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.     

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.     

Parental quality and egg size affect chick survival in a precocial bird, the lapwing Vanellus vanellus

Effects of egg size and parental quality on lapwing Vanellus vanellus chick survival were studied in southwestern Sweden over 6 years. Chicks from large eggs were heavier at hatching and survived significantly better than those from small eggs. To control for the confounding effect of parental quality on egg size and chick survival, we performed a cross-fostering experiment during 2 years, exchanging clutches between nests with large and small eggs. In control clutches, chicks from large eggs survived better than those from small eggs, but we found no significant difference in chick survival between exchanged clutches. Thus, egg size did not affect chick survival independently of parental quality. Fledging success increased with parental age and/or experience, and with female body mass. Hence, both egg size and parental quality affect chick survival in the lapwing. Received: 22 February 1996 / Accepted: 30 September 1996     

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.     

Brood reduction in the Blue-eyed Shag Phalacrocorax atriceps

Brood reduction is common in a population of Blue-eyed Shags on Signy Island, South Orkney Islands. This paper describes possible adaptations which may reduce the brood. In clutches of three, the last egg was smaller, and hatched 2.4 days later than its siblings. Whilst 78–84% of first and second ('A' & 'B') chicks fledged, only 11 % of 'C' chicks did. In a sample of artificially synchronized broods chick survival was as high as in normal asynchronously hatching broods, but there were more cases of total brood loss. The age at which the C chick died was related inversely to the length of the A-C hatching interval. Relative differences in sibling weights were highest during the first 12 days, when most of the C chick deaths occurred. At this age the daily food requirements of each brood of three was one-tenth that of each brood of two just prior to fledging. It is suggested that C chicks were unable to compete effectively for a food supply which was limited by the parents, rather than by the environment. The asymptotic weight attained by A chicks was inversely related to brood size, and was greater than that of B or C chicks. Normal asynchronous broods produced at least one heavy (A) chick and one medium weight (B) chick, whilst in synchronized broods the asymptotic weight attained was similar to that of B chicks in normal broods.     

Variations in size and growth of Great skua Catharacta skua chicks in relation to adult age, hatching date, egg volume, brood size and hatching sequence

The paper describes the relationships between adult age, hatching date, egg volume, brood size, hatching sequence, chick weight at hatching, chick growth rate and chick body size for a population of Great skuas in Shetland. The relative importance of these factors, and food availability, is assessed in determining chick size, weight, and growth rate.     

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.     

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     

Relationships between brood size and parental provisioning performance in chinstrap penguins during the chick guard phase

 Chinstrap penguins (Pygoscelis antarctica) normally lay two eggs, but brood size is often reduced by mortality during incubation or after hatching. We hypothesized that this variation in brood size would affect the parents’ foraging behavior and their chick provisioning performance. We studied patterns of adult foraging trip duration and frequency, food load delivery, and chick growth rates in relation to brood size during the guard phase in four breeding seasons (1991–1994) on Seal Island, Antarctica. Within a given year, parents with two chicks made more frequent foraging trips to sea and may have transported larger food loads to the nest; however, the duration of foraging trips was unrelated to brood size. Overall, parents with two chicks spent ∼15% more time at sea than parents with only one chick. Both the frequency and duration of foraging trips varied between years. Foraging trip duration may partly reflect the birds’ foraging radius, which probably varies with time in response to shifts in krill distribution. Chick growth rate varied betwen years, but was related to brood size only in 1992, when chicks from two-chick broods grew significantly more slowly than chicks from one-chick broods. Food loads transported to chicks, as well as chick growth rates, were highest in 1994, when concurrent hydroacoustic studies indicated that regional krill biomass was severely depressed. This apparent anomaly suggests that the spatial scale of the krill survey may have been too coarse to detect some high-density krill aggregations within the penguins’ foraging range. Received: 26 September 1995 / Accepted: 12 May 1996     

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.     

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.     

Penguin Chicks Benefit from Elevated Yolk Androgen Levels under Sibling Competition

Crested penguins (genus Eudyptes) have a peculiar hatching pattern, with the first-laid egg (A-egg) hatching after the second-laid egg (B-egg) and chicks from A-eggs typically having a much lower survival probability. Maternal yolk androgens have been suggested to contribute to the competitive superiority of the B-chick in southern rockhopper penguins Eudyptes chrysocome, given their important role in mediating sibling competition in other species. We therefore increased the yolk androgen levels in freshly-laid eggs and examined the consequences for sibling competition - via effects on embryonic developmental times, chick growth and early survival. We placed one androgen-treated egg and one control egg into each foster nest, matching them for mass, laying date and laying order. The androgen treatment did not significantly affect embryonic developmental times or chick measurements at hatching. However, elevated yolk androgen levels benefitted chick growth in interaction with the number of siblings in a brood. Chicks from androgen-treated eggs had faster growth in the presence of a sibling than chicks from control eggs. Under these circumstances they also had a higher survival probability. Thus maternal androgens appear to reinforce the observed hatching pattern, facilitating brood reduction. This contrasts to most previous studies in other species where yolk androgens have been shown to compensate for the negative consequences of delayed hatching within the brood hierarchy.     

