Synergism between sib-rearing and sex ratio in Hymenoptera

Summary In many bees and wasps, solitary females produce offspring without help from other females. The transition from lone mothers producing offspring to situations in which females often help to rear siblings is an important step in the origins of complex sociality and nonreproductive castes. Recent work on Hymenoptera has stressed the role of sex ratio variation in this transition; when a mother's brood is more female biased than average, older daughters are favored to help rear their younger siblings because they are more closely related to sisters than to their own offspring. Here the direction of causality is from biased sex ratios, which arise by some extrinsic mechanism, to the origins of sib-rearing (eusociality). We present a model in which there is a synergism between sib-rearing and female-biased sex ratios, which may either complement the sex ratio variation idea by increasing the rate at which helping spreads or be an alternative hypothesis about the origins of eusociality. The synergism in our model depends on three conditions. 1) Daughters that help cause more food to be provisioned per offspring, which in turn causes larger offspring. 2) Females gain more than males by being large, which favors mothers with helpers to produce a higher proportion of daughters. 3) A helper's inclusive fitness rises as her mother's brood becomes increasingly female biased because a female helper is more closely related to her sisters than to her brothers. A female helper may also be more closely related to her sisters than to her own offspring, but this particular sibling-offspring relatedness asymmetry is not required by the synergism model. These three conditions create a synergism which favors a rapid transition from solitary (subsocial) to eusocial. Demographic and ecological factors that facilitate the evolution of eusociality reduce the stringency of the relatedness asymmetry condition (3) required by our idea. The synergism model therefore complements factors other than relatedness that may have been important during the evolution of eusociality.     

Sex ratio manipulation and decreased growth of male offspring of undernourished golden hamsters (Mesocricetus auratus)

Summary The theory that female mammals in poor condition may increase individual fitness by skewing the sex ratio of their offspring toward daughters and by investing more resources in daughters than in sons was tested in hamsters. Newly mated experimental females were food-restricted during pregnancy and lactation (RR) or during lactation only (AR). Controls received food ad libitum. Maternal body weights were assessed daily from mating to 25 days postpartum. Litter survival (% litters with at least one pup surviving on any day), litter size, offspring sex ratios (=% males), and pup weights were monitored daily from birth (Day 1) to Day 25. All control and AR dams gave birth 16 days after mating. Gestation was extended by 1–3 days for 35.4% of RR dams. RR dams weighed significantly less at parturition than controls and AR females. During lactation, AR females showed the greatest weight loss and control females the least. AR weight loss exceeded that of RR females, possibly because the former maintained larger litters. Survival was highest for controls, intermediate for AR, and lowest for RR litters. Mean sex ratio at birth was significantly less for RR (40.7%) than for control (49.6%) and AR (48.8%) litters. RR sex ratio did not change significantly postnatally. Sex ratios of control and AR litters never differed statistically from 50%. Control male pups were significantly heavier than their sisters throughout the experiment. No significant gender differences were observed for AR pup weights after Day 2 postpartum. RR female offspring weighed more than their brothers throughout the experiment, and this difference was statistically significant immediately prior to the time that pups began to feed independently (Days 14–17). RR female pup weights were similar to, and sometimes significantly greater than, weights of control daughters during the period of postnatal maternal investment. Control males were always heavier than males from the other treatments. Patterns of weight gain by AR and RR males varied with age. We conclude that underfed female hamsters are able to adjust the sex ratio of offspring prenatally and parental investment postnatally to favor daughters.     

Do female roe deer in good condition produce more sons than daughters

In polygynous roe deer Capreolus capreolus, males are only slightly heavier than females and the overall sex ratio at birth is close to unity. We studied offspring sex ratio and litter size (range 1–4, n = 74) of culled females, in utero, which provided an opportunity to examine responses of sex ratio to maternal condition. Male embryos were heavier than their sisters, and male fawns (9 months old) heavier than female fawns, suggesting a higher growth rate in males. There was no evidence for differential mortality between the sexes from birth to 9 months old. Heavier adult females produced larger embryos than lighter, or primiparous females. The overall sex ratio of embryos did not differ from unity, but adult does had more male embryos (55%) than primiparous does (32%), and the proportion of male embryos in a litter increased with the mother's body mass. Litter size also tended to increase with maternal age and body mass. We argue that this pattern reflects adaptive variation in offspring sex ratio.     

