Relationships between egg size, chick size at hatching, and chick survival in the Whimbrel Numenius phaeopus

The relationships between egg size, chick size at hatching and chick survival in Whimbrels Numenius phaeopus were studied over a three-year period in the Shetland Isles. Three measurements of chick size at hatching were all positively correlated with egg volume, though the relationship was strongest with hatchling body-weight. In two of the three years the proportion of chicks from a brood which survived to fledging increased significantly with the mean hatching weight of chicks in the brood. Within broods, a significant effect of hatching weight on survival was detectable only up to 7 days after hatching. Between years the egg volumes and hatchling weights of individual female Whimbrels showed relatively little variability, indicating that these attributes could be controlled to a large extent by inheritance.     

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.     

Chick growth and survival in gentoo penguins (Pygoscelis papua): effect of hatching asynchrony and variation in food supply

Summary In the gentoo penguin, Pygoscelis papua, we examined the effects of intra-clutch egg size differences and hatching asynchrony on differential chick growth and survival (including post-fledging survival), in five years for which indices of food supply were available. An initial size hierarchy within-broods at hatching was due to hatching asynchrony not intra-clutch egg size differences. In 1988 only (a poor food year), the weight advantage gained by the first-hatched (A) chick persisted to the end of brooding (30 days), with more second-hatched (B) chicks dying. There was no difference between A- and B-chick weights at fledging (60 days) or in overall chick survival between synchronous and asynchronous broods in any year. Postfledging survival (measured in one year) was not related to fledging weight or hatching order. These results provide only partial support for the hypothesis that gentoo penguins operate a brood reduction strategy to optimise chick survival in years of low food availability. We suggest that hatching asynchrony in gentoo penguins may result from selection to keep the first egg warm as soon as it is laid, due to extreme low ambient temperatures.     

Shiny cowbird Molothrus bonariensis egg size and chick growth vary between two hosts that differ markedly in body size

In birds, egg size affects chick growth and survival and it is an important component of reproductive success. The shiny cowbird Molothrus bonariensis is an extreme generalist brood parasite that uses hosts with a wide range of body masses. Survival of cowbird chicks decreases with host body mass, as competition for food with nestmates is more intense in large than in small hosts. We studied variation in shiny cowbird egg size and chick growth in two hosts that differ markedly in body size: the chalk‐browed mockingbird Mimus saturninus (70–75 g), and the house wren Troglodytes aedon (12–13 g). We analyzed: 1) if females parasitizing mockingbirds lay larger eggs than those parasitizing wrens, and 2) the association between egg size and chick growth. We experimentally controlled for time of parasitism and number of host chicks and evaluated growth rate of male and female parasite chicks. Shiny cowbirds parasitizing mockingbird nests laid larger eggs than those parasitizing wren nests. Chick body mass after hatching was positively associated with egg size until chicks were five days of age, but there was no association between egg size and growth rate, or asymptotic mass. There were no sexual differences in egg size or body mass at the time of hatching, but growth rate was higher in males than in females leading to sexual dimorphism in asymptotic mass. Differences in egg size between hosts and the effect of egg size on body mass after hatching support the hypothesis that different females are specialized in the use of hosts that differ in body mass.     

Testing the relationship between clutch size and brood size in the Coot (Fulica atra)

The time between egg laying and chick fledging is of crucial importance for the survival of young birds. I analyzed breeding output at consecutive phases of growth of young Coots (Fulica atra) relative to the clutch size and laying date. Considering the specific breeding biology of the Coot, I tested whether chick survival reveals clutch size-dependent variability. Clutch size did not affect hatching success; it only affected brood size, and that merely temporarily. During the first 20 days after hatching, i.e. during the time of the highest chick mortality, birds with larger clutches lost chicks at a higher rate. As a result, the number of fledged chicks was independent of the initial number of chicks, and pairs with different clutch sizes had a similar number of fledglings. The laying date had no effect. This pattern of age-related chick survival points to the greater role of the type of chick growth (semi-precocial) and behavior in their survival.     

Factors affecting chick growth in the South Polar Skua (Catharacta maccormicki): food supply, weather and hatching date

This study investigated chick growth in a pelagic Antarctic seabird, the South Polar Skua (Catharacta maccormicki). The factors food supply, weather and hatching date were analysed in a population of 54 breeding pairs at King George Island/South Shetland Islands. Food supply was manipulated by offering fish corresponding to 20% of daily energy demand of chicks to half of the breeding pairs every second day. Growth of mass, head, wing and tarsus was followed and related to the treatment, weather conditions, hatching date and interactive effects. Food supply did not limit chick growth in the studied season. Parents seemed to try to feed their chicks at a maximum rate and succeeded in the studied season because the general food supply was very good. Low temperatures and strong winds depressed chick growth. A growth advantage of food-supplemented chicks could be observed when the natural conditions for chick growth were sub-optimal. Chick growth rate was strongly negatively associated with hatching date and worsening weather during the reproductive season could be excluded as explanatory variable for this finding.     

