Influence of hatching order on growth rate and resting metabolism of kestrel nestlings

Hatching asynchrony in altricial birds may result in a competitive disadvantage for the youngest nestlings compared to older siblings. We studied the effects of a size hierarchy on the growth rate of Eurasian kestrels Falco tinnunculus chicks in nests with and without access to supplemented food in western Finland. Body mass stopped increasing on the 19th day after hatching while body size, estimated by a combination of bone and feather lengths continued to increase at least until fledging at 26 days. Body condition, reflecting muscle and fat, did not change markedly during the growth period from the 12th day to fledging. Body temperature and resting metabolism were usually lower in nestlings 12 days old than in nestlings at fledging. Growth of body mass, size and condition, and resting metabolism were delayed in last-hatched nestlings aged 19 days. Just before fledging, last-hatched nestlings attained a similar body mass and size, and had a similar resting metabolism to those of older siblings. At fledging, only in nests without access to supplemented food was the body condition of last-hatched chicks lower than that of its siblings, but in nests with access to supplemented food no such difference was detected. Our results highlight that the level of lipids in the last-hatched nestling can be affected by the food restriction imposed by hatching order.     

Maternally derived yolk testosterone enhances the development of the hatching muscle in the red-winged blackbird Agelaius phoeniceus

Hatching asynchrony in avian species often leads to the formation of a size hierarchy that places last-hatched nestlings at a significant disadvantage. The hatching muscle (musculus complexus) is responsible for breaking the shell during hatching and for dorsal flexion of the neck during begging. An increase in its strength in last-hatched nestlings could mitigate the effects of hatching asynchrony by reducing the time required for hatching or enhancing the effectiveness of begging for parentally delivered food or both. We have previously found that yolk testosterone concentration increases with laying order in the red-winged blackbird Agelaius phoeniceus. In this study, we investigated the hypothesis that yolk testosterone has anabolic effects on the development of the complexus, thereby influencing competition among asynchronously hatched nestlings. We found that both yolk testosterone concentration and relative complexus mass (complexus mass/nestling body mass) increased with laying order and that these two variables were positively correlated in both newly hatched nestlings and in two-day-old broods. Moreover, direct injections of testosterone into egg yolks resulted in an increase in relative complexus mass, while injections of flutamide, a testosterone antagonist, resulted in a decrease in relative complexus mass. Neither yolk testosterone concentration nor relative complexus mass differed between male and female nestlings.     

Age and food deprivation affects expression of the glucocorticosteroid stress response in Magellanic penguin (Spheniscus magellanicus) chicks

We examined how the glucocortical stress response in free-living Magellanic penguin (Spheniscus magellanicus) chicks changes with age and whether adrenocortical function of chicks within a brood varies in relation to food provisioned by adults. Chicks showed little corticosterone response to capture stress shortly after hatching, an intermediate response around 45-d posthatch, and a robust stress response near fledging. However, in response to an adrenocorticotropic hormone (ACTH) challenge, hatchlings were capable of secreting corticosterone at adult-like levels. The larger sibling in broods of two showed a similar gradual stress-response development pattern. In contrast, by day 45, when differences in body condition were well established between siblings, the smaller, food-deprived chicks significantly increased baseline levels of corticosterone but showed normal stress-induced levels. Near fledging, baseline levels had returned to normal, but stress-induced levels were lower than expected. Similar to altricial species, normally developing semialtricial Magellanic penguin chicks do not express a robust corticosterone stress response until near fledging. Chronic stressors such as food deprivation cause corticosterone use to be up-regulated earlier than expected. However, in cases of extended chronic stress, down-regulation may ensue, thus avoiding the negative effects of chronically elevated levels of corticosterone.     

Dietary restriction causes chronic elevation of corticosterone and enhances stress response in red-legged kittiwake chicks

