Sex ratio and fledging success of supplementary-fed Tengmalm's owl broods

A nest box population of Tengmalm's owls (Aegolius funereus) in northern Sweden was studied to investigate the effects of extra food on the sex ratio between hatching and fledging in this sexually size-dimorphic species. The brood size and brood sex ratio of supplementary-fed and control broods were compared. Newly hatched nestlings were blood sampled and sexed by polymerase chain reaction (PCR) amplification of the sex-linked CHD1Z and CHD1W genes. The brood sex ratio at hatching was strongly male biased (65%); this was also the case in broods where all eggs hatched (72%). There was no relationship between hatch order and sex ratio, and hatching sex ratio did not vary significantly with laying date. Brood size decreased between hatching and fledging, but did not differ between fed and control broods at either stage. Brood sex ratio did not differ between hatching and fledging, and fledging sex ratio did not differ between fed and control broods. It was concluded that, at least during the year in which the study was carried out, feeding had no effect on brood reduction, and that male and female nestlings did not show any differential mortality. The mechanisms behind the male-biased sex ratio at hatching, and any possible adaptive reasons for it, are not known.     

Hatching asynchrony and growth trade‐offs within domesticated and wild zebra finch,Taeniopygia guttata,broods

The Australian zebra finch, Taeniopygia guttata, is a widely used model organism, yet few studies have compared domesticated and wild birds with the aim of examining its relevance as an evolutionary model species. Domestic and wild broods hatch over approximately 4 and 2 days, respectively, which is important given that nestlings can fledge after as little as 12 days, although 16–18 days is common. We aimed to evaluate the extent to which the greater hatching asynchrony in domestic stock may effect reproductive success through greater variance in size hierarchies, variance in within‐brood growth rates, and partial brood mortality. Therefore, by simultaneously controlling brood sizes and experimentally manipulating hatching intervals in both domesticated and wild birds, we investigated the consequences of hatching intervals for fledging success and nestling growth patterns, as well as trade‐offs. Fledging success was similarly high in domestic and wild broods of either hatching pattern. Nonetheless, between‐brood analyses revealed that domestic nestlings had significantly higher masses, larger skeletal characters, and longer wings than their wild counterparts, although wild nestlings had comparable wing lengths at the pre‐fledging stage. Moreover, within‐brood analyses revealed only negligible differences between domestic and wild nestlings, and larger effects of hatching order and hatching pattern. Therefore, despite significant differences in the hatching intervals, and the ultimate size achieved by nestlings, the domestication process does not appear to have significantly altered nestling growth trade‐offs. The present study provides reassuring evidence that studies involving domesticated zebra finches, or other domesticated model organisms, may provide reasonable adaptive explanations in behavioural and evolutionary ecology. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 763–773.     

Sexual dimorphism,survival, and parental investment in relation to offspring sex in a precocial bird

Differential growth rate between males and females, owing to a sexual size dimorphism, has been proposed as a mechanism driving sex‐biased survival. How parents respond to this selection pressure through sex ratio manipulation and sex‐biased parental investment can have a dramatic influence on fitness. We determined how differential growth rates during early life resulting from sexual size dimorphism affected survival of young and how parents may respond in a precocial bird, the black brant Branta bernicla nigricans. We hypothesized that more rapidly growing male goslings would suffer greater mortality than females during brood rearing and that parents would respond to this by manipulating their primary sex ratio and parental investment. Male brant goslings suffered a 19.5% reduction in survival relative to female goslings and, based on simulation, we determined that a female biased population sex ratio at fledging was never overcome even though previous work demonstrated a slight male‐biased post‐fledging survival rate. Contrary to the Fisherian sex ratio adjustment hypothesis we found that individual adult female brant did not manipulate their primary sex ratio (50.39% male, n = 645), in response to the sex‐biased population level sex ratio. However, female condition at the start of the parental care period was a good predictor of their primary sex ratio. Finally, we examined how females changed their behavior in response to primary sex ratio of their broods. We hypothesized that parents would take male biased broods to areas with increased growth rates. Parents with male biased primary sex ratios took broods to areas with higher growth rates. These factors together suggest that sex‐biased growth rates during early life can dramatically affect population dynamics through sex‐biased survival and recruitment which in turn affects decisions parents make about sex allocation and sex‐biased parental investment in offspring to maximize fitness.     

