Hatching order and sex ratio in Southern Grey Shrike Lanius meridionalis in relation to clutch size

Despite major advances in sex ratio theory, how offspring sex should vary with hatching order remains unclear. We examine nestling sex ratio in the Southern Grey Shrike Lanius meridionalis according to hatching order and clutch size. Southern Grey Shrike nestlings present a different sex ratio with body‐mass rank order depending on clutch size. When the clutch size was five eggs (with a very low risk of brood reduction; 13%) the less costly sex (male) was found at the end of the body mass hierarchy. However, when clutch size was six eggs (with a high risk of brood reduction; 42%) the larger sex (female) was found at intermediate positions in the hatching order, possibly to decrease competitive asymmetries.     

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.     

Hatching order affects offspring growth,survival and adult dominance in the joint‐laying Pukeko Porphyrio melanotus melanotus

In birds with asynchronous hatching, hatching order is an important factor in determining offspring phenotype. Many previous studies have demonstrated that later‐hatched offspring show reduced growth and survival during development. However, few studies have followed individuals from hatching to adulthood to test whether the effects of hatching order persist into later life. Here, we explore patterns of hatching order and fitness‐related traits in the Pukeko Porphyrio melanotus melanotus, a cooperatively breeding bird that lives in stable social groups that form linear dominance hierarchies. Pukeko groups sometimes contain two breeding females that lay eggs in the same nest (joint‐laying). Thus, competition between nest‐mates can influence the relative fitness of each laying female. We show that in both single‐clutch and joint‐clutch nests, earlier‐hatched Pukeko chicks grow faster and survive better than later‐hatched brood‐mates. Moreover, earlier‐hatched chicks achieve higher dominance ranks as adults, making this study one of the first to find a relationship between hatching order and adult dominance in wild birds. Finally, we show that in groups with two breeding females, the chicks of the primary female hatch earlier than the chicks of the secondary female. As a result, the offspring of the primary female may be at a competitive advantage, which could have important implications for social dynamics in this species.     

Experimental evidence for parental,but not parentally biased,favouritism in relation to offspring size in Blue Tits Cyanistes caeruleus

In species with biparental care, males and females share the benefits of investing in offspring but pay the costs individually. As a result of these evolutionary conflicts of interest between the sexes, it is expected that the two parents should follow different behavioural rules when providing food to the young. Such a discrepancy may be accentuated when parents have to choose between different subsets of offspring (e.g. large and small nestlings). We manipulated the degree of hatching asynchrony in Blue Tits Cyanistes caeruleus and quantified male and female feeding behaviour when nestlings were 7 and 10 days old. First, we tested for a difference in the role of the sexes during the nestling rearing period between experimentally asynchronous and synchronous control broods. We then used experimentally asynchronous broods to assess differences between the sexes in the pattern of food distribution in terms of number of feedings and prey types, between junior and senior siblings. When nestlings in experimental nests were 7 days old, females fed young more often than did males despite facing a trade‐off between brooding the smallest nestlings and bringing food to the nest. At this age, there was also a skew in food delivery in favour of senior siblings, whereas food was more evenly distributed across the brood when nestlings were 10 days old. We found no difference in how male and female Blue Tits distributed feeding visits among junior and senior nestlings. However, females fed the smallest nestlings with more spiders in comparison with their senior siblings. This could be related to their more suitable size relative to other prey types, their high content of essential nutrients, or both, and may represent a more cryptic form of parentally biased favouritism. We compare these findings with previous work on other species and discuss why parents did not feed junior siblings more frequently.     