Release of corticosterone in hungry kittiwake chicks facilitates begging and allows them to restore depleted energy reserves by increasing parental food provisioning. However, in order to avoid detrimental effects of chronic elevation of corticosterone, chicks might suppress adrenocortical activity in response to prolonged food shortages. In this study we examined temporal dynamics of corticosterone release in red-legged kittiwake (Rissa brevirostris) chicks exposed to prolonged restrictions in energy content and/or nutritional quality (low versus high lipid content) of their food. Starting at the age of 15 days, chicks were fed either high- or low-lipid fish at 40%, 65%, and 100% of ad libitum energy intake. Body mass measurements and baseline plasma samples were taken on a weekly basis after beginning of the treatment. After 3 weeks of treatment, chicks were exposed to a standardized acute handling and restraint stress protocol, where in addition to a baseline sample, three plasma samples were taken at intervals up to 50 min. We found that food-restricted chicks had lower body mass, chronically (during 2-3 weeks) elevated baseline and higher acute stress-induced levels of corticosterone compared to chicks fed ad libitum. Low lipid content of food further exacerbated these effects. An increase in baseline levels of corticosterone was observed within a week after energy requirements of food-restricted chicks exceeded their daily energy intake. A tendency for suppression of adrenocortical activity was observed in treatments fed low-lipid diets only at the end of the experiment. We suggest that nest-bound chicks, if food-stressed, might suffer deleterious effects of chronic elevation of corticosterone.     

Brood Reduction via Intra-clutch Variation in Testosterone - An Experimental Test in the Great Tit

In birds, yolk androgen concentrations in eggs can increase or decrease over the laying sequence and common hypotheses hold that this serves to favour the competitive ability of either first- or last-hatched chicks depending on the prevailing conditions, and thus promote brood reduction or maintenance of original brood size respectively. Intra-clutch variation of testosterone can shift relative competitive ability of siblings and hence competitive dynamics. In a natural population of great tits, we experimentally investigated the effects and function of maternal testosterone on offspring phenotype in relation to the laying position of the egg in a context of hatching asynchrony. To this end, we created three types of clutches where either the first three or the last three eggs of a clutch were injected with testosterone (T) dissolved in sesame oil, and the remaining eggs with sesame oil only, or where all eggs of a clutch were injected with sesame oil. Increased levels of yolk T in the last-laid eggs resulted in the last-hatched chicks being significantly lighter and smaller than their siblings, while increased levels of T in the first-laid eggs had no direct effect on the first-hatched chicks, but an indirect negative effect on their siblings. Our results suggest that females can potentially adjust offspring phenotype by modulating, over the laying sequence, the amounts of T deposited in the eggs. These results are in contradiction, however, with current hypotheses and previous findings, which suggest that under good conditions higher levels of maternally derived T in the last-laid eggs should mitigate the negative effects of hatching asynchrony.     

Hatching asynchrony,sibling hierarchies and brood reduction in the Chinstrap penguin Pygoscelis antarctica

We studied patterns of chick growth and mortality in relation to egg size and hatching asynchrony during two breeding seasons (1991 and 1992) in a colony of chinstrap penguins sited in the Vapour Col rookery, Deception Island, South Shetlands. Intraclutch variability in egg size was slight and not related to chick asymmetry at hatching. Hatching was asynchronous in 78% (1991) and 69% (1992) of the clutches, asynchrony ranging from 1 to 4 days (on average 0.9 in 1991 and 1.0 days in 1992). Chicks resulting from oneegg clutches grew better than chicks in families of two in 1991. In 1992, single chicks grew to the same size and mass at 46 days of age as chicks of broods of two, suggesting food limitation in 1991 but not in 1992. In 1991, asymmetry between siblings in mass and flipper length was significantly greater in asynchronous than in synchronous families during the initial guard stage, but these differences disappeared during the later créche phase. In 1992, asymmetry in body mass increased with hatching asynchrony and decreased with age. Only the effect of age was significant for flipper length and culmen. Asymmetries at 15 days were similar in both years, but significantly lower in 1992 than in 1991 at 46 days of age. There were relatively frequent reversals of size hierarchies during both phases of chick growth in the two years, reversals being more common in 1991 than in 1992 for small chicks. In 1991, survivors of brood reduction grew significantly worse than chicks in nonreduced broods. In both years, chicks of synchronous broods attained similarly large sizes before fledging as both A and B chicks of asynchronous broods. In 1991, chick mortality rate increased during the guard stage due to parental desertions, decreased during the transition to crèches (occurs at a mean age of 29 days) and returned to high constant levels during the crèche stage, when it is mostly due to starvation (in total 66% of hatched chicks survived to fledging). In contrast, in 1992, mortality was relatively high immediately after hatching and almost absent for chicks older than 3 weeks (87% of chicks survived to fledging). Mortality affected similarly one- and two-chick families. In 1991, asynchronous families suffered a significantly greater probability of brood reduction than synchronous families, but this probability was not significantly related to degree of asymmetry between siblings. No association between asynchrony and mortality was found in 1992. These results show that there is food limitation in this population during the crèche phase in some years, that asynchronous hatching does not facilitate early brood reduction and that it does not ensure stable size hierarchies between siblings. Brood reduction due to starvation is not associated to prior asymmetry and does not facilitate the survival or improve the growth of the surviving chick. Asynchronous hatching may be a consequence of thermal constraints on embryo development inducing incubation of eggs as soon as they are laid.     