Chick mortality leads to male‐biased sex ratios in endangered Great Lakes Piping Plovers

Few investigators have studied the offspring sex ratios of monomorphic shorebirds because visually determining the sex of juveniles is not possible. We investigated the ontogeny of an observed male‐biased adult sex ratio in the federally endangered Great Lakes population of Piping Plovers (Charadrius melodus). We determined sex ratios at hatching, banding ( = 9.0 d old), and fledging (23 d old) to determine if the bias arises during the pre‐fledging period and, if so, at what stage. For three consecutive years (2012–2014), we used a molecular technique to determine the sex of 307 chicks and followed individuals to a stage where survival to fledging could be inferred. Within fully‐sexed broods at hatching, the average proportions of male chicks (2012–2014) were 0.47, 0.58, and 0.54, respectively. At banding, the sex ratio remained unbiased in 2012 (0.51), but was male‐biased in 2013 (0.59) and 2014 (0.57). Overall, the sex ratio did not differ significantly from parity at fledging in 2012, but did differ during 2013 (P = 0.01) and 2014 (P = 0.03). Using logistic regression models fit using Bayesian inference, we found strong support for a sex effect on chick survival to fledging age, with higher male than female survival (μ_male = 0.83 [95% credible interval: 0.75–0.90]; μ_female = 0.71 [0.61–0.80]). These results suggest that the male‐biased adult sex ratio in Piping Plovers arises, in part, due to differential survival during the pre‐fledging period. This difference did not result from female chicks hatching later in the season or weighing less at banding than male chicks, factors that could potentially affect the likelihood of survival. Future investigations into possible behavioral‐ or weather‐related influences on sex‐specific survival are needed. Our results have important implications for (1) identifying management efforts needed to increase recruitment given female‐biased chick mortality, and (2) conducting population viability analyses, which frequently assume an unbiased fledgling sex ratio.     

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.     

Brood‐parasite‐induced female‐biased mortality affects songbird demography: negative implications for conservation

Parasites, of all sorts, can profoundly affect host population dynamics. Parasites commonly cause sex‐biased mortality and this can add to their impact. Female‐biased mortality in particular can destabilize dynamics and promote population collapse. We previously reported in a correlative study that brown‐headed cowbird Molothrus ater brood parasitism of song sparrows Melospiza melodia appears to cause female‐biased host nestling mortality. Here, we report results from ‘infestation’ and ‘de‐infestation’ experiments designed to test whether brood parasitism causes female‐biased mortality, and we document the resulting demographic impact using a simulation model. Experimental cowbird infestation of song sparrow nests halved the proportion of female host nestlings (0.31±0.07 vs 0.59±0.06; infested vs unparasitized nests at day 6) replicating the halving reported in naturally cowbird‐parasitized nests (0.28±0.01 vs 0.57±0.05; parasitized vs unparasitized). De‐infestation of naturally cowbird‐parasitized nests in turn wholly eliminated any effect on the proportion of female host nestlings (0.53±0.13 vs 0.54±0.06; de‐infested vs unparasitized) confirming that brood parasitism is the cause. This halving of the proportion of females fledging is likely to be as significant as nest predation in affecting population dynamics, based on the elasticities derived from our demographic model (–0.50 vs –0.59). Experimental infestation reduced the testosterone levels, begging behaviour, and body mass of six day old female host nestlings, whereas males were largely unaffected, suggesting that it is the exacerbation of intra‐brood competition that may be primarily responsible for the resulting female‐biased mortality. The brown‐headed cowbird is invasive in most of North America and has been implicated in regional population declines of many native species. We suggest that female‐biased host offspring mortality is likely to be commonplace among the 144 host species the cowbird successfully parasitizes, and we discuss the negative implications for songbird conservation, given the projected demographic impact.     

