egg in the laying sequence.     

Sex-biased environmental sensitivity: natural and experimental evidence from a bird species with larger females

The larger sex is often more vulnerable, in terms of developmentand survival, to poor conditions during early life. Differentialvulnerability has implications for parental investment strategiessuch as sex ratio theory. When males are larger, it is not possibleto separate the effects of larger size per se and other aspectsof the male phenotype on vulnerability. Furthermore, offspringcompetition might favor the larger sex and thereby mask intrinsic,size-related effects. We studied sex-specific mortality in abird species with reversed size dimorphism, the great skua Stercorariusskua, under natural and experimentally created poor conditions.Small eggs from extended laying sequences were used to createpoor early conditions for the offspring, which were raised assingletons. Daughters had a lower survival in all treatmentgroups. Survival in natural broods was additionally affectedby hatch date and position. Hatch weight was not different forsons and daughters but was lower in experimental than in naturalnests. In natural nests, daughters fledged 10% heavier thansons, but in experimental nests, they did not reach a highermass. The average survival difference between sons and daughterswas not increased in experimental broods. However, hatch weighthad a strong sex-specific effect. Very light females never survived,and survival probability of daughters increased with increasinghatch weight. By contrast, survival of sons over the same rangeof hatch weights was not related to weight. These findings supportthe hypothesis that larger (final) size per se is related tosex-specific offspring vulnerability during early life.     

Sex allocation in yellow-legged gulls (Larus michahellis) depends on nutritional constraints on production of large last eggs

Male and female offspring can differ in their susceptibility to pre-natal (e.g. egg quality) and post-natal (e.g. sib–sib competition) conditions, and parents can therefore increase their individual fitness by adjusting these maternal effects according to offspring sex. In birds, egg mass and laying/hatching order are the main determinants of offspring viability, but these effects can act differently on each sex. In a previous study, relatively large last-laid (c-)eggs of yellow-legged gulls (Larus michahellis) were more likely to carry a female embryo. This suggests compensatory allocation of maternal resources to daughters from c-eggs, which suffer reduced viability. In the present study, we supplemented yellow-legged gulls with food during the laying period to experimentally test whether their nutritional conditions were responsible for the observed covariation between c-egg sex and mass. As predicted, food supplementation enhanced female c-eggs'' mass more than that of male c-eggs. Thus, this experiment indicates that mothers strategically allocated their resources to c-eggs, possibly in order to compensate for the larger susceptibility of daughters to hatching (and laying) order. The results also suggested that mothers decided on resource allocation depending on the sex of already ovulated c-eggs, rather than ovulating ova of either sex depending on food availability.     

Hatching sex ratio and sex specific chick mortality in common terns Sterna hirundo

Bias in sex ratios at hatching and sex specific post hatching mortality in size dimorphic species has been frequently detected, and is usually skewed towards the production and survival of the smaller sex. Since common terns Sterna hirundo show a limited sexual size dimorphism, with males being only about 1–6% larger than females in a few measurements, we would expect to find small or no differences in production and survival of sons and daughters. To test this prediction, we carried out a 2-year observational study on sex ratio variation in common terns at hatching and on sex specific post hatching mortality. Sons and daughters hatched from eggs of similar volume. Post hatching mortality was heavily influenced by hatching sequence. In addition, we detected a sex specific mortality bias towards sons. Overall, hatching sex ratio and sex specific mortality resulted in fledging sex ratios 8% biased towards females. Thus, other reasons than body size may be influencing the costs of rearing sons. Son mortality was not homogeneous between brood sizes, but greater for two-chick broods. Since adults rearing two-chick broods were younger, lighter and bred consistently later than those rearing three-chick broods, it is suggested that lower capacity of two-chick brood parents adversely affected offspring survival of sons. Though not significantly, two-chick broods tended to be female biased at hatching, perhaps to counteract the greater male-biased nestling mortality. Thus, population bias in secondary sex ratio is not limited to strongly size dimorphic species, but species with a slight sexual size dimorphism can also show sex ratio bias through a combination of differential production and mortality of sons and daughters.     