Complex patterns of food allocation in asynchronously hatching broods of crimson rosellas

In birds, asynchronous hatching typically leads to lower growth and survival of last-hatched chicks. However, all crimson rosella Platycercus elegans, chicks grow at the same rate, although first-hatched chicks can be as much as seven times heavier than last-hatched chicks at the end of hatching. We examined the delivery and distribution of food to 18 rosella broods by videotaping feeds and simultaneously recording mass changes in the nestbox using a digital balance. Parents visited the nest infrequently and delivered loads of up to 25% of their body weight during a feeding visit. Male rosellas consistently delivered larger loads and consequently had higher feeding rates (g/h) than females. Parents distributed food between chicks by direct regurgitation in a series of up to 51 food transfers. Overall, chicks of all hatching ranks received equal numbers of transfers, but parents differed in how they distributed food within the brood. Males fed first-hatched chicks more than last-hatched chicks, whereas females distributed food equally to all chicks. Selective feeding of small chicks might be costly to females since they delivered food more slowly than males and spent more time in the nestbox. Thus female rosellas may invest more in current reproduction than males. Parents also distributed food differently to male and female chicks. Large males were fed more than all other nestlings, while female nestlings were fed equally irrespective of size. This study confirms that complex patterns of parental allocation occur in wild populations. Copyright 1999 The Association for the Study of Animal Behaviour.     

Effects of intraclutch variation in egg size and hatching asynchrony on nestling development and survival in semi‐precocial Herring Gulls

Intraclutch egg size variation may non‐adaptively result from nutritional/energetic constraints acting on laying females or may reflect adaptive differential investment in offspring in relation to laying/hatching order. This variation may contribute to size hierarchies among siblings already established due to hatching asynchrony, and resultant competitive asymmetries often lead to starvation of the weakest nestling within a brood. The costs in terms of chick mortality can be high. However, the extent to which this mortality is egg size‐mediated remains unclear, especially in relation to hatching asynchrony which may operate concomitantly. I assessed effects of egg size and hatching asynchrony on nestling development and survival of Herring Gulls (Larus argentatus), where the smaller size and later hatching of c‐eggs may represent a brood‐reduction strategy. To analyze variation in egg size, I recorded the laying order and laying date of 870 eggs in 290 three‐egg clutches over a 3‐yr period (2010–2012). I measured hatchlings and monitored growth and survival of 130 chicks from enclosed nests in 2011 and 2012. The negative effect of laying date (β = ?0.18 ± SE 0.06, P = 0.002) on c‐egg size possibly reflected the fact that late breeders were either low quality or inexperienced females. The mass, size, and condition of hatchling Herring Gulls were positively related to egg size (all P < 0.0001). C‐chicks suffered from increased mortality risk during the first 12 d, identified as the brood‐reduction period in my study population. Although intraclutch variation in egg size was not directly related to patterns of chick mortality, I found that smaller relative egg size interactively increased differences in relative body condition of nestlings, primarily brought about by the degree of hatching asynchrony during this brood‐reduction period. Thus, the value of relatively small c‐eggs in Herring Gulls may lie in reinforcing brood reduction through effects on nestling body condition. A reproductive strategy Herring Gulls might have adopted to maintain a three‐egg clutch, but that also enables them to adjust the number of chicks they rear relative to the prevailing environmental conditions and to their own condition during the nestling stage.     