Growth and survival of neotropic cormorant (Phalacrocorax brasilianus) chicks in relation to hatching order and brood size

We investigated chick development and feeding rate in the neotropic cormorant, Phalacrocorax brasilianus, in a colony in Central Chile. The year of our study was characterized by relatively good foraging conditions. Brood sizes varied from two to five chicks, and hatching was asynchronous, with gaps of 0 to 6 days between the youngest and the oldest chick. Egg size declined over laying order in three-egg clutches, but not in four-egg clutches. Hatch weight did not vary with hatching position, irrespective of brood size. Chicks increased mass on average by 60 g/day between 8 and 20 days of age. Growth rates and survival to fledging depended on hatching position only in broods of four, where D-chicks grew slower and showed a higher pre-fledging mortality. There was a non-significant tendency that also A-, B-, and C-chicks in broods of four grew slower than in smaller broods. Average number of fledglings was 2.76. Feeding frequency decreased with chick age between the ages of 10–40 days. Four-chick broods received more feeds per day than smaller broods, leading to a similar per-chick feeding frequency across all brood sizes. D-chicks were clearly disadvantaged in growth and survival, and facultative brood reduction occurred.     

Hatching plasticity in a Southeast Asian tree frog mitigates submergence-induced mortality

Environmentally cued hatching has been well-documented in amphibians in response to a wide range of abiotic and biotic factors. The hatching of terrestrial amphibian eggs in response to flooding may be basal within the group, but amphibian lineages in tropical Asia and sub-Saharan Africa have not received as much attention as their Neotropical counterparts. We investigated submergence-induced hatching in Feihyla hansenae, a Rhacophorid tree frog with terrestrial eggs. We quantified natural rates of clutch submergence at our study site in Thailand. Using submergence experiments, we found that embryos are capable of hatching early to escape flooding, and that failure to hatch results in mortality. Among the embryos that were able to hatch early, only the earliest, youngest hatchlings experienced a trade-off in body size that persisted for 6 days, while later, older hatchlings were not significantly smaller than spontaneous hatchlings under control conditions. By incorporating our natural and experimental data into Monte Carlo methods to simulate and compare survival probabilities with and without hatching plasticity, we found an overall 3.1% increase in submergence survival due to hatching plasticity. Our findings support the idea that flooding-induced hatching is widespread across amphibians with terrestrial eggs and highlight the importance of researching understudied tropical regions. As climate change is projected to affect rainfall patterns, the ability of embryos to escape abiotic egg-stage threats may be an indicator of species' ability to flexibly navigate a changing environment.     

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.     

The evolution of obligate interspecific brood parasitism in birds

We present a simple analytical model to investigate the conditionsfor the evolution of obligate interspecific brood parasitismin birds, based on clutch size optimization, when birds canlay more eggs than their optimal clutch size. The results showthat once intraspecific parasitism has appeared (i.e., femalesstart to spread their eggs over their own and other nests) the evolutionarily stable number of eggs laid in its own nest decreases.Two possible ESSs exist: (1) either the evolutionarily stablenumber of eggs laid in its own nest is larger than zero, anda fraction of the total number of eggs is laid parasitically(i.e., intraspecific parasitism); and (2) either the evolutionarilystable number of eggs laid in its own nest is zero and all eggs are laid parasitically. Since all females lay parasitically,this could favor the evolution of obligate interspecific broodparasitism. The key parameter allowing the shift from intraspecificto obligate interspecific parasitism is the intensity of density-dependentmortality within broods (i.e., nestling competition). Strongnestling competition, as in altricial species, can lead toan ESS where all eggs are laid parasitically. Altricial speciesare, therefore, predicted to evolve more easily toward obligate interspecific parasitism than precocial species. These predictionsfit the observed distribution of brood parasitism in birds,where only one species out of 95 obligate interspecific parasitesexhibits a precocial mode of development. Different nestlingsurvival functions provided similar findings (i.e., obligatebrood parasitism is more likely to evolve in altricial species),suggesting that these results are robust with respect to themain assumption of the model.     