A stab in the dark: chick killing by brood parasitic honeyguides

The most virulent avian brood parasites obligately kill host young soon after hatching, thus ensuring their monopoly of host parental care. While the host eviction behaviour of cuckoos (Cuculidae) is well documented, the host killing behaviour of honeyguide (Indicatoridae) chicks has been witnessed only once, 60 years ago, and never in situ in host nests. Here, we report from the Afrotropical greater honeyguide the first detailed observations of honeyguides killing host chicks with their specially adapted bill hooks, based on repeated video recordings (available in the electronic supplementary material). Adult greater honeyguides puncture host eggs when they lay their own, but in about half of host nests at least one host egg survived, precipitating chick killing by the honeyguide hatchling. Hosts always hatched after honeyguide chicks, and were killed within hours. Despite being blind and in total darkness, honeyguides attacked host young with sustained biting, grasping and shaking motions. Attack time of 1-5 min was sufficient to cause host death, which took from 9 min to over 7 h from first attack. Honeyguides also bit unhatched eggs and human hands, but only rarely bit the host parents feeding them.     

CLUTCH SIZE, EGG SIZE, HATCH WEIGHT AND LAYING DATE IN RELATION TO EARLY MORTALITY IN RED GROUSE LAGOPUS LAGOPUS SCOTICUS CHICKS

Clutches of Red Grouse eggs were collected from the wild and subsequent hatching and rearing done in standard conditions in captivity. Variations in chick survival from one clutch to another in the same year were related to differences in hatch weight. Hatch weight was determined only partly by egg size. Weight loss between laying and hatching was related to survival independently of egg size. Variation in this weight loss obscured any simple relationship between egg size and survival, except in eggs laid by captive hens. Intrinsic differences amongst hens caused some variations in laying date, egg size, hatch weight and chick survival. Variations in egg size and hatch weight accounted for less than half the variation in survival; other unmeasured intrinsic factors were also important. Big clutches hatched earlier than small ones. The commonest clutches were of seven and eight eggs, with six and nine frequent. Very big clutches of ten or more eggs were infrequent and chicks from them sometimes survived worse than from smaller clutches. As in other species, the commonest clutch sizes were not the most productive. There was no simple relationship between egg size and clutch size.     

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.     

Seasonal variation in reproductive measures of tropical Roseate Terns Sterna dougallih previously undescribed breeding patterns in a seabird

Seasonal variation in egg-laying, egg size, hatching success, hatchling mass, fledging success and chick growth of Roseate Terms Sterna dougallii breeding on Aride Island (Seychelles), Indian Ocean, were studied in 1997 and 1998. I investigated to what extent two patterns, common in a range of species, were followed by tropical Roseate Terns: (a) seasonal decrease in clutch size, egg size and breeding success and (b) an increase in breeding success with increasing egg weight. In 1997 (a poor year), the earliest nesting birds laid significantly smaller eggs, and chicks were lighter at hatching than those of peak nesting birds. The mean clutch size, of 1.04 eggs, showed no seasonal variation and no 'b'-eggs hatched. In 1998 (a good year) the earliest nesting birds laid eggs of similar size and their chicks were of similar weight to those of peak nesting birds. Mean clutch size, of 1.25 eggs, increased significantly through the season and about 60% of the 'b'-eggs hatched. In 1997, hatching success was 57% whereas in 1998 it was 80%. In both years, breeding success declined significantly through the season. The fact that the earliest breeding birds laid smaller eggs in a poor year and smaller clutches in a good year is in marked contrast to a range of other species, and to temperate-nesting Roseate Terns. Egg volume explained about half of the variance in hatchling mass in both years, but only 15% of the variation in linear growth rate. Hatching date was the only variable with a significant effect on fledging success. Roseate Terns on Aride seemed to sacrifice egg size and clutch size for earliness of laying. Presumably it is a strategy of older birds to lay as early as possible and may be regarded as a response of tropical Roseate Terns to breeding under relatively poor, and seasonally declining, food conditions.     