Sex bias or equal opportunity? Patterns of maternal investment in bison

Summary In polygynous mammals, it may be adaptive for mothers to invest more in sons and/or to adjust the sex ratio of offspring in relation to body condition. Calving patterns were examined over an 8-year period (1982–1989) for a population of Bison bison in which barren females are not selectively culled. From these data, we tested predictions of the sex ratio adjustment hypothesis as well as two assumptions: (1) that offspring weight at the end of the period of parental investment (PI) is correlated with later condition, and (2) that maternal and offspring condition during the period of PI are correlated. In contrast to predictions, there was little evidence that mothers in better condition bear more sons. Short- and long-term measures of maternal condition (previous reproductive status, age, dominance status, pre-pubertal body weight, age at first reproduction, birth date, and the duration of the mother's own suckling period) were little related to offspring sex ratio, although the last calves of old females were nearly always female. Similarly, there was little evidence for sex-biased PI. Weights at about 7 months of age were greater for males than females; males also had somewhat later birth dates, suggesting either longer gestation or later conception. However, maternal reproductive costs, as measured by subsequent fecundity, weight loss, and interbirth intervals, did not vary with calf sex. Both assumptions of the model received some support. However, while maternal condition was correlated with offspring condition, there may be sex differences in investment patterns. Mothers appear better able to influence the condition of daughters than of sons. This sex difference may negate any benefit from male-biased investment.     

Variations in the birth sex ratio and neonatal mortality in a natural herd of horses

Variations in birth sex ratios and sex differences in juvenile mortality occur in a number of mammalian species, and in many cases have been linked to resource availability. Most of these biases in offspring sex ratios concern polygynous species with pronounced sexual dimorphism, and where females only are philopatric. Data on species with unusual life-history strategies, such as slight sexual dimorphism or dispersal by both sexes, are of particular interest. In this study of a natural herd of horses (Equus caballus) which experienced an eruptive cycle, and therefore a period of nutritional stress, male offspring had higher neonatal mortality rates in nutritionally poor years than in good ones, whereas “year quality” had no effect on the mortality of female offspring; year quality could therefore be used by mares as predictor of sex-specific offspring survival. We show that the environmental conditions that predicted lower survival of males were negatively related to their production: the birth sex ratio the following year was female-biased; and mares were less likely to produce a son when they had produced a son the preceding year. There was no significant effect of mother's parity, age or rank, or the timing of conception or birth on offspring sex ratios. The mechanism leading to biases in the birth sex ratio could have been the loss of male embryos by mares that did not foal. As there was no evidence for selective abortion of male foetuses in females that did foal the next year, it is not necessary to invoke maternal adjustment, though this remains a possibility. Finally, there was a suggestion that male offspring were more costly to raise than females, since mothers that reared a son in poor years tended to experience an increase in the interbirth interval between their two subsequent offspring. Received: 28 December 1996 / Accepted after revision: 27 July 1997     

Maternal investment in rhesus macaques (Macaca mulatta): reproductive costs and consequences of raising sons

Maternal investment in offspring is expected to vary according to offspring sex when the reproductive success of the progeny is a function of differential levels of parental expenditure. We conducted a longitudinal investigation of rhesus macaques to determine whether variation in male progeny production, measured with both DNA fingerprinting and short tandem repeat marker typing, could be traced back to patterns of maternal investment. Males weigh significantly more than females at birth, despite an absence of sex differences in gestation length. Size dimorphism increases during infancy, with maternal rank associated with son’s, but not daughter’s, weight at the end of the period of maternal investment. Son’s, but not daughter’s, weight at 1 year of age is significantly correlated with adult weight, and male, but not female, weight accounts for a portion of the variance in reproductive success. Variance in annual offspring output was three- to fourfold higher in males than in females. We suggest that energetic costs of rearing sons could be buffered by fetal delivery of testosterone to the mother, which is aromatized to estrogen and fosters fat accumulation during gestation. We conclude that maternal investment is only slightly greater in sons than in daughters, with mothers endowing sons with extra resources because son, but not daughter, mass has ramifications for offspring sirehood. However, male reproductive tactics supersede maternal investment patterns as fundamental regulators of male fitness. Received: 23 July 1999 / Received in revised form: 23 February 2000 / Accepted: 13 March 2000     

A field test of the effects of familiarity and relatedness on social associations and reproduction in prairie voles

