Sex-biased nestling mortality in the Montagu's harrier Circus pygargus

I evaluate causes and patterns of nestling mortality in a sexually dimorphic species, the Montagu's harrier Circus pygargus , and their relationship with sex and condition. Starvation was apparently the main reason for nestling death. Condition of birds that died was lower than those that survived. Both probability of nestling death and the proportion of nestlings that died within a brood increased with the number of hatched nestlings in a brood, and with increasing hatching date. For the nestlings that died after being sexed, when controlling for brood effects, probability of death was significantly related to nestling sex, with smaller males having a higher probability of dying. The probability of nestling death if hatched late in the season was relatively greater for males than for females. There was also a significant interaction between sex and hatching date on nestling condition: the decline in condition if hatched late in the season was steeper for males than for females. Males did not have a higher probability of death when having more sisters: neither the probability of brood reduction nor the proportion of nestlings that died were significantly related to within-brood sex ratio. Results suggest that mortality may partly result from sibling competition: females, being the larger sex, might be better able to compete for food within a brood than their male siblings. Additionally, smaller males may be less able to recover from periods of declining body weight.     

Unusual pattern of sex‐specific mortality in relation to initial brood sex composition in the black‐billed magpie Pica pica

In sexually size‐dimorphic species, brood sex composition may exert differential effects on sex‐specific mortality. We investigated the sex‐specific mortality and body condition in relation to brood sex composition in nestlings of the black‐billed magpie Pica pica. Neither significantly sex‐biased production at hatching nor overall sex‐biased mortality during the nestling period was found. Sex‐specific mortality as a function of brood sex composition, however, differed between female and male nestlings. We found higher mortality for females in male‐biased broods and higher mortality for males in female‐biased broods, a phenomenon that we call ‘rarer‐sex disadvantage’. As a result, fledging sex ratios became more biased in the direction of bias at hatching, a phenomenon that cannot be readily explained by previous hypotheses for sex‐specific mortality. Two temporal variables, fledging date and laying date, were also correlated with sex‐specific mortality: female nestlings in earlier broods experienced higher mortality than male nestlings whereas male nestlings in later broods experienced higher mortality. We suggest that this unusual pattern of mortality may be explained by adaptive adjustments of brood sex composition by parents, either through the effects of a slight sex difference in offspring dispersal patterns on parental fitness, or owing to sex differences as regards the benefits of early fledging.     

Antagonistic natural selection revealed by molecular sex identification of nestling collared flycatchers

Natural selection may act in different directions during different life-history stages, or in different directions on different classes of individuals. Antagonistic selection of this kind may be an important mechanism by which additive genetic variation for quantitative traits is maintained, and can prevent populations or species reaching local adaptive peaks. This paper reports the results of a study of viability selection on morphological traits of nestling collared flycatchers Ficedula albicollis . Analyses performed without knowledge of the sex of nestlings suggested that no selection was occurring on these traits. However, using molecular sex identification with the avian CHD gene, it is shown that selection acts in different directions on male and female body size from fledging to breeding, apparently favouring relatively small males and large females. The results suggest that differential selection on male and female nestlings may contribute to purely phenotypic sexual size dimorphism in this species. These findings highlight the potential of newly developed molecular sexing techniques to reveal the consequences of an individual's gender for many aspects of its life history in taxa where gender cannot be determined on the basis of external appearance.     

Growth of yellow-headed blackbird Xanthocephalus xanthocephalus nestlings in relation to maternal body condition, egg mass, and yolk carotenoids concentrations