Larval sensory abilities and mechanisms of habitat selection of a coral reef fish during settlement

In most animals, siblings from a given reproductive event emerge over a very short period of time. In contrast, many species of birds hatch their young asynchronously over a period of days or weeks, handicapping last-hatched chicks with an age and size disadvantage. Numerous studies have examined the adaptive significance of this atypical hatching pattern, but few have attempted to explain the considerable intrapopulation variation that exists in hatching asynchrony. I explored proximate determinants of hatching asynchrony by monitoring 112 Burrowing Owl (Athene cunicularia) nests in the grasslands of southern Saskatchewan, Canada, over 4 years. Age disparities between first- and last-hatched siblings (i.e., hatching spans) varied considerably, ranging between 1 and 7 days (mode = 4 days). These hatching spans increased with increased hatching success. Hatching spans also increased with larger clutches, but the increase was less than predicted given the increased time required to lay more eggs. Hatching span was unrelated to number of prey cached in the nest during egg laying (an index of food availability), and was unaltered by a year of super-abundant prey. Furthermore, pairs given extra food during laying had hatching spans equal to those of unsupplemented control pairs. These results were inconsistent with both the energy constraint and facultative manipulation hypotheses, which predict that hatching asynchrony should vary with the level of food during laying, when incubation onset is determined. Burrowing Owls were apparently free of food limitation early in breeding, yet may not have been able to optimize hatching spans because food conditions during laying were largely unrelated to food conditions during brooding. Thus, one of the premises for facultative manipulation of hatching asynchrony—that laying females are able to forecast post-hatch food conditions—may not have been met for this population of Burrowing Owls.     

Food allocation in crimson rosella broods: parents differ in their responses to chick hunger

Food allocation in many asynchronously hatching bird species favours large, competitively superior chicks. In contrast, food is usually distributed equally within broods of crimson rosellas, Platycercus elegans, implying that parents do not simply feed the most competitive chick. We used two temporary removal experiments to manipulate hunger of: (1) individual first- or last-hatched chicks, or (2) the whole brood. When only a single chick was hungry, parents compensated fully and chicks gained the same mass over the day as during controls. Mothers and fathers, however, responded in different ways to chick hunger. Mothers did not strongly alter their food allocation when a single chick was hungry, and controlled the distribution of food by refusing to feed first-hatched chicks when they were hungry and by moving more during feeds. In contrast, fathers allocated more food to hungry last-hatched chicks. When the whole brood was hungry, parents were unable to compensate chicks and all chicks lost mass over the day. In these conditions, mothers preferentially fed first-hatched chicks, while fathers fed all chicks equally. Our results show that both mothers and fathers were able to discriminate and selectively feed chicks, but that parents responded differently to changes in chick hunger within the brood. Fathers responded more strongly to variation in chick hunger within the brood, suggesting they reallocate food based on short-term changes in hunger. Mothers distributed food preferentially to last-hatched chicks except when the whole brood was hungry, when they switched to favouring first-hatched chicks. This pattern is consistent with a strategy of adaptive brood reduction when food is scarce. Copyright 2000 The Association for the Study of Animal Behaviour.     

Adrenocortical responses in zebra finches (Taeniopygia guttata): individual variation, repeatability, and relationship to phenotypic quality

Although individual variation is a key requirement for natural selection, little is known about the magnitude and patterns of individual variation in endocrine systems or the functional significance of that variation. Here we describe (1) the extent and repeatability of inter-individual variation in adrenocortical responses and (2) its relationship to sex-specific phenotypic quality, such as song duration and frequency and timing of egg laying. We measured adrenocortical responses to a standardized stressor in zebra finches (Taeniopygia guttata) at two life history stages: approximately day 16 (nestlings) and 3 months of age (sexually mature adults). Subsequently, we assessed phenotypic (reproductive) quality of all individuals as adults. Marked inter-individual variation in the adrenocortical response was seen in both sexes and ages, e.g., stress-induced corticosterone ranged from 2.2 to 62.5 ng/mL in nestlings and 5.0-64.0 ng/mL in adults. We found sex differences in (a) inter-individual variation in the adrenocortical response, (b) repeatability, and (c) relationships between corticosterone levels and phenotypic quality. In males, variation in nestling corticosterone was weakly but positively correlated with brood size and negatively correlated with nestling mass (though this relationship was dependent on one individual). There was no significant correlation of adrenocortical responses between two stages in males and adult phenotypic quality was significantly correlated only with adult corticosterone levels. In contrast, in females there was no relationship between nestling corticosterone and brood size or mass but adrenocortical response was repeatable between two stages (r2=0.413). Phenotypic quality of adult females was correlated with nestling baseline and adrenocortical response.     