Sex-specific hatching order, growth rates and fledging success in jackdaws Corvus monedula

In the jackdaw Corvus monedula , eggs hatch asynchronously with the youngest chicks in the brood often starving to death. So far, it is unknown whether there are sex differences in vulnerability to starvation. Adult females are smaller than males suggesting that daughters should be cheaper to produce than sons and so, less likely to starve when nest conditions are poor. Here, we determine whether sex, laying order and season interact to influence growth and fledging success. In a nestbox population of jackdaws, we found a non-significant female bias at both hatching (112:120) and fledging (37:52). Generalised linear models revealed that parents seemed to be investing differently in sons and daughters depending on their chances of success. Broods produced late in the season were significantly female biased, particularly those from small clutches. Females hatched towards the end of the season, when conditions were poor, were more likely to fledge than males. Nestlings that were relatively large at hatching were more likely to fledge. This effect was particularly important for last hatched individuals. Overall, males had a higher mortality rate than females. The most likely cause was starvation due to higher energetic requirements, because males were larger than females at fledging. We suggest that in species with brood reduction, sex-biased mortality may be at least as important as primary sex ratio manipulation in determining avian sex ratios.     

COURTSHIP FEEDING AND THE FITNESS OF FEMALE KATYDIDS (ORTHOPTERA: TETTIGONIIDAE)

Katydid males (Requena verticalis) produce spermatophores with a large sperm-free spermatophylax, which is eaten by the female after mating. In this study, I asked 1) how do spermatophylax nutrients affect the fitness of the mated female and her progeny? 2) does this male-produced food substitute for other food in the diet of the female, or is it a source of specialized nutrients? and 3) does an increase in the size of the spermatophylax eaten influence female reproduction in the same way as the additional spermatophylaxes that would be obtained from multiple mating? An experimental increase either in dietary protein or the number of spermatophylaxes eaten increased the number of eggs produced. However, spermatophylax size had no effect on egg number. An increase in either the size or number of spermatophylaxes eaten resulted in an increase in egg size. There was no influence of protein in the general diet on egg size. This suggests that males provide nutrition not available from other sources. Although there were no direct effects of number of spermatophylaxes eaten by females on the overwintering survival of their progeny, offspring from females producing larger eggs had a relatively higher probability of surviving winter. The amount of spermatophylax eaten had no influence on the mean adult size of progeny but significantly increased the mean date at which sons matured to adulthood. There was no influence of dietary protein on these variables. Since maturation date is positively correlated with adult size in both sexes, it is suggested that the influence of courtship feeding on maturation date may result in an increase in adult size and thus the fitness of sons. A significant correlation between the size of the female and the mean size of her sons (but not daughters) suggests that there is also a heritable component to body size.     

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.     

Ural owl sex allocation and parental investment under poor food conditions

Parents are expected to overproduce the less costly sex under poor food conditions. The previously regular 3-year cycle in the abundance of voles, the main prey of the Ural owl, Strix uralensis, temporarily disappeared in 1999–2001. We studied Ural owls' parental feeding investment and sex allocation during these poor-quality years. We sexed hatchlings and embryos in unhatched eggs of all 131 broods produced during these years. Population wide, the owls produced significantly more males (56%). The parental food investment in the brood was estimated by sorting out the prey remains in the bottom of nest boxes. Food delivered to 83 broods without chick mortality showed no clear sex-specific investment. Nestling mortality was equal in both sexes. Thus, evidence for an investment-driven sex allocation is weak. Neither laying date, brood size nor the female's condition correlated with offspring sex ratios. In these poor years, parents provided less food per chick and the fledging weight of daughters was reduced more than the weight of sons compared with years of high food abundance (1983 and 1986). We discuss, in relation to published studies, the possibility of a sex-allocation scenario where, under poor food conditions, a daughter's long-term fitness is reduced more than a son's.     