Breeding success in a Houbara Bustard Chlamydotis [undulata] macqueenii population on the eastern fringe of the Jungar Basin, People's Republic of China

Nesting success and chick survival of a migratory population of Houbara Bustard Chlamydotis [undulata] macqueenii were studied during three consecutive years (1998–2000) in the Xinjiang province of north‐west China. A total of 45 nests was monitored and 85 broods comprising 227 chicks were captured, of which 82 chicks were radio‐tracked. Start of laying varied between 6 and 17 April between years but the laying mode fell consistently between 26 and 30 April. Mean clutch size was 4.0 (sd = 0.8) (range 2–6) for early clutches and 3.3 (sd = 1.1) for late clutches (range 2–5). The average nesting success was 0.588 (sd = 0.270) but great variations were observed between years –0.882 in 1998, 0.530 in 1999 and 0.351 in 2000. This was related to increased predation in 1999 and 2000, which is reflected by increased predator density (chiefly Corsac Fox Vulpes corsac and Long‐legged Buzzards Buteo rufinus). The overall hatchability, defined as the proportion of eggs hatched in successful nests was 0.839 sd = 0.238). The average brood size at hatching varied from 2.9 (sd = 0.8) to 3.3 (sd = 0.9) according to years, and no significant decrease in brood size was observed in the first 5 days post‐hatching. In 1999 and 2000 the brood size diminished sharply (14% and 27%, respectively) in prefledging chicks. A further severe decrease (37%) was observed in fledglings in 2000, probably due to predation by raptors. For the 3 years of the study, a successful female Houbara would bring on average 2.3 (sd = 0.9) chicks to fledging and would have lost 30.2% (sd = 14.9%) of its brood to adversity during the rearing process. The proportion of females that lost their entire brood was 0.181 in 1998, 0.708 in 1999 and 0.453 in 2000. For the 3 years of the study, only 55.3% (sd = 26.3%) of the females hatching eggs brought chicks to fledging. The overall chick production was 0.827 per breeding female per year and the probability of an egg laid producing a fledgling of 8 weeks old was 0.190.     

Reproductive success of the Kittiwake Rissa tridactyla: the roles of clutch size, chick growth rates and parental quality

Kittiwake growth rates and breeding success are examined in relation to survival between fledging and breeding and to adult survival rates. High chick growth rates lead to increased survival after fledging. Broods of three (the maximum brood size) did not suffer lower fledging success than broods of two and clutches of three fledged appreciably more chicks per pair than did clutches of two or one. On average, the a- and b -chicks in broods of three grew at a faster rate and had a higher survival before breeding than those from smaller broods. Chicks from broods of two with experienced female parents grew at a faster rate than those of inexperienced female parents. Female parents which laid three egg clutches had a higher survival rate than those which laid clutches of two or one. We contend that three egg clutches were laid by higher quality individuals. We believe that clutch size indicates the condition of the Kittiwakes forming the pair. This condition probably has a genetical component, but is modified by environmental factors.     

Effects of egg size, parental quality and hatch-date on growth and survival of Common Tern Sterna hirundo chicks

We examined the relative contributions of egg size, parental quality and hatch-date to growth and survival of second-hatched chicks (those chicks making the greatest contribution to differences in productivity among pairs) by exchanging clutches among nests of Common Terns Sterna hirundo matched for lay-date (range 13 May to 9 June). The mass of a second-laid egg in an exchanged clutch ranged from 17.70 to 23.80 g. Growth and survival were studied during three periods: early (days 0–3), middle (days 3–12) and late (days 12–25). Both egg mass and hatch-date were important predictors of hatchling mass (positive relationships), although there was no seasonal trend in egg mass. During the middle period, hatch-date was a significant predictor of mass gain and survival (inverse relationships). After controlling for hatch-date, other indices of parental quality made only small contributions to chick mass gain and survival. Our results suggest that although breeding early generally leads to greater overall survival of chicks, several important interactions among egg 'quality', parental quality and early laying may affect breeding success under specific conditions.     

Sex of the first hatched chick influences survival of the brood in the herring gull (Larus argentatus)

Differences in the growth rate of male and female offspring can result in different parental rearing costs for sons and daughters. Such differences may also influence the survival chances of male and female offspring when conditions are unfavourable. In birds, hatching asynchrony leads to hierarchical competition for food between siblings. Therefore, the sex of the chick in the first hatched position in the brood may influence breeding success by affecting the extent to which the later hatched chicks can compete for resources. The interaction between brood sex composition and chick performance in the herring gull Larus argentatus was examined under different environmental conditions. When environmental conditions were relatively good, chick survival within broods was better when a female was first to hatch, an effect that was most obvious later in the season. When conditions were poorer however, sex of the first hatched chicks was not related to brood survival. In neither situation did the overall primary sex ratio differ from equality. However in the year of relatively good food availability, the first chick in the brood was more likely to be male early in the season, which was when the disadvantageous effects on brood survival of males being in this position are weakest.