Sex allocation and sex‐specific parental investment in an endangered seabird

Biased offspring sex ratio is relatively rare in birds and sex allocation can vary with environmental conditions, with the larger and more costly sex, which can be either the male or female depending on species, favoured during high food availability. Sex‐specific parental investment may lead to biased mortality and, coupled with unequal production of one sex, may result in biased adult sex ratio, with potential grave consequences on population stability. The African Penguin Spheniscus demersus, endemic to southern Africa, is an endangered monogamous seabird with bi‐parental care. Female adult African Penguins are smaller, have a higher foraging effort when breeding and higher mortality compared with adult males. In 2015, a year in which environmental conditions were favourable for breeding, African Penguin chick production on Bird Island, Algoa Bay, South Africa, was skewed towards males (1.5 males to 1 female). Males also had higher growth rates and fledging mass than females, with potentially higher post‐fledging survival. Female, but not male, parents had higher foraging effort and lower body condition with increasing number of male chicks in their brood, thereby revealing flexibility in their parental strategy, but also the costs of their investment in their current brood. The combination of male‐biased chick production and higher female mortality, possibly at the juvenile stage as a result of lower parental investment in female chicks, and/or at the adult stage as a result of higher parental investment, may contribute to a biased adult sex ratio (ASR) in this species. While further research during years of contrasting food availability is needed to confirm this trend, populations with male‐skewed ASRs have higher extinction risks and conservation strategies aiming to benefit female African Penguin might need to be developed.     

Costs of rearing and sex‐ratio variation in southern grey shrike Lanius meridionalis broods

Several non‐mutually exclusive hypotheses predict adaptive variation in the offspring sex ratio. When conditions for breeding are adverse, parents are predicted to produce more offspring of the less costly sex to rear (‘the cost‐of‐reproduction hypothesis’). Moreover, they also should produce the more dispersing sex in order to diminish future competition (‘the local‐resource‐competition hypothesis’). Here, we analyse brood sex ratio according to rearing conditions in the southern shrike Lanius meridionalis, a species with moderately reversed sexual dimorphism. Our results suggest that females are more costly to rear than males in this species. Adult females proved heavier than males, and female nestling tended to be heavier than male nestlings. Moreover, the greater brood reduction, the more male‐biased was the brood, suggesting that brood reduction implied higher mortality in female nestlings. Consistent with these findings, the brood sex ratio was biased to the less costly sex (males) when breeding conditions were adverse (bad years or low‐quality male parents), supporting the cost‐of‐reproduction hypothesis. By contrast, these findings did not support the local‐resource‐competition hypothesis, which predicted female‐biased brood sex ratio under adverse conditions. As a whole, our results support the idea that birds adaptively modulate sex ratio in order to minimize reproduction costs.     

Hatching order and size-dependent mortality in relation to brood sex ratio composition in chinstrap penguins

The differential environmental sensitivity of the sexes hasstrong implications in the evolutionary history of species asit can alter sexual size dimorphism, population sex ratios,and the faculty of parents to manipulate offspring sex in relationto environmental conditions. We studied sexual differences inhatching patterns and evaluated sex- and size-related mortalityin relation to hatching order and brood sex ratios in the chinstrappenguin Pygoscelis antarctica, a moderately size-dimorphic species,with a modal clutch size of 2 eggs. We found that male, second-hatched,and large eggs showed shorter hatching periods than female,first-hatched, and small eggs. We also found a male-biased mortalityof nestlings in the colony. However, male mortality patternsdiffered depending on the brood sex ratio composition. Mortalityof male chicks in all-male broods was higher than in mixed broodsand higher than female mortality in all-female broods. Contrary,females from mixed brood showed higher mortality than theirmale nest mates and higher too than females in all-female broods.Second-hatched chicks also suffered from higher mortality thanfirst-hatched chicks. Our results indicate that both the superiorcompetitive capacity and the higher energy demand of the largersex constitute 2 causal factors explaining patterns of sex-biasedmortality. Both factors occur in the same species and in differentsituations of sibling competition shaped by brood sex ratiocomposition. This study constitutes a good example of how patternsof sex-related mortality can vary depending on nest environmentalcircumstances. Furthermore, our study suggests that hatchingperiod can be a mechanism underlying sexual differences in theembryonic period of birds.     