Facultative sex allocation in snow skink lizards (Niveoscincus microlepidotus)

Mathematical models suggest that reproducing females may benefit by facultatively adjusting their relative investment into sons vs. daughters, in response to population‐wide shifts in operational sex ratio (OSR). Our field studies on viviparous alpine skinks (Niveoscincus microlepidotus) document such a case, whereby among‐ and within‐year shifts in OSR were followed by shifts in sex allocation. When adult males were relatively scarce, females produced male‐biased litters and larger sons than daughters. The reverse was true when adult males were relatively more common. That is, females that were courted and mated by few males produced mainly sons (and these were larger than daughters), whereas females that were courted and mated by many males produced mainly daughters (and these were larger than sons). Maternal body size and condition also covaried with sex allocation, and the shifting pattern of sexual size dimorphism at birth may reflect these correlated effects rather than a discrete component of an evolved sex‐allocation strategy.     

FACTORS AFFECTING THE EGG SIZE OF RED-BILLED GULLS LARUS NOVAEHOLLANDIAE SCOPULINUS

The factors influencing the egg size of the Red-billed Gull Larus novaehollandiae scopulinus were studied at Kaikoura, New Zealand, between 1964 and 1972. In two- and three-egg clutches there was a trend for the eggs to become smaller in the sequence of laying. Length, breadth and volume of eggs of one-, two- and three-egg clutches declined significantly as the season progressed. The size of eggs from single-egg clutches tended to be smaller than eggs from two-egg clutches laid at the same time. There were correlations between the proportions of one-egg and of three-egg clutches being laid at a given period and the mean egg volume of two-egg clutches. When the mean egg volume of two-egg clutches increased there was a corresponding increase in the proportion of two- and three-egg clutches laid. When the mean egg volume of two-egg clutches decreased there was an increase in the proportion of single-egg clutches laid. The egg size of the Red-billed Gull showed no direct correlation with the abundance or availability of food; the largest eggs were produced early in the season when food was in short supply. In spite of an increase in the food supply in the middle of the breeding season, birds laying at this time produced smaller eggs than birds which laid earlier in the season. However, early breeders which relayed at the peak in food abundance on average produced a larger replacement clutch than originals laid early in the season. It is suggested that the birds nesting early in the season are able to produce the largest eggs because they are the most efficient foragers for food, and those which nest later in the season produce smaller eggs, even at peak food abundance, because of their inefficiency or inexperience. Early breeders laying replacement clutches tended to lay larger eggs and larger clutches than birds which are producing their first clutches at the same time. Two-year-old females laid eggs which were significantly shorter than older aged birds while the breadth and volume of the egg increased with the age of the female up to the fifth year. There was a trend for females to lay larger eggs when mated with older rather than younger males. No statistical differences in egg size were detected between females changing or retaining the partner of the previous season. Female body weight and egg volume were positively correlated in females weighing less than 275 g but not for heavier females. It is suggested that the seasonal decline in egg size and clutch size results from a decrease in the availability of food and the ability of the individual to exploit the resource.     

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.     

Parents’ genetic dissimilarity and offspring sex in a polygynous mammal

Offspring quality may benefit from genetic dissimilarity between parents. However, genetic dissimilarity may trade‐off with additive genetic benefits. We hypothesized that when sexual selection produces sex‐specific selective scenarios, the relative benefits of additive genetic vs. dissimilarity may differ for sons and daughters. Here we study a sample of 666 red deer (Cervus elaphus) microsatellite genotypes, including males, females and their foetuses, from 20 wild populations in Spain (the main analyses are based on 241 different foetuses and 190 mother‐foetus pairs). We found that parental lineages were more dissimilar in daughters than in sons. On average, every mother was less related to her mate than to the sample of fathers in the population when producing daughters not sons. Male foetuses conceived early in the rutting season were much more inbred than any other foetuses. These differences maintained through gestation length, ruling out intrauterine mortality as a cause for the results, and indicating that the potential mechanism producing the association between parents’ dissimilarity and offspring sex should operate close to mating or conception time. Our findings highlight the relevance of considering the sex of offspring when studying genetic similarity between parents.     