Condition‐dependent resource allocation to eggs can affect offspring growth and survival, with potentially different effects on male and female offspring, particularly in sexually dimorphic species. We investigated the influence of maternal body condition (i.e., mass‐tarsus residuals) and two measures of female resource allocation (i.e., egg mass, yolk carotenoid concentrations) on nestling mass and growth rates in the polygynous and highly size dimorphic yellow‐headed blackbird Xanthocephalus xanthocephalus. Egg characteristics and carotenoid concentrations were obtained from the third‐laid egg of each clutch and were correlated with the mass and growth rates of the first two asynchronously hatched nestlings. Maternal body condition was associated with the growth of first‐hatched, but not second‐hatched nestlings. Specifically, females in better body condition produced larger and faster growing first‐hatched nestlings than females in poorer body condition. As predicted for a polygynous, size‐dimorphic species, females that fledged first‐hatched sons were in better body condition than females that fledged first‐hatched daughters. Associations between egg mass, yolk carotenoid content, and nestling growth were also specific to hatching‐order. Egg mass was positively correlated with the mass and growth rates of second‐hatched nestlings, and yolk concentrations of β‐carotene were positively correlated with second‐hatched nestling mass. Surprisingly, the relationship between yolk lutein and hatchling growth differed between the sexes. Females with high concentrations of yolk lutein produced larger and faster growing first‐hatched sons, but smaller first‐hatched daughters than females with lower lutein concentrations. Mass and growth rates did not differ between first‐ and second‐hatched nestlings of the same sex, despite asynchronous hatching in the species. Results from this study suggest that maternal body condition and the allocation of resources to eggs have carotenoid‐, sex‐, and/or hatch‐order‐specific effects on yellow‐headed blackbird nestlings.     

Sex ratio comparisons between nestlings and dead embryos of the Sparrowhawk Accipiter nisus

Sex ratios that differ from unity have been reported for several bird species, but are poorly understood. Skewed sex ratios may originate at ovulation (primary sex ratio) or arise through differential mortality between the sexes (secondary sex ratio). To estimate the primary sex ratio from nestlings is difficult because in some nests not all the offspring can be sexed. Both when including and excluding such nests, there is a risk of overestimating the proportion of the better-surviving sex. Here we sexed dead Sparrowhawk embryos to determine whether unhatched eggs affect primary sex ratio estimates that are based on nestling data. In nests in which embryo mortality occurred, there was up to a 9% discrepancy in the primary sex ratio estimates based on nestlings alone compared to nestlings and dead embryos together. There was no evidence that these differences were based on sex-specific causes of mortality of embryos.     

Fledgling sex ratios in relation to brood size in size-dimorphic altricial birds

In six species of dimorphic raptors (females larger than males)and one passerine (males larger than females), the sex ratioat fledging varied systematically with brood size at fledging.In all species the strongest bias toward the smaller sex wasestablished in the largest as well as the smallest broods; amore even distribution of males and females was observed inbroods of intermediate size. We explored a specific differentialmortality explanation for this sex ratio variation. Our hypothesispostulates that variation in mortality is caused by differencesin food demand between broods of the same size, due to theirsex composition. Data from the marsh harrier Circus aeruginosuson gender-related food demand and overall nestling mortalitywere used to predict the frequency of surviving males and femalesat fledging, assuming an even sex ratio at hatching and randommortality with respect to both sexes within broods. The modelquantitatively fits the marsh harrier data well, especiallyin broods originating from large dutches. Although we anticipatethat other mechanisms are also involved, the results supportthe hypothesis of sex-ratio-dependent mortality, differentialbetween broods, as the process generating the observed brood-sizedependence of fledgling sex ratios in sexually dimorphic birds.     

Pair bond duration influences paternal provisioning and the primary sex ratio of brown thornbill broods

Parents should vary their level of investment in sons and daughters in response to the fitness costs and benefits accrued through male and female offspring. I investigated brood sex ratio biases and parental provisioning behaviour in the brown thornbill, Acanthiza pusilla, a sexually dimorphic Australian passserine. Parents delivered more food to male-biased than female-biased broods. However, factors determining parental provisioning rates differed between the sexes. Female provisioning rates were related to brood sex ratio in both natural and experimental broods with manipulated sex ratios. In contrast, male provisioning rates were not affected by brood sex ratio in either natural or experimental broods. However, males in established pairs provisioned at a higher rate than males in new pairs. Data on the sex ratio of 109 broods suggest that female brown thornbills adjust their primary sex ratio in response to pair bond duration. Females in new pairs produced broods with significantly fewer sons than females in established pairs. This pattern would be beneficial to females if the costs of rearing sons were higher for females in new than established pairs. This may be the case since females in new pairs provisioned experimental all-male broods at elevated rates. The condition of nestlings also tended to decline more in these all-male broods than in other experimental broods. This will have additional fitness consequences because nestling mass influences recruitment in thornbills. Female thornbills may therefore obtain significant fitness benefits from adjusting their brood sex ratio in response to the status of their pair bond. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Intraseasonal effects of clutch manipulation on parental provisioning and residual reproductive value of eastern phoebes (Sayornis phoebe)