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.     

Facultative and non‐facultative sex ratio adjustments in a dimorphic bird species

If parental allocation to each offspring sex has the same cost/benefit ratio, Fisher's hypothesis predicts a sex ratio biased towards the cheaper sex. However, in dimorphic birds there is little evidence for this, especially at hatching. We investigated the pre‐fledgling 1) sex ratio, 2) body condition and 3) sex‐differential mortality in a population of the glossy ibis Plegadis falcinellus, in southern Spain between 2001 and 2011. We defined two age groups for the period between hatching and fledging. We also compared pre‐fledgling with the autumn sex ratio. Metabolic rates were estimated by the doubly labeled water (DLW) technique to establish that sons (the bigger sex) were 18% more energy demanding than daughters, and to compute the predicted Fisher's sex ratio (0.465). As population size increased between years, body condition decreased in both sexes, and mortality increased more for daughters than sons prior to fledging. At the same time, the proportion of males among chicks close to fledging increased (average sex ratio: 0.606) while the proportion close to hatching decreased (average sex ratio: 0.434, in line with Fisher's prediction). Furthermore, the proportions of males at fledging and the following autumn were negatively correlated across years. We suggest that, as population density increased and conditions worsened the larger sex had relatively higher survival. These differences in survival produce a shift from a facultative female‐biased sex ratio at hatching into a non‐facultative male‐biased sex ratio of fledglings. Additionally, the excess of males at fledging was counterbalanced by sex‐related dispersal during the autumn. Overall, glossy ibis sex ratio is a product of a combination of facultative and non‐facultative adjustments triggered by environmental conditions, driven by rapid population growth, and mediated by highly interrelated life‐history traits such as body condition, mortality, and dispersal.     

Hatching asynchrony and brood reduction influence immune response in Common Kestrel Falco tinnunculus nestlings

The onset of incubation before the end of laying imposes asynchrony at hatching and, therefore, a size hierarchy in the brood. It has been argued that hatching asynchrony might be a strategy to improve reproductive output in terms of quality or quantity of offspring. However, little is known about the mediating effect of hatching asynchrony on offspring quality when brood reduction occurs. Here, we investigate the relationship between phenotypic quality and hatching asynchrony in Common Kestrel Falco tinnunculus nestlings in Spain. Hatching asynchrony did not increase breeding success or nestling quality. Furthermore, hatching asynchrony and brood reduction had different effects on nestlings’ phytohaematogglutinin (PHA)‐mediated immune response and nestling growth. In asynchronous and reduced broods (in which at least one nestling died), nestlings showed a stronger PHA‐mediated immune response and tended to have a smaller body size compared with nestlings raised in synchronous and reduced broods. When brood reduction occurred in broods hatched synchronously, there was no effect on nestling size, but nestlings had a relatively poor PHA‐mediated immune response compared with nestlings raised in asynchronous and reduced broods. We suggest that resources for growth can be directed to immune function only in asynchronously hatched broods, resulting in improved nestling quality, as suggested by their immune response. We also found that males produced a greater PHA‐mediated immune response than females only in brood‐reduced nests without any effect on nestling size or condition, suggesting that females may trade off immune activities and body condition, size or weight. Overall, our results suggest that hatching pattern and brood reduction may mediate resource allocation to different fitness traits. They also highlight that the resolution of immune‐related trade‐offs when brood reduction occurs may differ between male and female nestlings.     