Limited effects of egg size on chick growth in the chinstrap Penguin Pygoscelis antarctica

We studied the effect of egg size on chick growth in the chinstrap penguin under natural non-manipulative, conditions. The influence of egg size on chick mass at hatching, 15 and 45 days of age was analysed controlling for the effect of hatching date, as hatching date has been repeatedly associated with chick growth in this species. Egg volume explained 20% of the variation in the body mass of chicks at hatching. Its effect disappeared as chicks grew, being statistically not significant at the age of 15 and 45 days. Egg volume asymmetry also had no role in determining chicks' growth asymmetries. Accepted: 6 June 1998     

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.     

Adaptive hatching hypotheses do not explain asynchronous hatching in Brown-headed Parrots Poicephalus cryptoxanthus

At the core of the suite of adaptive hatching hypotheses advanced to explain asynchronous hatching in birds is the assumption that if food is not limited then all the hatchlings will develop normally to adulthood. In this study Brown-headed Parrot Poicephalus cryptoxanthus chicks were hand fed and weighed on a daily basis. Their respective masses were compared using a reparamatised Richards growth model. The results show that although the second and third hatched chicks initially grow faster than the first, the final mass attained by the third chick is significantly less than either of its siblings. Adaptive hatching hypotheses do not explain this result. We propose that the third chick is opportunistic and that is has evolved as insurance against prior chicks surviving because of interspecific pressure on the breeding biology of the species.     

Immunocytochemical studies of chicken somatotrophs and somatotroph granules before and after hatching

Immunocytochemical methods were used to gain information about the embryonic development of chicken somatotrophs before and after hatching. To localize growth hormone, anterior pituitary sections were incubated with growth-hormone antibody, and then an indirect peroxidase method was used for light microscopy and an immunogold method for electron microscopy. The earliest evidence of embryonic somatotrophs was seen at 12 days. At this stage somatotrophs were sparse (0.2% of parenchymal cells) and their granules were pleomorphic with elongated ovoid and lozenge shapes predominating. Few of the immunogold-labeled somatotroph granules of the embryo were spherical until 15 days after fertilization. At 18 days, most of the granules were spherical (their shape in the adult chicken). During the six days between the 15-day-old embryo and the 1-day-old chick, the number of gold particles per granule section approximately doubled suggesting an increase in growth hormone content of the granules. This rise was the result of increases in the size of the granule sections and in the concentration of gold particles in the sections. During the embryonic period of 12–20 days, somatotrophs were not more than 3.6% of the anterior pituitary cell population. During the following two days, between the 20-day-old embryo and the 1-day-old chick, the percentage of somatotrophs in the pituitary parenchymal cell population rose rapidly from 3.6% to 20.7% and then increased slowly to 24.6% during the period of 1–5 days after hatching. Both the sharp percentage rise in somatotrophs (20-day-old embryo to 1-day-old chick) and the rise in growth hormone content of the granules (15-day-old embryo to 1-day-old chick) suggested by gold-particle counts occur close to the time of hatching. These morphological changes may reflect an increased synthesis of growth hormone that is responsible for the rise in plasma growth-hormone concentration that begins about the same time and is especially abrupt two days later (1–3 days after hatching).     

Growth of nestling black kites Milvus migrans: effects of hatching order, weather and season

The growth of black kites Milvus migrans was studied in 1981, 1982 and 1984 at the Doñana Biological Reserve (south-west Spain). Fifteen variables were submitted to a principal component analysis in order to characterize the growth pattern of the young. In general, in the nestlings that fledged, rapid weight gain corresponded to rapid growth of the tarsus and feathers. For nestlings that starved, tarsus growth took priority over weight gain. Within the same brood, the last-hatched sibling grew more slowly than its older sibling and the differences between them increased in proportion to the hatching asynchrony. Brood size did not affect growth as clearly as hatching order and hatching asynchrony. Increased rainfall and lower maximum temperature, factors that seem to make the search for food more difficult, had negative effects on the youngest sibling in a multiple brood and on single nestlings. The only environmental variable that had an effect on all of the nestlings was the hatching date.     