Inbreeding depression is a well-documented phenomenon. In animals, one means of avoiding the costs of inbreeding is through the recognition and avoidance of kin as mates. Prairie voles (Microtus ochrogaster) are short-lived, socially monogamous rodents that demonstrate inbreeding depression in the laboratory. Field data indicate that pair formation in nature is opportunistic but pairing among close relatives seems uncommon. We examined the role of relatedness and familiarity on prairie vole social associations and reproduction by placing adult voles into 0.1-ha enclosures with familiar siblings, unfamiliar siblings, and unrelated, unfamiliar conspecifics. Live-trapping data indicated that indices of social pair bonding were random with respect to relatedness and familiarity. Among females whose litters were sired by a single male, litters were significantly more likely to be sired by unfamiliar than familiar males, but the number of litters sired by males that were unrelated to their partner was not different from the number of litters sired by males that were related to their partner. Additionally, females that produced offspring with familiar siblings were significantly more likely to have litters with multiple paternity than females not producing offspring with familiar siblings. However, multiple paternity was not influenced by relatedness of sires. Finally, older individuals were more likely to produce offspring with each other than with younger individuals. Our findings suggest that prior association is a more important mechanism of inbreeding avoidance than phenotype matching for prairie voles mating under ecologically relevant conditions.     

Does it matter that male beaugregory damselfish have a mate preference?

Male beaugregory damselfish (Stegastes leucostictus) spent more time courting larger females in both two-choice and single presentations. Female size was significantly correlated with gonad weight. We also verified that female fecundity was extremely variable within a natural population. We found that male reproductive success was highly correlated with both clutch size and clutch number. However, clutch size was not significantly correlated with clutch number, indicating that males that received larger clutches did not receive more egg clutches. Furthermore, there was no difference between the number of offspring produced by males that mated with the largest females and by males that mated with the most females. Thus, although males preferred larger females, males produced similar numbers of offspring by mating with large females or mating with many females. Received: 7 March 1997 / Accepted after revision: 1 November 1997     

Sex ratio and maternal rank in wild spider monkeys: when daughters disperse

Summary Data from a long-term field study of the spider monkey, Ateles paniscus, in Peru indicate that a strongly female-biased sex ratio exists from birth in this population. Of 46 infants born between July 1981 and June 1986, 12 were male, 32 were female and 2 were of undetermined sex. This effect is consistent between years as well, with more females than males born in each year of the study (Table 1). This bias is driven by the fact that low-ranking females produce daughters almost exclusively, while high-ranking females bias their investment somewhat less strongly towards sons (Table 2). The unusual pattern of female-biased maternal investment observed in this population of Ateles probably occurs for a combination of the following reasons: (1) maternal investment in individual male offspring is somewhat greater than in individual female offspring; (2) males remain with their natal groups, and the sons of high-ranking females are likely to be competitively superior to the sons of low-ranking females; (3) males compete for mates, and only the one or two most dominant males within a community are likely to achieve significant reproductive success. Two possible mechanisms of sex-ratio adjustment and the evidence for each are discussed.     

Sex ratios,mating behavior and sexual size dimorphism of the northern water snake,Nerodia sipedon

Competition among males to mate is generally associated with male-biased size dimorphism. In this study we examine mating behavior in the northern water snake (Nerodia sipedon), a species in which males are much smaller than females despite substantial competition among males to mate. Competition among males was a consequence of a male-biased operational sex ratio due to slightly higher female mortality from a birth sex ratio of 1 : 1, and, in 1 year, more synchronous and longer mating activity by males. Approximately one-third of both males and females appeared not to mate in a given year. Larger males were generally more likely to attempt mating, but size did not explain the variance in the number of aggregations in which individual males participated. Within aggregations, males that were successful at achieving intromission were larger than unsuccessful males in 1 of 2 years. Variation in condition (mass relative to length) and relative tail length were not generally useful predictors of either mating effort or success in males. Because large size was often advantageous to males, sexual size dimorphism appeared not to be a consequence of sexual selection favoring smaller males. Because sexual dimorphism was evident at birth, and both males and females matured sexually at about 4 years, sexual dimorphism was not simply a consequence of one sex growing at the maximum rate for longer. Female fecundity increased with size, and sex differences in size-fecundity relations may underly the pattern of sexual size dimorphism. However, because multiple mating by females is common, sperm competition is likely to be important in determining male reproductive success. Therefore, allocation of energy to sperm rather than growth may also prove to be an important influence on male growth rates and sexual size dimorphism.     