Brood parasitism causes female‐biased host nestling mortality regardless of parasite species

Inequality in male and female numbers may affect population dynamics and extinction probabilities and so has significant conservation implications. We previously demonstrated that Brown‐headed Cowbird Molothrus ater brood parasitism of Song Sparrows Melospiza melodia results in a 50% reduction in the proportion of female host offspring by day 6 post‐hatch and at fledging, which modelling demonstrated is as significant as nest predation in affecting demography. Many avian brood parasites possess special adaptations to parasitize specific hosts so this sex‐ratio effect could be specific to the interaction between these two species. Alternatively, perturbations associated with brood parasitism per se (e.g. the addition of an extra, larger, unrelated nestling), rather than a Cowbird nestling specifically, may be responsible. We experimentally eliminated the effects of Cowbird‐specific traits by parasitizing nests with a conspecific nestling rather than a Cowbird, while otherwise emulating the circumstances of Cowbird parasitism by adding an extra, larger (2‐day‐older), unrelated Song Sparrow nestling to Song Sparrow nests. Our parasitism treatment led to few host offspring deaths and no evidence of male‐biased sex ratios by day 6 post‐hatch. However, after day 6, female nestling mortality rates increased significantly in experimentally parasitized nests, resulting in a 60% reduction in the proportion of females fledging. Cowbird‐specific traits are thus not necessary to cause female‐biased host nestling mortality and far more general features associated with Cowbird parasitism instead appear responsible. Our results suggest, however, that Cowbird‐specific traits may help accelerate the pace of female host deaths. The conservation implications of our results could be wide reaching. Cowbirds are unrelated to all their hosts, are larger than the great majority, and a Cowbird nestling's presence can mean there is an extra mouth to feed. Thus, sex‐biased mortality in parasitized nests could be occurring across a range of host species.     

Brood sex ratio in the Kentish plover

How and why do the mating opportunities of males and femalesdiffer in natural population of animals? Previously we showedthat females have higher mating opportunities than males inthe Kentish plover Charadrius alexandrinus. Both parents incubatethe eggs, and males provide more brood care than females; thusit is not obvious why the females find new mates sooner thanthe males. In this study we investigated whether the sex-biasedmating opportunities stem from biased offspring sex ratios.We determined the sex of newly hatched, precocial chicks usingCHD gene markers. Among fully sexed broods, 0.461 ± 0.024(SE) of chicks (454 chicks in 158 broods) were male, and thissex ratio was not significantly different from unity. The proportionof males at hatching decreased significantly over the breedingseason, which occurred consistently in all 3 years of the study.Large chicks were more likely to be males than females. Neitherparental age nor body size of male and female parents was relatedto brood sex ratio. We also sexed a number of chicks that werecaught after they left their nest (range of estimated ages 0–17days) and found that the proportion of males increased withbrood age. This relationship remained highly significant whencontrolling statistically for hatching date. As brood size decreaseddue to mortality after the chicks left their nest, these resultssuggest that the mortality of daughters was higher than thatof the sons shortly after hatching. Taken together, our resultsshow that the female-biased mating opportunities in the Kentishplover are not due to biased brood sex ratio at hatching but,at least in part, are due to female-biased chick mortality soonafter hatching.     