The effect of hatching date on parental care, chick growth, and chick mortality in the chinstrap penguin Pygoscelis antarctica

We studied the effect of hatching date on breeding performance (chick growth and mortality) and phenology (creching and fledging ages) of the chinstrap penguin during three years. The year affected every variable considered, probably due to pack-ice persistence and food availability differences between years. Hatching date had slight or no effect on mortality and early growth, but was negatively correlated with creching age, which, in turn, was positively related to final size. The decision to leave the chicks unguarded does not seem to be based on the condition of the chicks, but on that of adults. Fledging age was negatively correlated with hatching date, and this effect was more marked in the year with poor growth performance. Given the short time available for breeding in Antarctica, there must be conflicting pressures between investing in feeding chicks and advancing the period of premoult resource storage, this explaining the strong relationship between hatching dates and subsequent phenological events (creching and fledging). In this kind of study, it may be important to remove the effect of inter-year variation before assessing the possible effects of other variables.     

Relationship between clutch size,egg volume and hatching success in a Yellow-legged Gull Larus michahellis colony in south-eastern Tunisia

This study aimed to test the hypothesis that clutch size covaries with egg volume and hatching success in the Yellow-legged Gull Larus michahellis. We determined clutch size and egg volume in a sample of 131 nests, and we used the data to check whether egg volume varied among nests according to clutch size, while taking into account the effects of egg laying order. We also estimated hatching success rate and investigated the relationship between hatching success and clutch size. Egg volume varied among clutches according to clutch size, with eggs being larger in three-egg clutches than in two-egg clutches. Moreover, three-egg clutches showed higher daily survival rates, and hence hatching success, than two-egg clutches. Overall, our results suggest that in the Yellow-legged Gull clutch size covaries with egg volume and hatching success, which could possibly reflect an age effect through different mechanisms. Indeed, older females could be hypothesised to exhibit greater breeding performance than younger females because of their higher experience in tapping energy resources for egg formation and defending nests from dangers. Moreover, due to their age, older females are likely to have lower residual reproductive potential and should invest more heavily in current breeding attempts.     

Altricial and precocial developmental features in modern meat-type chicks

Variability of egg weight, egg yolk content, neonatal growth rate and relationships of these parameters were studied in meat-type chicks. As it had been established the level of variability in neonatal growth traits was greater than variability of the egg morphology parameters. Egg weight had stronger influence on the chicks' neonatal growth rate than egg yolk content did. Low egg size was associated with limited neonatal growth rate variability, declined chick weight at hatching and increased relative growth rate throughout four days post hatch. Comparison of egg morphological parameters in two species having the same female definitive body weight--meat-type domestic fowl (precocial type) and brown pelican (altricial type) has shown, that, in contrary to predicted on the basis of avian developmental typology, egg weight to female body ratio was greater in brown pelican, egg yolk content was equal in both species.     

Reproductive measures and chick provisioning of Cory's Shearwater Calonectris diomedea borealis in the Azores

Capsule Timing of breeding influenced wing-length at fledging, and egg size may be an indicator of fledging weight and the amount of food received by chicks.

Aims To investigate chick growth, temporal patterns of chick food provisioning and the importance of indices of parental condition or quality, egg size and hatching date, to predict nestling body mass and wing-length at fledging, and compare breeding and chick feeding characteristics between colonies in the northeast Atlantic.

Methods A survey of Cory's Shearwater nests was carried out at Vila islet. A sample of 52 chicks, ringed and weighed at hatching, was selected to study chick growth and food provisioning.

Results Hatching success (51%) was much lower than fledging success (87%). Both hatching date and egg size contributed to explain wing-length at fledging, but hatching date, which was negatively correlated with wing-length at fledging, had the most important contribution (22%). There was some indication that egg size may explain variation in fledging weight and the amount of food received by chicks. Food delivery and feeding frequency of chicks varied throughout the chick development stage and three phases were distinguished: (1) 0–29 days, the highest feeding frequency values and a linear increase in food delivery; (2) 30–69 days, an oscillation in food delivery and medium feeding frequencies; (3) 70–90+ days, a sharp decrease in both food delivery and feeding frequency.

Conclusion Variation in food availability did not seem sufficient to override the overall importance of indices of parental quality in determining reproductive measures and chick provisioning. Breeding and feeding characteristics were similar between colonies in the northeast Atlantic, with variability in chick provisioning higher further south.     

Offspring size and timing of hatching determine survival and reproductive output in a lizard

Selection on offspring size and timing of birth or hatching could have important consequences for maternal investment strategies. Here we show consistent viability selection on hatchling body length across 2 consecutive years in a lizard that lays several clutches per season. There was no effect of hatching date on survival to maturity. However, both early hatching and large hatchling size increased adult size, which has a positive effect on total reproductive output. Earlier hatching also led to an earlier onset of reproduction. Overall, increased survival probability for large hatchlings and a positive effect of clutch size on recruitment suggest consistent directional selection on both egg size and clutch size within and across years. Because offspring size and timing of hatching are strongly affected by environmental and maternal effects, there should be potential for strong transgenerational effects on reproductive output in this species. We briefly discuss the implications of these results for the evolutionary ecology of maternal investment and population fluctuations in short-lived lizards.     