Summary First clutches of double-brooded eastern phoebes Sayornis phoebe were manipulated (up two eggs, down 2 eggs or no change) to test for intraseasonal reproductive tradeoffs and to test whether size of first brood influenced food delivery rates to nestlings and nestling quality in second broods.Considering all nests from both broods, rate of feeding nestlings increased linearly with brood size but nestling mass per nest decreased with increasing brood size. High nestling weights in small broods may have resulted from parents delivering better quality food, but we did not test this.Among treatment groups in first broods, nestlings from decreased broods weighed more than those in control or increased broods. Treatment did not influence the likelihood that second nests would be attempted after successful first nests nor did it alter the interval between nests. Nestlings of parents that renested weighed more than those of parents that did not, regardless of treatment, suggesting that post-fledging care may preclude renesting. Mass of individual females did not change between broods, regardless of brood size. Clutch sizes of second attempts were not affected by manipulations of first broods but increasing first broods reduced the number of nestlings parents were able to raise to day 11 in their second broods. However, manipulation of first broods did not affect mean nestling mass per nest of nestlings that survived to day 11.In phoebes, parents of small first broods are able to raise nestlings in better condition. We predict that in harsh years, parents of small first broods would be more likely to renest. Parents of enlarged first broods sacrificed quality of offspring in second broods, which seems a reasonable strategy if nestlings from second broods have lower reproductive value.     

Offspring sex ratio variation in relation to brood size and mortality in a promiscuous species: the Aquatic Warbler Acrocephalus paludicola

We analyse nestling sex ratio variation in the Aquatic Warbler Acrocephalus paludicola to test for predictions from sex allocation theory that the brood sex ratio is close to parity. We also tested Fiala's (1980) prediction that there is no difference in sex ratio between broods affected and not affected by mortality, and whether a shift in primary sex ratios or simple differential mortality by sex underlies that difference. Furthermore, we explore additional analytical possibilities for inferring proximate mechanisms through simulation modelling. In the Aquatic Warbler, which is promiscuous, the overall sex ratio determined by molecular sexing of nestlings at 8–11 days of age did not deviate significantly from parity (proportion of females 0.509), nor did we find any predictive effect of brood size, maternal body mass, fat condition, wing and bill length, laying date, mean daily temperature, and multiple-male mating. However, extensive simulation suggested that the whole pattern of sex ratio variation is unlikely to arise purely by chance: (1) there is a diverging sex ratio between complete and partial broods, (2) large broods tended to be female-biased and small broods male-biased, and (3) low ambient temperature prior to the laying period seemed to increase the proportion of female offspring in complete broods. We conclude that most variation in nestling sex ratio is non-adaptive in nature, and results from variation in female nestlings mortality dependent on brood size and sex ratio.     

Manipulating rearing conditions reveals developmental sensitivity in the smaller sex of a passerine bird, the European starling Sturnus vulgaris