The effects of hatching asynchrony on growth and mortality patterns in Eurasian Hoopoe Upupa epops nestlings

Growth is a fundamental life history trait in all organisms and is closely related to individual fitness. In altricial birds, growth of many traits is restricted to the short period between hatching and fledging and strongly depends on the amount of food that parents deliver and the extent of hatching asynchrony. However, empirical studies of energy allocation to growth of different body size traits as a function of hatching asynchrony are scarce. We studied growth and mortality of Eurasian Hoopoe Upupa epops, a species with a long breeding season and high brood size variance, whose nestlings show pronounced hatching asynchrony, in order to test how hatching asynchrony affects different growth traits in the context of territory quality, season and brood size. The growth of five body traits (body mass, and lengths of tarsus, third primary, bill and longest crest feather) was investigated to understand how it was affected by brood size, hatching date and order, and territory quality. In total, 241 nestlings from 39 nests were measured every 4 days in 2014 in south‐western Switzerland. Brood size, hatching date and hatching order had the strongest influence on growth trajectories, although tarsus growth was only marginally affected by these variables. Nestlings that hatched earlier than their siblings were heavier and had longer third primaries, bills and crest feathers compared with later‐hatched siblings. In territories of high quality, hatching order differences disappeared for body mass growth, but persisted for lengths of third primary, bill and crest feathers. Brood size was inversely associated with third primary, bill and crest feather lengths, but positively associated with body mass. Nestling mortality was higher in later‐hatched nestlings and in broods that were raised in territories of lower quality. Our study shows that in nestlings, energy was allocated differentially between body traits and this allocation interacted with hatching order and territory quality. Rapid mass gain by nestlings was prioritized in order to increase competitive ability. Our results provide support for the brood reduction hypothesis as an explanation of hatching asynchrony in Hoopoes.     

Sex allocation in the sexually monomorphic fairy martin

Offspring sex ratios were examined at the population and family level in the sexually monomorphic, socially monogamous fairy martin Petrochelidon ariel at five colony sites over a 4-year period (1993–1996). The sex of 465 nestlings from 169 broods was determined using sex-specific PCR at the CHD locus. In accordance with predicted sex allocation patterns, population sex ratios at hatching and fledging did not differ from parity in any year and the variance in brood sex ratios did not deviate from the binomial distribution. Further, brood sex ratio did not vary with hatching date during the season, brood number, brood size or colony size. The sex ratio of broods with extra-pair young did not differ from those without, while the sex ratio of broods fathered by males that gained extra-pair fertilizations did not differ from broods fathered by other males. Extra-pair chicks were as likely to be male as female. Neither the total number of feeding visits to the brood nor the relative feeding contribution by the sexes varied significantly with brood sex ratio. Brood sex ratios were also unrelated to paternal size, condition and breeding experience or maternal condition and breeding experience. However, contrary to our prediction, brood sex ratio was negatively correlated with maternal size. Generally, these results were consistent with our expectations that brood sex ratios would not vary with environmental factors or parental characteristics, and would not influence the level of parental provisioning. However, the finding that females with longer tarsi produced an excess of daughters is difficult to reconcile with our current understanding of fairy martin life history and breeding ecology.     

Evidence for Strategic Sex Allocation in the Guppy (Poecilia reticulata): Brood Sex Ratio and Size Predict Subsequent Adult Male Mating Success

Sex allocation theory predicts that females should produce more sons when the reproductive success of sons is expected to be high, whereas they should produce more daughters, not daughters when the reproductive success of sons is expected to be low. The guppy (Poecilia reticulata) is a live‐bearing fish, and female guppies are known to produce broods with biased sex ratios. In this study, we examined the relationship between brood sex ratio and reproductive success of sons and daughters, to determine whether female guppies benefit from producing broods with biased sex ratios. We found that sons in male‐biased broods had greater mating success at maturity than sons in female‐biased broods when brood sizes were larger. On the other hand, the reproductive output of daughters was not significantly affected by brood sizes and sex ratios. Our results suggest that female guppies benefit from producing large, male‐biased brood when the reproductive success of sons is expected to be high.     