Discrepancy between factors affecting nestling growth and survival and maternal success in Common Grackles

ABSTRACT Multiple factors potentially affect nestling survival and maternal reproductive success. However, little is known about the relative importance of different factors when operating simultaneously or whether the same factors are important for nestlings and their mothers. We determined the effect of hatching asynchrony, individual egg size, mean egg size, nestling sex, and clutch initiation date on the survival of individual nestlings and on maternal reproductive success in Common Grackles (Quiscalus quiscula) from 2004 to 2006 in central Illinois. Factors most important to maternal success differed from those important for individual nestling growth and survival. Hatching asynchrony had the greatest within‐nest influence on the fate of nestlings; the earlier a nestling hatched relative to siblings, the greater its mass and likelihood of fledging. Clutch size had the greatest influence on maternal reproductive success, with females with larger clutches fledging more young. Thus, both nestling survival and maternal success were largely determined by a single, albeit different, factor. A possible explanation for the apparent unimportance of most factors we measured in determining maternal success is that we did not consider variation among females. Individual variation in maternal attributes such as condition, size, age, experience, or mate quality may result in females tailoring clutch attributes (i.e., egg size, sex, and degree of hatching asynchrony) in ways that allow them to maximize their reproductive success. The discordance between factors that benefited mothers versus their offspring illustrates the importance of considering the maternal consequences of any factor that appears to affect offspring survival. Factors that increase the mass and survival of some offspring may not result in increased maternal reproductive success.     

Within clutch co-variation of egg mass and sex in the black-headed gull

Female birds of several species have control over the production of daughters and sons. However, most studies failed to find a relationship between egg size and sex. This is intriguing as adjustment of egg size would constitute a powerful tool for the female to meet different resource demands of the sexes, particularly in size dimorphic species. Our results show that, within clutches of black-headed gulls (Larus ridibundus) the proportion of males was positively associated with egg mass. This applied for all three laying positions, independently of the absolute egg mass. There was a significant relationship between the distribution of the sexes over the laying sequence and the egg mass change. When egg mass decreased over the sequence, first-laid eggs were male biased and last-laid eggs female biased, and vice versa. The potential adaptive value of this allocation strategy is evaluated with regard to male sensitivity to egg quality and competitive differences between the sexes.     

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.     

Variation in incubation effort during egg laying in the Mountain Bluebird and its association with hatching asynchrony

Females in many bird species reportedly begin incubation prior to clutch completion, but the nature of such incubation and the degree to which it varies among females remains undescribed for almost all species. We used continuous recording of nest‐cup temperatures to document incubation effort during egg laying at 57 Mountain Bluebird (Sialia currucoides) nests in a high‐elevation Wyoming population. We then asked whether such effort predicted the degree to which eggs hatch asynchronously. Although substantial egg heating could begin abruptly late in laying (previously reported as the norm for this species) or even after clutch completion, we found that most (>90%) females began incubation gradually, engaging in a few (usually 1–8), brief (<10 min) bouts of heating on the day they laid their first or second egg. Thereafter, females varied markedly in when they increased incubation effort and by how much. The onset of nocturnal incubation also varied, with females beginning to incubate at night after laying their prepenultimate, penultimate, or last egg and not always initially incubating through the night. As an index of the total amount of heat applied to eggs during laying, we calculated the cumulative number of degrees by which nest‐cup temperatures exceeded the threshold temperature required for embryonic development. This value varied by more than 150‐fold between nests and explained >50% of the variation in hatching asynchrony. Our results thus provide strong support for the widely held, but rarely tested, assumption that parent birds can have substantial control over the degree of hatching asynchrony by varying the amount of incubation done prior to clutch completion.     

Length of herring larvae in relation to age and time of hatching

The lengths of newly hatched herring Clupea harengus larvae were generally larger than the lengths of embryos released from eggs on the same day. Over subsequent days, the daily increase in length was higher in groups that were released from the eggs as opposed to groups that hatched naturally. Thus by the end of the hatching period (18 days after fertilization) there were no significant differences in mean lengths.     

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.     