Traditionally, studies of sexually size-dimorphic birds and mammals report that the larger sex is more sensitive to adverse environmental conditions during ontogeny. However, recent studies in avian species that exhibit moderate size-dimorphism indicate that the smaller sex may be more sensitive to poor rearing conditions. To better understand sex-specific sensitivity in a passerine exhibiting moderate size-dimorphism, we examined growth, cell-mediated immunity (CMI) and survival of European starling Sturnus vulgaris nestlings following an experimental reduction of maternal rearing ability (via a feather-clipping manipulation). Contrary to conventional theory, daughters showed reduced growth in both body mass and measures of structural size in response to the maternal treatment. In contrast, sons showed no reductions in any of these traits in relation to the treatment. No sex-specific differences in nestling CMI were found for either group, although CMI of nestlings raised by manipulated mothers were higher than those of control nestlings. Finally, fledging sex ratios did not change from those at hatching indicating that neither sex appeared differentially sensitive to the maternal treatment in terms of mortality. These results reveal that variation in the quality of the rearing environment can have significant effects on the smaller sex of a passerine exhibiting moderate dimorphism and as such support recent studies of species with small-moderate sexual size-dimorphism. Combined results suggest that sex-specific effects of environmental variation on nestling development may be both context- (i.e., brood size, resource level, hatching order) and temporally- (when during development they occur) specific. Furthermore, more studies are needed that examine multiple traits at several developmental stages and then follow the sexes over the longer-term to examine potential effects on fitness.     

Plumage colour in nestling blue tits: sexual dichromatism,condition dependence and genetic effects

Sexual-selection theory assumes that there are costs associated with ornamental plumage coloration. While pigment-based ornaments have repeatedly been shown to be condition dependent, this has been more difficult to demonstrate for structural colours. We present evidence for condition dependence of both types of plumage colour in nestling blue tits (Parus caeruleus). Using reflectance spectrometry, we show that blue tit nestlings are sexually dichromatic, with males having more chromatic (more 'saturated') and ultraviolet (UV)-shifted tail coloration and more chromatic yellow breast coloration. The sexual dimorphism in nestling tail coloration is qualitatively similar to that of chick-feeding adults from the same population. By contrast, the breast plumage of adult birds is not sexually dichromatic in terms of chroma. In nestlings, the chroma of both tail and breast feathers is positively associated with condition (body mass on day 14). The UV/blue hue of the tail feathers is influenced by paternally inherited genes, as indicated by a maternal half-sibling comparison. We conclude that the expression of both carotenoid-based and structural coloration seems to be condition dependent in blue tit nestlings, and that there are additional genetic effects on the hue of the UV/blue tail feathers. The signalling or other functions of sexual dichromatism in nestlings remain obscure. Our study shows that nestling blue tits are suitable model organisms for the study of ontogenetic costs and heritability of both carotenoid-based and structural colour in birds.     

Bill length: a reliable method for sexing Purple Sandpipers

ABSTRACT.   Purple Sandpipers ( Calidris maritima ) are sexually dimorphic, with females larger than males. We tested the reliability of using bill length to sex individuals and estimate the sex ratio at a stopover site in Iceland in May 2003 and 2005. Feather samples from 65 of 324 captured birds were used for molecular sexing, and generalized linear models (GLMs) were used to discriminate the sexes from body measurements of the molecularly sexed birds. About 3% of the 324 individuals were misclassified by the Harding-Cassie method, and the proportion of males was 0.657 compared to 0.656 according to the best GLM. Our results showed that the reliability of determining the sex and sex ratio of Purple Sandpipers using a Harding-Cassie plot of bill length measurements was high, but that reliability was improved by including other variables in a GLM. The estimate of an uneven sex ratio in the population we studied was not due to a systematic error, and supports the conclusion of earlier studies that Purple Sandpipers exhibit an uneven sex ratio in favor of males.     

Sex identification by alternative polymerase chain reaction methods in falconiformes

A number of avian species are difficult to sex morphologically, especially as nestlings. Like other avian species, many species of Falconiformes are sexually monomorphic. Therefore, it is desirable that new methods based on DNA analysis are established in Falconiformes and other sexual monomorphic species. We identified sex in Falconiformes by two alternative methods. First, we used a sexing method based on the intronic length variation between CHD1W and CHD1Z using primers flanking the intron. In this method, two species of Falconidae could be identified for sexing. However, six species of Accipitridae could not, because they have few length variations. The second method used was based on differences in sequences between CHD1W and CHD1Z. From sequence analysis, a 3'-terminal mismatch primer on point mutation conserved among Falconiformes was designed, and identification of sex with the amplification refractory mutation system (ARMS) was performed. This method could identify sex in all species tested. In addition, because the 3'-terminal mismatch primer was designed on a point mutation conserved among Falconiformes, ARMS with these primers may identify sex in all Falconiformes. These are simple and rapid sexing methods, since only polymerase chain reaction (PCR) and agarose electrophoresis are required. In conclusion, sex identification by an alternative PCR approach based on intronic length variation and on differences in sequences between CHD1W and CHD1Z proved applicable to and useful for Falconiformes.     