A novel PCR‐based genetic marker shows that sex of offspring does not account for hatch‐order effects on growth,survival and dominance in Pūkeko Porphyrio melanotus melanotus

Intra‐brood competition can influence a variety of fitness‐related traits in birds. Previous research on the joint‐nesting Pūkeko Porphyrio melanotus melanotus, a New Zealand subspecies of Australasian Swamphen, showed that chicks that hatched earlier in a brood tended to grow faster, were more likely to survive and had higher dominance status as adults than later hatched nest‐mates. However, this finding could be due to changes in offspring sex ratio across hatch order (e.g. if males tend to hatch earlier), which was not previously examined because of methodological challenges associated with sexing nestling Pūkeko. Here, we report a useful PCR‐based genetic marker to determine the sex of Pūkeko. We then used new sex‐specific data to re‐examine patterns of offspring growth, survival and dominance. We found that the sex of offspring does not account for the hatching‐order patterns related to social dominance, growth or survival. Furthermore, changes in offspring sex ratio across hatching‐order were negligible and offspring sex ratios did not differ significantly between the primary female and secondary female broods (in joint‐clutch nests), or when comparing primary female and single female broods. We found no clear evidence for sex ratio bias according to hatching‐order and conclude that hatching‐order and not offspring sex explain patterns of growth, survivorship and adult dominance in Pūkeko.     

Similarity in the begging calls of nestling Red‐winged Blackbirds

ABSTRACT Although individually distinct begging calls may permit parents to recognize their offspring, birds nesting in dense breeding colonies where fledglings intermingle might benefit from additional adaptations. For example, if the calls of all nestlings in a brood were similar, parents would need to recognize only one brood call instead of the identity calls of each nestling. We recorded nestling Red‐winged Blackbirds (Agelaius phoeniceus) to determine whether their calls function to identify individuals (identity call hypothesis) or broods (brood call hypothesis). We used spectrogram cross‐correlation and dynamic time warping as well as call duration, peak frequency, and frequency range to estimate the similarity of begging calls of nestling Red‐winged Blackbirds. We recorded individual nestlings on day 5 and on day 9 of the nestling period to determine whether calls of individuals were more similar than calls of different nestlings, and whether calls of broodmates were more similar than calls of nestlings from different broods. We found that calls of 8‐d‐old individuals were more similar than calls of different nestlings, but the calls of broodmates were not more similar than those of nestlings from different broods. These results were consistent with the identity call hypothesis. We then compared begging calls of pairs of nestlings recorded separately and together on day 9. We found that the calls of 8‐d‐old nestlings recorded together were more similar than when they were recorded separately. In addition, using playback of begging calls from normal broods and artificial “broods” constructed from the calls of single nestlings, we found that females returned with food sooner in response to the calls of single nestlings (with enhanced call similarity) than to those of normal broods. Our results suggest that similar begging calls may be beneficial for both nestlings and parents, with broodmates fed at higher rates when their calls are more similar and, after fledging, parents needing to recognize only one brood call instead of the identity calls of each fledgling.     

Age‐related prenatal maternal effects and postnatal breeding experience have different influences on nestling development in an altricial passerine

Reproductive success and nestling performance are related to the age of parents across several vertebrate taxa. However, because breeding experience and prenatal maternal investment in reproduction often covary, the source of these age‐related differences can be difficult to determine. In this study, we evaluated the influence of prenatal maternal effects and postnatal breeding experience on the performance of nestling tree swallows Tachycineta bicolor by conducting a carefully controlled partial cross‐fostering experiment. We swapped half‐broods of nestlings between the nest of a young first‐time breeding female and the nest of a female known to have previously raised and fledged young. Our manipulation did not influence the within‐brood nestling hierarchies, and controlled for the effects of egg laying order. We found that nestlings of older females were heavier just prior to fledging regardless of the breeding experience of the attending female. In addition, fledglings raised by experienced females grew their flight feathers faster, and had greater probability of fledging. Our study demonstrates that prenatal investment in reproduction by older females can have long‐term consequences on nestling mass, and suggests limited potential for compensatory mass gains prior to fledging. Because our analyses controlled for feeding rates, our results also suggest that foraging quantity and quality are not the only benefits nestlings gain by being raised by an experienced female.