Leucocyte profiles and corticosterone in chicks of southern rockhopper penguins

The immune system is essential for health and survival of vertebrates, yet still little is known about the ontogeny of the immune system in wild birds. The southern rockhopper penguin (Eudyptes chrysocome chrysocome) is a semi-altricial seabird with a long developmental period and reversed hatching asynchrony, favouring the survival of B-chicks. We compared leucocyte counts and baseline corticosterone levels of southern rockhopper penguin chicks under different preconditions such as sex and origin from an A- or B-egg from 4 to 51 days of age. We conducted an experiment to compare leucocyte profiles and baseline corticosterone levels in A- and B-chicks in single-egg clutches as well as in B-chicks from normal two egg-clutches (one A- and one B-egg). None of these treatments influenced leucocyte counts or corticosterone levels, indicating a similar investment in the immune system. Our main finding was an increase of leucocytes/10,000 erythrocytes with age, which was mainly caused by an increase in lymphocyte numbers. This suggests differential investment into acquired immunity at this stage of development. As the granulocyte/lymphocyte (G/L) ratio did not change with age or body condition, G/L ratios seem not to reflect stress caused by poor provisioning of penguin chicks. This was also reinforced by the decrease of plasma corticosterone levels with age. Body condition was negatively correlated with monocyte numbers, suggesting a poorer health status of penguin chicks in poorer body condition. Yet, there was no link between body condition and other leucocyte parameters, indicating that chicks in a good body condition did not additionally invest into their immune system.     

Lesser double‐collared sunbirds Nectarinia chalybea do not compensate for hatching asynchrony by adjusting egg mass or yolk androgens

Substantial amounts of maternal androgens are found in birds’ eggs and have been shown to benefit offspring development. Within‐clutch patterns of increasing androgen concentrations over the laying sequence are often hypothesized to compensate for the negative effects of hatching asynchrony. However, detrimental effects to offspring fitness of exposure to high yolk androgen levels have also been demonstrated. This suggests that mothers should forego these costs to their offspring when the need for compensation for hatching asynchrony is low or when alternative compensatory strategies, e.g. in terms of increasing egg mass, are available. Here we show that in the south‐temperate lesser double‐collared sunbird Nectarinia chalybea, a species with hatching asynchrony but also with high survival of last‐hatched chicks, mothers do not deposit resources differentially in terms of either yolk androgen concentration or egg mass across the laying sequence. We discuss to what extend this challenges the original explanation of within‐clutch variation in these egg parameters and offer some explanation for their between‐clutch variation which was related to female body mass.     

Asynchronous hatching in the laughing gull: Cutting losses and reducing rivalry

The adaptive significance of hatching asynchrony was investigated in the laughing gull (Larus atricilla). Staggered hatching of the brood is frequently correlated with reduced survival in the later-hatching chicks, and the phenomenon has therefore been the subject of speculation about adaptive function. The relative timing of hatching within the brood was experimentally manipulated in order to compare the effects of synchrony versus asynchrony on parental reproductive success. The average number of young fledging per nest was significantly higher in the asynchronous group. Staggered hatching of the brood is hypothesized to be a parental strategy offering the option of brood reduction under conditions of food shortage. The possibility is discussed that asynchronous hatching benefits parents independently by reducing sibling rivalry over parental investment and minimizing wasteful competition.     

Family dynamics through time: brood reduction followed by parental compensation with aggression and favouritism

Parental food allocation in birds has long been a focal point for life history and parent–offspring conflict theories. In asynchronously hatching species, parents are thought to either adjust brood size through death of marginal offspring (brood reduction), or feed the disadvantaged chicks to reduce the competitive hierarchy (parental compensation). Here, we show that parent American coots (Fulica americana) practice both strategies by switching from brood reduction to compensation across time. Late‐hatching chicks suffer higher mortality only for the first few days after hatching. Later, parents begin to exhibit parental aggression towards older chicks and each parent favours a single chick, both of which are typically the youngest of the surviving offspring. The late‐hatched survivors can equal or exceed their older siblings in size prior to independence. A mixed allocation strategy allows parents to compensate for the costs of competitive hierarchies while gaining the benefits of hatching asynchrony.     

