Experimental evidence for the relationship between food supply, parental effort and chick survival in the Lesser Black-backed Gull Larus fuscus

We studied breeding success, chick growth, parental effort and chick behaviour in two groups of Lesser Black-backed Gulls Larus fuscus whose chicks were provided with additional food until 7 days after hatching or until fledging. These data were compared with those from control pairs which we studied simultaneously to test the hypotheses that food was in short supply during the chick stage at the colony site and that in such circumstances the behaviour of adults and young is mainly responsible for the low success. Pairs whose chicks were fed with additional food until fledging showed a higher fledging success than control pairs (intermediate for pairs of first experimental group). During the first week after hatching, experimental adults of both groups were present together at the territory for longer than control pairs. In adult females of experimental pairs, the length of feeding trips was shorter than in females of control pairs, whilst the rate of chick feeding was more frequent in the experimental broods. After the chicks were 7 days old, differences were significant only for the experimental pairs whose chicks were provided with additional food until fledging. Chicks fed until fledging showed a higher daily mass and wing-length increments and reached a higher fledging mass at an earlier age than both control chicks and chicks which were provided with additional food until day 7. Starvation occurred only in control chicks and in chicks of the first experimental group after we had stopped providing food. When food was in short supply, fledging success of gulls was adversely affected as a result of both starvation (because of the lower feeding rates of chicks) and a higher predation rate (arising from changes in behaviour of both adults and chicks).     

Fly or die: the role of fat stores in the growth and development of Grey-headed Albatross Diomedea chrysostoma chicks

Chicks of albatrosses, like other Procellariiformes, become independent at a mass similar to their parents but during growth attain a peak mass some 30% or more greater, before losing mass prior to fledging. The current views are that this high peak mass represents chicks storing fat reserves as an energy sink, or as an insurance against periodic food scarcity, or as a Consequence of natural stochastic variation in provisioning rate. We analysed growth and body composition of Grey‐headed Albatross Diomedea chrysostoma chicks at Bird Island, South Georgia in 1984 and 1986, two years of very different food availability. In 1984 when overall breeding success was only 28% (the lowest in 20 years and less than halt that in 1986), chicks were significantly smaller in terms of peak mass (by 37%), primary length (by 25%), liver, lung, heart and kidney size (by 18–34%) and fat (by 75–80%) but not significantly different in terms of skeletal (tarsus, culmen, ulna, sternum) or muscle (pectoral, leg) size. Despite these differences, there were some important similarities in the patterns of growth in both years. Up to the attainment of peak mass, most of the growth of organs and of skeletal structures was completed and little fat was deposited. In the remaining part of the chick‐rearing period, feather growth and acquisition of fat stores were undertaken. Thus Grey‐headed Albatross chicks begin to acquire substantial fat stores only during the later part of the development period; this is contrary to the predictions of any of the existing hypotheses concerning provisioning patterns and the role of fat stores in Procellariiformes. We propose that the deposition of fat in the later stages of chick growth is an adaptation to: (a) ensure against energy demands and/or nutritional stress affecting the quality of flight feathers (many of which are not renewed for up to three years after fledging); and (b) provide an energy reserve for chicks to use in the critical period immediately after independence.     

Chick provisioning rates and growth in Blacklbrowed Albatross Diomedea melanophris and Grey-headed Albatross D. chrysostoma at Bird Island, South Georgia

We compared the parental division of labour and the pattern and rate of parental provisioning by two sympatric species of albatross of similar mass and breeding timetable but differing in diet and in the duration of chick‐rearing. Using electronic weighing platforms inside artificial nests, we recorded chick mass of Black‐browed Albatross and Grey‐headed Albatross at Bird Island, South Georgia every 10 minutes for both species in 1993 and 1994 and for each species in two other years between 1990 and 1996. The chick mass data (nearly one million weighings) were used to calculate meal mass (over 5000 meals) and intervals between meals. Adult birds were fitted with radio‐transmitters which allowed each meal to be allocated to the appropriate parent. The combination of meal mass and foraging trip duration were used to calculate provisioning rates for chicks and individual adults. Overall, Black‐browed Albatrosses delivered significantly lighter meals (569 g) than Grey‐headed Albatrosses (616 g) but more frequently (every 2.07 days and 2.50 days respectively). Thus combining foraging trip data for both parents, Black‐browed Albatross chicks received a meal every 1.22 days compared with 1.26 days for Greyheaded Albatross. These rates did not differ significantly. The contribution of each sex of each species in chick provisioning fluctuated between years, being similar in some years or biased towards males in others. Chicks of both species that failed to fledge received smaller, less frequent meals than successful chicks. In 1990 and 1994, Black‐browed Albatross chick provisioning rates were lower than in 1992 and 1993. In 1990, both meal mass and trip duration were affected, but only in 1994 was trip duration longer. Greyheaded Albatross chick provisioning rate was lower in 1994 than in other years but trip duration was longer. In each species, significant changes in meal mass and trip duration occurred within the chick‐rearing period. Chick provisioning rates invariably declined before chicks attained their peak mass. For both species, chick growth rates and peak and fledging mass, but not fledging age, were affected by differences in provisioning rate.     