Parasites favour intermediate nestling mass and brood size in cliff swallows

A challenge of life‐history theory is to explain why animal body size does not continue to increase, given various advantages of larger size. In birds, body size of nestlings and the number of nestlings produced (brood size) have occasionally been shown to be constrained by higher predation on larger nestlings and those from larger broods. Parasites also are known to have strong effects on life‐history traits in birds, but whether parasitism can be a driver for stabilizing selection on nestling body size or brood size is unknown. We studied patterns of first‐year survival in cliff swallows (Petrochelidon pyrrhonota) in western Nebraska in relation to brood size and nestling body mass in nests under natural conditions and in those in which hematophagous ectoparasites had been removed by fumigation. Birds from parasitized nests showed highest first‐year survival at the most common, intermediate brood‐size and nestling‐mass categories, but cliff swallows from nonparasitized nests had highest survival at the heaviest nestling masses and no relationship with brood size. A survival analysis suggested stabilizing selection on brood size and nestling mass in the presence (but not in the absence) of parasites. Parasites apparently favour intermediate offspring size and number in cliff swallows and produce the observed distributions of these traits, although the mechanisms are unclear. Our results emphasize the importance of parasites in life‐history evolution.     

Sib-mating and sex ratio strategies in scelionid wasps

Abstract. 1. Theoretical models by Hamilton and others predict that the optimal sex ratio (proportion males) for a species should decrease with increasing levels of sib-mating. A test of this hypothesis is made for members of the parasitoid family Scelionidae.
2. Scelionid wasps differ in levels of sib-mating as a consequence of differences in host egg mass size, male—male aggression and female—female aggression.
3. Sex ratios for thirty-one species vary with inferred changes in sib-mating as predicted by the models.
4. Interspecific differences in sex ratio may represent different fvted strategies or different manifestations of a single variable strategy (sex ratio game) in response to parasitoid-produced stimuli.
5. Males and females are assigned non-randomly to egg masses in a manner which ensures mixed broods in the proper proportions for a particular species.     

Do females adjust the sex of their offspring in relation to the breeding sex ratio?

When the adult sex ratio differs between years in local populations, but still is predictable between adjacent years, it has been proposed that the best strategy would be to bias the offspring sex ratio in favour of the rare sex. We tested this hypothesis using a data set of great reed warbler offspring, sexed by molecular techniques, that were collected over 11 breeding seasons at two adjacent reed marshes. Three important assumptions for this hypothesis are fulfilled in the studied great reed warbler population. First, a substantial proportion of great reed warblers are living in small local populations where sex ratio distortions would be sufficiently large and common. Second, breeding adults and their offspring return to breed in the local population to a high degree. Third, females have a possibility to assess the breeding sex ratio before laying their eggs. At our study site, the breeding sex ratio was positively correlated between successive years. However, contrary to our prediction, female great reed warblers seemed not to adjust their offspring sex ratio in relation to the local breeding sex ratio.     

Cold season mortality and hatching behavior of Aedes aegypti L. (Diptera: Culicidae) eggs in Buenos Aires City,Argentina

In temperate regions, populations of Aedes aegypti survive the cold season in the egg stage. In the present work, we studied the cold‐season mortality of Ae. aegypti eggs and their subsequent hatching pattern in Buenos Aires city. Eggs were exposed during the winter season (three months) in three neighborhoods located along a gradient of distance toward the Río de la Plata River, coincident with a gradient of activity of Ae. aegypti. Results showed mortalities lower (30.6%) than those from tropical regions during the dry season. Significant differences were detected among the egg mortalities of each site with a maximum value at the site nearest the Río de la Plata River (50%), and a minimum value at the most continental site (9%). Post‐experimental hatching response of eggs differed between sites, with the highest proportion of hatched eggs during the first immersion in the site nearest to the river and the lowest proportion in the most continental site. The hatching proportion also differed between age classes, with older (early‐laid) eggs hatching later than new (late‐laid) ones. Our results provide the first information of Ae. aegypti egg mortality in temperate South America and support the hypothesis that differences in egg mortality are associated with abundance patterns of Ae. aegypti in Buenos Aires city.