The effects of age and relatedness on mating patterns in thornbug treehoppers: inbreeding avoidance or inbreeding tolerance?

The social environment of many species includes synchronous maturation of siblings in family groups, followed by limited dispersal of adults from their natal site. Under these conditions, females may experience high encounter rates with same-age siblings during mate searching, increasing their risk of inbreeding. If inbreeding depression occurs, mating with a sibling is often considered maladaptive; however, in some contexts, the inclusive fitness benefits of inbreeding may outweigh the costs, favoring females that tolerate some level of inbreeding depression. We evaluated mating patterns in the treehopper Umbonia crassicornis, a semelparous species in which females encounter same-age siblings during mate searching. A female U. crassicornis that mates with a brother suffers from inbreeding depression. We used a free-choice mating design that offered females simultaneous mating opportunities with three groups of males: siblings, same-age nonsiblings, and older nonsiblings. These groups represent the types of males typically encountered by females during mate searching. Our goal was to assess whether mating patterns were influenced by inbreeding avoidance by evaluating two hypotheses: kin discrimination and age-based mating (older males cannot be siblings in this species). There was no difference in the proportions of females mating with siblings vs nonsiblings, suggesting an absence of kin discrimination. However, females mated with a greater proportion of older vs younger males. Given that females do not avoid siblings as mates despite a cost to inbreeding, our results provide a possible example of inbreeding tolerance. We also discuss some factors that may have contributed to the mating advantage of older males.     

Sex allocation and local mate competition in Old World non-pollinating fig wasps

The populations of many species are structured such that mating is not random and occurs between members of local patches. When patches are founded by a single female and all matings occur between siblings, brothers may compete with each other for matings with their sisters. This local mate competition (LMC) selects for a female-biased sex ratio, especially in species where females have control over offspring sex, as in the parasitic Hymenoptera. Two factors are predicted to decrease the degree of female bias: (1) an increase in the number of foundress females in the patch and (2) an increase in the fraction of individuals mating after dispersal from the natal patch. Pollinating fig wasps are well known as classic examples of species where all matings occur in the local patch. We studied non-pollinating fig wasps, which are more diverse than the pollinating fig wasps and also provide natural experimental groups of species with different male morphologies that are linked to different mating structures. In this group of wasps, species with wingless males mate in the local patch (i.e. the fig fruit) while winged male species mate after dispersal. Species with both kinds of male have a mixture of local and non-local mating. Data from 44 species show that sex ratios (defined as the proportion of males) are in accordance with theoretical predictions: wingless male species<wing-dimorphic male species<winged male species. These results are also supported by a formal comparative analysis that controls for phylogeny. The foundress number is difficult to estimate directly for non-pollinating fig wasps but a robust indirect method leads to the prediction that foundress number, and hence sex ratio, should increase with the proportion of patches occupied in a crop. This result is supported strongly across 19 species with wingless males, but not across 8 species with winged males. The mean sex ratios for species with winged males are not significantly different from 0.5, and the absence of the correlation observed across species with wingless males may reflect weak selection to adjust the sex ratio in species whose population mating structure tends not to be subdivided. The same relationship is also predicted to occur within species if individual females adjust their sex ratios facultatively. This final prediction was not supported by data from a wingless male species, a male wing-dimorphic species or a winged male species. Received: 27 July 1998 / Received in revised form: 11 January 1999 / Accepted: 16 January 1999     

Parental investment and its ecological consequences in the solitary wasp Sceliphron assimile (Dahlbom) (Sphecidae)

Summary The sphecid wasp Sceliphron assimile Dahlbom builds mud cells whose volumes are positively correlated to maternal length, the relationship being highly significant for daughters (Fig. 1). However, small females build cells longer and wider in relation to their own body length than large females do (Fig. 3). Cell volume closely regulates the total weight of prey stocked per cell (Figs. 4, 10). Maternal length is directly related to prey size and inversely related to the number of prey stocked (Fig. 5). Female length, above a threshold of 19 mm, probably does not influence her chance of starting a nest (Fig. 6) but is strongly positively correlated to achieved fecundity. Large females have larger offspring and a higher proportion of daughters than small females do (Fig. 7), but this is largely controlled by maternal behaviour rather than by genetical mechanisms (Fig. 10). The results are in general agreement with those of Trivers and Hare (1976) for solitary wasps, but large progeny are cheaper to produce per unit weight than small ones, and for a given weight sons are cheaper to produce than daughters.     