Body condition is associated with adrenocortical response in the barn swallow (Hirundo rustica L.) during early stages of autumn migration

Migration is an energy-demanding life-history period and also a significant population-limiting factor of long-distance migratory birds. It is important to understand how corticosterone, the main energy regulating hormone in birds, is associated with behavioural and physiological changes during migration. According to the migration modulation hypothesis (MMH), individual birds may express elevated levels of baseline corticosterone to facilitate fuelling, but down-regulate the adrenocortical response in order to protect skeletal muscles from the catabolic effects of the hormone. We measured the baseline and stress-induced levels of corticosterone in barn swallows (Hirundo rustica L.) during early stages of autumn migration. Here, we show that, while barn swallows clearly responded to the capture and handling stress by increasing the corticosterone level, the strength of this acute response was related to their energetic condition: birds with high body mass responded more rapidly and had lower peak values of corticosterone than lighter birds. Further, the baseline levels of corticosterone correlated negatively with the magnitude of the adrenocortical response. Barn swallows did not show elevated baseline levels of corticosterone in the course of autumn, which suggests that, instead of fuelling, the birds were actively migrating. Our results indicate that MMH also applies to aerial feeders, whose foraging habits differ from model birds of previous studies.     

Hormonal correlates of siblicide in Galápagos Nazca boobies

Nazca boobies (Sula granti) show unconditional obligate siblicide immediately after hatching, reducing the typical two-egg clutch size to one. We studied body mass changes and levels of testosterone (T), corticosterone (CORT), and progesterone (P) for A-chicks (dominant, first hatched), B-chicks (subordinate, second hatched), and singletons, during the first 7 days after hatching, when siblicide normally occurs. Mass increase with age was higher for A-chicks than for singletons and B-chicks. This exaggerated the existing developmental advantage of A- over B-chicks that is due to hatching asynchrony. In nests with two chicks, CORT titer was significantly higher in B-chicks than in A-chicks. During ontogenetic development, CORT decreased with age for A-chicks, but did not change for singletons. P showed qualitatively similar ontogenetic changes to CORT, remaining unchanged for A-chicks but increasing for singletons. Thus, both CORT and P levels were lower for A-chicks than for singletons, and both hormones varied inversely with body mass. Overall, T levels did not differ between different categories of chicks. However, one B-chick in the process of reversing the dominance relationship with its older, but weakened, sibling had significantly elevated T. We suggest that CORT and P are regulated to promote exaggerated mass gain in socially challenged A-chicks, facilitating siblicide. Whether T induces aggressiveness during short time intervals of intense sibling rivalry needs further attention.     

Maternal compensation for hatching asynchrony in the collared flycatcher Ficedula albicollis

Several hypotheses have been proposed to explain the adaptiveness of hatching asynchrony for the parents, but delayed hatching is generally detrimental for the late hatched young. These offspring often experience competitive disadvantage and delayed development. If hatching asynchrony has a reason other than producing competitive differences among offspring, it would be advantageous, not only for the offspring but even for the parents, to compensate for its detrimental effects. In some species, increasing investment into later laid eggs has been reported and discussed as a compensation mechanism, but its effect on nestling growth and fledging size has not been examined in details. In this study we investigated nestling growth and size at fledging in terms of body mass and length of primaries in relation to the accurate laying and hatching order in collared flycatcher Ficedula albicollis broods. We found that females laid larger eggs at the end of the laying sequence, and this helped to decrease the disadvantages for the last offspring. The last offspring had lower body mass growth rate and fledged with shorter feathers, but in both cases the larger the last egg was, the smaller the lag of the offspring was. We conclude, that even if females were not able to fully compensate for the detrimental effects of hatching asynchrony, larger eggs may improve the survival prospects of late hatched nestlings.     

Influence of hatching asynchrony and within-brood parental investment on size, condition, and immunocompetence in nestling red-billed choughs

Studies of hatching asynchrony have rarely assessed the effect of parental investment strategies on the development of intra-brood hierarchies using indicators of nestling quality. The influence of hatching asynchrony and other variables related to parental investment on immunocompetence was evaluated measuring T-cell mediated immune response in broods of red-billed choughs ( Pyrrhocorax pyrrhocorax ), a large long-living passerine with a pronounced sexual size dimorphism. The results showed that T-cell mediated immune response depends on parental effects as shown by the differences between broods on hatching asynchrony. Male nestlings were both heavier and larger than female nestlings, but there was no effect of hatching order on these traits, nor on sex differences in immunocompetence. Differential investment strategy in relation to laying order did not favour older offspring or either of the sexes. Successful reproduction in this species might be so unpredictable and infrequent that strategies of parental investment in the brood could have evolved to attempt to maximize the survival of all nestlings by avoiding within-brood hierachies of size, body condition, and immunocompetence.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 675–684.