Do sex ratio and development differ in sexually size‐dimorphic shiny cowbirds (Molothrus bonariensis) parasitizing smaller and larger hosts?

Two possible patterns of bias in primary sex ratio have been proposed for size‐dimorphic brood parasites that do not evict host chicks: (1) larger males should be laid at greater frequency in hosts larger than the parasite because they compete better (increasing their survival) than females with large host nest‐mates, and (2) more costly males (i.e. the larger sex) should be laid at greater frequency in hosts smaller than the parasite because, in these hosts, parasite nestlings are provisioned at a higher rate and grow faster than in larger hosts. We tested these hypotheses in two hosts of the sexually size‐dimorphic shiny cowbird, Molothrus bonariensis, one smaller (house wren, Troglodytes aedon) and one larger (chalk‐browed mockingbird, Mimus saturninus) than the parasite. We measured: (1) sex ratio at laying; (2) development of sexual differences in body mass during the nestling stage; and (3) chick survival and sex ratio of chicks before fledging. In both hosts, we found sexual differences in body mass of nestlings from 7 days of age onwards, although we did not find a bias in the sex ratio of eggs laid and chicks fledged. The results of the present study do not support the hypothesis that shiny cowbird females benefit from biasing the primary sex ratio depending on the size of the hosts they parasitize. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 442–448.     

Sexual size dimorphism and morphometric sexing in a North African population of Laughing Doves Spilopelia senegalensis

Like the majority of Columbiformes, the Laughing Dove Spilopelia senegalensis is sexually monomorphic in plumage, but seems to be slightly dimorphic in size. However, due to the lack of studies little is known about the sexual size dimorphism in this species. In this work, we used morphometric data on a sample of 61 Laughing Doves from southern Tunisia, and sexed using a DNA-based method, to assess size differences between males and females and to determine a discriminant function useful for sex identification. The results showed that wing length was the most dimorphic trait, which could be due to the effects of sexual selection. The best function for the discrimination between sexes included wing length and head length, which is comparable with findings on other dove species. This discriminant function accurately classified 89% of birds, providing a rapid and accurate tool for sex identification in the studied population. Further data from different populations are needed for firmer conclusions about the extent of sexual size dimorphism and the reliability of the morphometric sexing approach in this dove species.     

Morphological and genetic sex identification of white-tailed eagle Haliaeetus albicilla nestlings

Identifying the sex of bird nestlings is relevant to studies of behaviour and ecology and is often a central issue in the management of endangered or captive populations. The white-tailed eagle Haliaeetus albicilla is a formerly threatened Eurasian raptor which is closely monitored in many countries due to its high exposure to environmental pollutants in the food chain. The aim of this study was to evaluate the reliability of sex identification methods for white-tailed eagle nestlings based on morphological measurements that can be recorded at the nest by a single person and with minimum disturbance. The sex of each bird was independently determined using molecular (genetic) methods. One measure of tarsus width allowed the correct identification of sex for 96% of the nestlings from southern and central Sweden. However, we found that the criteria for sex identification were not directly applicable to the population in Swedish Lapland, where nestlings are typically thinner, probably due to a limited food supply. These results show that sexing in the field of white-tailed eagle nestlings can be feasible with high accuracy based on a limited number of measurements. However, the criteria employed to separate sexes may have to be adjusted for each population.     