Local mate competition and precise sex ratios in Telenomus fariai (Hymenoptera: Scelionidae), a parasitoid of triatomine eggs

Telenomus fariai is a gregarious endoparasitoid of the eggs of several species of Triatominae (Hemiptera) with a high degree of sibmating: males fertilize their sisters inside the host egg before emergence or emerge first and copulate with their sisters as these emerge. Our results show that, when laying alone, T. fariai behaves adaptively, minimizing offspring mortality and conforming to the prediction of local mate competition (LMC) theory by laying a single male, which is sufficient to fertilize all the sisters. When more than one wasp was placed with one host, sex ratios still conformed to LMC predictions but, despite the decreasing number of eggs laid per wasp, clutch size could not be completely adjusted to avoid mortality. This is not surprising, as superparasitism is rare in the field. Offspring production was independent of the contacts between conspecifics but was affected by the number of mothers laying on a single host egg. The sex of the progeny was precisely determined: a female produced one male per clutch when laying on both unparasitized or previously parasitized hosts. On the other hand, a mother produced less daughters when superparasitizing. Under crowded conditions, the number of eggs laid per female wasp and per host decreased as the number of mothers increased. Developmental mortality also increased with the number of T. fariai eggs per host, determining a maximum of approximately 14 emerged adults. Host resources per individual affected male and female adult size with similar intensity, and male adult mortality was slightly higher than that for females. These results, and previous findings, suggest that T. fariai attains Hamiltonian sex ratios by laying one male and a variable number of females, and that the detection of chemical marks left by conspecifics provides information on the number of foundresses sharing a patch. Received: 4 February 2000 / Received in revised form: 19 April 2000 / Accepted: 20 May 2000     

Parent–offspring and sibling conflict in Galápagos fur seals and sea lions

Parent–offspring conflict theory is well supported by theoretical arguments. However, empirical observations are often difficult to interpret and have contradicted one of its most appealing predictions that parent and offspring should disagree over killing of nest or littermates. We present the first examples of deadly conflict between siblings of different cohorts. In Galápagos fur seals (Arctocephalus galapagoensis) and sea lions (Zalophus wollebaeki), mothers often wean their single offspring at 2 years. This leads to a situation where up to 23% of all pups are born while the older sibling is still being nursed. Younger siblings are disadvantaged by being born lighter than neonates without older still dependent siblings. Pups born while an older sib is still dependent grow less in early life (fur seal) and suffer increased early mortality (both species) through direct aggression or scramble competition with the older sibling. This effect is much stronger in years of high sea surface temperature (El Niño) indicating low marine productivity and if the older offspring is a male. In both species, mothers interfere aggressively in this conflict by defending the younger offspring. In years of El Niño, intense resistance to maternal aggression by the older offspring happens frequently in the fur seal. Such resistance against weaning can induce maternal neglect of the newborn. Given substantial year to year variation in offspring growth, maternal aggression forces weaning in the older sibling only if it has reached sufficient size to support itself by foraging. In Galápagos fur seals, pups with older siblings can either represent insurance against loss of older offspring or extra reproductive value.     

Male limitation of female reproductive success in a pipefish: effects of body-size differences

Summary In the pipefish Syngnathus typhle, a species with exclusive male parental care, males limit female reproductive success because of their limited brood pouch space and long pregnancy. Sexual size dimorphism is absent in these 1-year-old animals but increases with age so that older females are larger than similarly aged males. Because fecundity is related to size in both sexes and increases more rapidly with body size in females than in males, the difference in growth increases female fecundity more, relative to male fecundity, as the fish get older. We therefore predicted that male limitation of female reproductive success is even more severe when all age classes are considered. To measure a female's maximum reproductive rate, she was provided with three males. Small 1-year-old females produced as many eggs, or produced eggs at the same rate, as a male of similar size could care for. Small females filled on average 1.06 males within the time span of one male pregnancy and actually produced on average 10 eggs fewer than needed to fill a similarly sized male. Large 2-year-old females, in contrast, produced on average a surplus of 149 eggs and filled 2.7 similarly sized males within the course of one pregnancy. The difference between females of the two size classes was highly significant. Males prefer to mate with larger females if given a choice. In nature sex ratios are equal, and males limit female reproductive success in the whole population. Therefore, small females are more severely constrained by mate availability than are larger females because males choose to mate with larger females. Offprint requests to: A Berglund     