Azure‐winged Magpies Cyanopica cyanus trade off reproductive success and parental care by establishing a size hierarchy among nestlings

It is common in birds that the sizes of nestlings vary greatly when multiple young are produced in one nest. However, the methods used by parents to establish size hierarchy among nestlings and their effect on parental provisioning pattern may differ between species. In the Azure‐winged Magpie Cyanopica cyanus, we explored how and why parents controlled the sizes of nestlings. Asynchronous hatching was the main cause of size hierarchy within the brood, although the laying of larger eggs later in the laying sequence reduced this effect. Parents with asynchronous broods produced more eggs and fledged more nestlings than those with synchronous broods but their brood provisioning rates, food delivery per feeding bout and feeding efficiency did not differ. We performed a cross‐fostering experiment to synchronize some asynchronous broods. Provisioning rates of asynchronous broods were lower than those of synchronized broods, but the daily growth rates and fledging body mass of their nestlings were not different. Our findings indicate that parents of asynchronous broods can achieve higher reproductive success than those of synchronous broods based on the same parental care, and the same reproductive success as those of synchronized broods based on less parental care. It appears that parent birds can better trade off reproductive success and parental care by establishing a size hierarchy among nestlings.     

Offspring are lighter at birth and smaller in adulthood when born after a brother versus a sister in humans

In mammals, including humans, it is more costly to produce sons than it is to produce daughters, with maternal survival and subsequent reproductive success diminished more by producing male over female offspring. It is therefore predicted that offspring who are produced by mothers who have previously produced sons versus daughters will be compromised by the relatively high cost their mother incurred in the previous reproductive episode. Such effects are potentially important because characters that determine offspring survival and fecundity ultimately contribute to maternal fitness. Using questionnaire-based data from a contemporary human population, I show that birthweight (irrespective of their sex) is lower in individuals born after an elder brother than in those born after an elder sister. In addition, I show that both men and women who were born after a male versus a female sibling have reduced adulthood height, a known correlate of reproductive success in both sexes. The results suggest that producing sons may have a negative effect on the fitness of subsequent offspring, which has implications for calculations of maternal fitness and for optimal sex allocation.     

Consequences of sex-specific growth on sibling competition in black-headed gulls: a sexually-size dimorphic species with scramble competition

Biased mortality of the larger sex during the early developmental period has been reported for a number of size-dimorphic bird species. This can partly be explained by the fact that growing to larger size renders the larger sex more vulnerable to food shortage. However, since sibling rivalry is often size-dependent, chicks of the larger sex should have a competitive advantage. This raises the question as to why the larger sex does not always benefit from its size in sibling competition. We studied sibling competition in the black-headed gull (Larus ridibundus), a sexually-size dimorphic species with male-biased mortality. We manipulated the natural brood sex ratio and placed one male chick in direct competition with one female chick while concurrently controlling for differences in age, size and laying order. Male chicks outgrew their female siblings by 15% in asymptotic body mass and did not suffer from enhanced mortality. Female chicks tended to be more alert when the parents returned to the nest and were more persistent in gull-typical begging displays. Females were more likely to get the first food item, but they did not get more food, possibly due to a size-mediated dominance over the non-monopolizable regurgitated food. Thus, it is unlikely that sex differences in competitiveness significantly contribute to male-biased mortality in black-headed gulls. The previously reported male-biased mortality is more likely due to a disadvantage of a higher food demand and a higher sensitivity towards low egg quality, as has been shown in previous studies.