Complex sex allocation in the laughing kookaburra

In groups of the cooperatively breeding laughing kookaburra(Dacelo novaeguineae), offspring sex varied with the type ofsocial group and with hatch rank. Groups with female helpers,especially if all helpers were female, had male-biased clutchand fledging sex ratios. Groups without female helpers (unassistedpairs or male-only helpers) had female-biased clutch and fledgingsex ratios. Breeding females responded facultatively to increasesin the number of female helpers in their group by producingmore male eggs. These biases may occur if breeding femalestry to limit the number of daughters recruited into their groupbecause unlike male helpers, female helpers depress the breedingsuccess of their parents. Across all nests, two-thirds of first-hatchedyoung were male, two-thirds of second-hatched young were female, and the sex ratio of third-hatched young was even. Hatch ranksex ratios also varied dramatically between different typesof social groups, from 16.7% for second-hatched nestlings ofunassisted pairs to 100% for first-hatched nestlings of groupswith only female helpers. A corollary of the relationship betweenhatch rank and sex was that hatching sex sequences were distributed nonrandomly: all groups avoided hatching a daughter first followedby a son (FM). Sibling competition is aggressive and sometimesfatal. Since females grow to be 15% larger than males the hatchingsequence of sexes could affect nestling growth and mortality.However, an exhaustive analysis found little evidence thatgrowth or survival of males was compromised if hatched aftera sister. The small number of FM sequences may only have occurredin nests that were able to ameliorate any negative consequences.Alternatively, when clutch size is small and fledging successunpredictable because of brood reduction, the preferred broodsex ratio may be contingent on the number of fledged young,making it advantageous to order the sexes in the brood.     

Androgynous rex - the utility of chevrons for determining the sex of crocodilians and non-avian dinosaurs

The sex of non-avian dinosaurs has been inferred on numerous occasions using a variety of anatomical criteria, but the efficacy of none has been proven. Nearly 50 years ago Romer suggested that the cranial-most or first chevron in the tails of some reptiles, including crocodilians, is sexually dimorphic. Recent work on this subject purportedly substantiated that the female first chevron articulates in a more caudal position than in males. Furthermore, it was concluded that this element is shorter in females. These phenotypic attributes theoretically provide a broader cloacal passageway for eggs by ovipositing females and a greater attachment area for male “penile retractor muscles”. Because theropod dinosaurs such as Tyrannosaurus rex presumably show similar variation in chevron anatomy, the same criteria has been advocated for sexing dinosaurs. We tested the neontological model for the chevron sexual dimorphism hypothesis using a skeletonized growth series of American alligators (Alligator mississippiensis) of known sex. No statistical support for the hypothesis was found. Furthermore, analysis of a diversity of crocodilian taxa from museum collections revealed similar findings suggesting the alligator results are not taxon specific. Study of well-preserved tyrannosaurid dinosaurs in museum collections showed nearly invariant chevron positioning like that seen in crocodilians. This suggests the usefulness of chevron anatomy for sexing dinosaurs is tenuous.     

Limits to sexual size dimorphism in red‐winged blackbirds: the cost of getting big?

A fundamental assumption of sexual selection theory is that the reproductive advantage of large size is balanced by a survival disadvantage. Previous studies of the sexually size-dimorphic red-winged blackbird ( Agelaius phoeniceus ) have indicated that the largest adult males have a survival advantage, suggesting that the limit to male size may be the cost of getting big rather than the cost of being big. If the cost of getting big limits male size, then starvation rates for male nestlings should exceed those of female nestlings. In addition, given high heritability of body size, larger parents should lose more nestlings, particularly males, to starvation. We tested these predictions for red-winged blackbirds using data on the sex of 1356 fledglings from 465 nests collected over 10 years. We found no disadvantage for male nestlings relative to females – 49% of fledglings were male and previous research had shown that 48% of hatchlings are male. We also found no disadvantage for male nestlings that would become large adults (i.e. those with larger parents) – partial brood loss and fledging sex ratios did not vary with mid-parent size. Given no apparent disadvantage to large size for males either as adults or as nestlings, this leaves only the period between fledging and adulthood during which natural selection might limit sexual size dimorphism, although other mechanisms might explain the failure to find a limit to male size.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 353–361.