Prenatal social conditions shape offspring adult phenotype and reproductive success

The prenatal social environment affects offspring development in most studied taxa with potentially lifelong consequences. To understand the adaptive significance of such maternal influences on offspring development, it is important to study their effects on fitness. In guinea pigs, social instability during pregnancy leads to delayed development of male offspring. This has been interpreted as an adaptation to high social densities, where young males need to queue for reproductive opportunities since they cannot out-compete older dominant males. The consequences for male reproductive success are, however, so far unknown. To study the effects of different prenatal social densities on offspring reproductive performance, we housed females individually or in small groups during late pregnancy. Offspring from both treatments were reared together in large groups until independence and thereafter housed in same-sex pairs of the same treatment. We then observed courtship, aggressive behavior, and reproductive success in a low-density context with one male from each treatment competing over access to two females. Sons born to individually housed females initiated more fights, had more social contacts, courted females more, and had a higher reproductive success than sons of group-housed females. Sons born to mothers experiencing low social densities before birth therefore perform better at low social group sizes, suggesting that male development may be adaptively adjusted to anticipated social densities, although performance under high densities still needs to be compared.     

Sex differences in maternal investment by Macaca mulatta

Summary The possibility that male offspring are more costly to produce than female offspring was investigated in captive Macaca mulatta. Differential maternal investment into 187 infants was evaluated by comparing birth weight, infant growth rate, month of birth, and interbirth interval of males and females. There were no statistically significant differences between male and female infants for any of these variables.     

Mate choice,fecundity and sexual dimorphism in two pipefish species (Syngnathidae)

Summary In order to understand the causes of sexual dimorphism, mate choice and size-related fecundity were studied in two pipefish species, Syngnathus typhle and Nerophis ophidion. Sexual dimorphism is more pronounced in N. ophidion; females are larger, have sexual colourings, and are more active during courtship. In S. typhle the sexes are alike in all these respects. Males brood their offspring in both species. In N. ophidion fecundity was positively correlated with both body size and the amount of sexual colouring in females. In males no correlation between body size and fecundity, or between body size and embryo size existed. Predictably, in mate choice experiments with equal-sized females, males chose females with more extensive sexual colourings. We explain sexual dimorphism in this species as a consequence of both natural selection (fecundity increases with size in females but not in males) and sexual selection (males prefer larger females). We argue that sexual size dimorphism did not evolve by selection minimizing overlap in food niches between the sexes, because food production is high in the Zostera beds where the fishes live, and no size dimorphism was found in the sympatrically occurring S. typhle. Furthermore, in N. ophidion dimorphism is not greater in a particular mouth character than in overall body size. In S. typhle egg size and the average number of eggs transferred per spawning were positively correlated with female body size. Apparently more energy per offspring was provided by larger males than by smaller males, and larger males also carried more offspring. As predicted, large mates were preferred by both sexes in mate choice experiments. This is explicable in terms of both natural selection (fecundity increases with size in both sexes) and sexual selection (both sexes prefer large mates). As a consequence of selection acting in the same direction in both sexes, sexual dimorphism is absent in S. typhle.     

Androgen-dependent maternal effects on offspring fitness in zebra finches

There is accumulating evidence that maternal hormones may play a role in offspring sex adjustment, but little is known about the costs of such hormone-mediated mechanisms. Recent studies have reported sex-specific effects of hormones on offspring viability. Specifically, we previously found that elevating the plasma androgen level in mothers results in a male-biased offspring primary sex ratio, but it affects the viability of sons negatively and daughters positively in zebra finches (Taeniopygia guttata; Rutkowska and Cichoń, Anim Behav, 71:1283–1288, 2006). In this study, we studied further fitness consequences of exposure to elevated yolk androgen levels in zebra finches. We measured growth rate and cellular immune response of nestlings that hatched from eggs laid by females injected with testosterone during egg laying and nestlings of unaffected control females. We found that sons of testosterone-treated females grew slower in comparison to sons of control females. The significant interaction between experimental group and offspring sex indicates that sons of testosterone-treated mothers suffered impaired immune responsiveness while daughters seemed to benefit from elevated androgen level in terms of enhanced immune responsiveness. We found no effects of androgens on offspring performance at adulthood—neither fecundity of females nor attractiveness of males was affected. We conclude that the benefits of biasing sex ratio towards males by increasing androgen level in the yolk may be limited due to negative effects on male offspring performance early in life.