Development of intraspecific size variation in black coucals,white-browed coucals and ruffs from hatching to fledging

Most studies on sexual size dimorphism address proximate and functional questions related to adults, but sexual size dimorphism usually develops during ontogeny and developmental trajectories of sexual size dimorphism are poorly understood. We studied three bird species with variation in adult sexual size dimorphism: black coucals (females 69% heavier than males), white-browed coucals (females 13% heavier than males) and ruffs (males 70% heavier than females). Using a flexible Bayesian generalized additive model framework (GAMM), we examined when and how sexual size dimorphism developed in body mass, tarsus length and bill length from hatching until fledging. In ruffs, we additionally examined the development of intrasexual size variation among three morphs (Independents, Satellites and Faeders), which creates another level of variation in adult size of males and females. We found that 27–100% of the adult inter- and intrasexual size variation developed until fledging although none of the species completed growth during the observational period. In general, the larger sex/morph grew more quickly and reached its maximal absolute growth rate later than the smaller sex/morph. However, when the daily increase in body mass was modelled as a proportion, growth patterns were synchronized between and within sexes. Growth broadly followed sigmoidal asymptotic models, however only with the flexible GAMM approach, residual distributions were homogeneous over the entire observation periods. These results provide a platform for future studies to relate variation in growth to selective pressures and proximate mechanisms in these three species, and they highlight the advantage of using a flexible model approach for examining growth variation during ontogeny.     

三种鹭异步孵化与雏鸟生长的比较研究

根据1996～1999年的野外工作和实验饲养,研究了池鹭、白鹭、夜鹭3种鹭的繁殖、雏鸟生长和异步孵化对雏鸟生长的影响.3种鹭于4月上、中旬迁到常山、余杭鹭保护区,9月下旬陆续迁离.孵卵期池鹭为23.0 d(n=26),白鹭23.9 d(n=32),夜鹭25.3 d(n=34);孵化率池鹭为76.32%,自鹭86.96%,夜鹭95.45%,池鹭繁殖力为3.21只,白鹭3.38只,夜鹭3.50只.雏鸟体重增长与成体体形大小呈负相关.幼雏体重增长与孵化顺序相关.在生长早期(≤8 d)全部雏鸟正常发育,之后差异显著.第1和第2孵化雏鸟生长曲线相似,但明显高于第4雏鸟,表明较早孵化者获得食物能力较强,而迟孵出雏鸟有食物不足现象.该3种鹭孵育幼雏能力的最适度估计为3只左右.     

Activation of the serotonergic system in chick spinal cord during hatching.

The objectives of the present study were to determine the levels of serotonin (5-HT), its major catabolic metabolite, 5-hydroxyindoleacetic acid (5-HIAA), and norepinephrine (NE) in chick spinal cord before, during, and after hatching and also to determine if changes in the levels of these chemicals are directly related to the hatching behavior. The levels of 5-HT, 5-HIAA, and NE were measured by high performance liquid chromatography with electrochemical detection in whole spinal cords of 20-day-old "pre-hatching" embryos, 21-day-old "normal hatching" embryos, 0-day-old "post-hatching" chicks, and 0-day-old "glass egg hatching" chicks. NE was measured but no significant differences were found in NE levels among experimental groups. The concentration of 5-HT was elevated in chick embryo spinal cords during normal hatching compared to pre-hatching embryos and post-hatching chicks. The concentration of 5-HIAA was elevated during and after normal hatching compared to pre-hatching embryos. However, neither 5-HT nor 5-HIAA levels were found to be elevated in chick spinal cords during glass egg hatching compared to pre-hatching embryos or post-hatching chicks. Therefore, there appears to be an activation of the serotonergic system in chick spinal cord related to the specific event of hatching but this activation is not directly related to the movements common to both hatching and glass egg hatching.     

Egg size, offspring sex and hatching asynchrony in zebra finches Taeniopygia guttata

Experimental synchronization of onset of incubation was employed in laboratory held zebra finches Taeniopygia guttata to study whether differential resource allocation and possible bias of offspring sex in subsequent eggs in the laying order could mitigate the effects of hatching asynchrony. We found that egg mass increased with laying order, but offspring sex was not related to laying order. Among synchronized clutches, eggs hatched more synchronously than eggs from control nests. Survival probability was related to egg mass, and as expected, this effect differed between experimental groups: it was positive among synchronized broods and not significantly related among asynchronous broods. This suggests that increase in egg mass with the laying order might reduce disparities between early and late hatching chicks. Female nestlings survived better than male nestlings. However their growth was impaired in synchronized broods, whilst growth of males was not affected by hatching synchronization.