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.     

Variation in growth in Sandwich Tern chicks Sterna sandvicensis and the consequences for pre- and post-fledging mortality

Fitness consequences of variation in body mass growth and body condition were studied in a Sandwich Tern Sterna sandvicensis colony on Griend, Dutch Wadden Sea, during 1990–2000. Body mass increment during the linear growth phase predicted nestling survival probabilities accurately. Chicks growing less than 8 g per day had low survival probabilities until fledging, but within a range of 8–11 g per day growth only small effects on chick survival were observed. Effects of slow growth on survival became obvious after about 10 days after hatching. Slow growing chicks reached a much lower fledging mass, whereas slow growth had only small effects on structural size at fledging. Body condition of the chicks was highly variable and had strong effects on survival until fledging. However, body condition during the nestling stage did not influence post-fledging survival. Body condition at fledging had no effects on post-fledging survival and did not affect final mass or body size. It is argued that low fledging mass can be overcome soon after fledging, as parents take their fledglings closer to the foraging areas, thereby avoiding high rates of kleptoparasitism by Black-headed Gulls Larus ridibundus .     

Nestling feeding strategy of the British storm-petrel Hydrobates pelagicus in a Mediterranean colony

The pattern of chick feeding of the British storm-petrel Hydrobates pelagicus in a Mediterranean colony was examined by weighing chicks at 24-h intervals on different days during the nestling period. In order to calibrate daily mass increments (NET) against number of feedings, daily mass changes of chicks were regressed upon the sum of positive mass increments recorded overnight (SUM) during four nights. The average meal size delivered to chicks per night by one parent was 3.5 ß 1.3 g or 12% of adult weight. This was insufficient for sustaining constant chick mass during a day. On average 85% of chicks were fed each night, and the mean interval between feeds was 1.2 ß 0.5 nights. Nightly feeding frequencies differed among days, but this night-to-night variation was not related to meteorological conditions. Both food requirements necessary for chick body maintenance (zero-growth) and meal size were relatively constant for ages up to 59 days. However, feeding frequency decreased throughout the fledging period, accounting for agespecific variation in growth rates until fledging. Food requirement and feeding patterns at Benidorm were different from North Atlantic colonies. None the less, growth patterns were almost identical, suggesting adjustment to maintain chick body mass at a determined level, as food delivery to nestling appears to be regulated to chicks'nutritional requirements.     

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.     

Rates and consequences of relaying in little auks Alle alle breeding in the High Arctic an experimental study with egg removal

Most seabirds have a small clutch size. Thus, replacement of a clutch after loss can make important contributions to an individual’s lifetime reproductive success. However, in the condition of short polar summer, relaying propensity may be time‐constrained. In this study, we investigated rates and consequences of relaying in a small High Arctic seabird, the little auk Alle alle. We performed an experiment in which we removed the single egg from 20 nests of early‐laying breeders. We measured relaying rates, and compared chick body mass and breeding success between the experimental and control nests. Despite the narrow window of the Arctic summer and the closely synchronized breeding, 75% of females produced a replacement egg just 2.7% smaller in volume than the first egg. This indicates that in little auks, the demographic effects of disruptions to breeding attempts (by predators, adverse weather or human activity) may be mitigated to some extent by replacement clutches. However, peak body mass and fledging body mass were lower in the experimental than the control chicks. This effect was rather a consequence of late hatching – chicks from replacement clutches followed seasonal decline in peak body mass and fledging mass. Finally, breeding success and chick survival up to 20 d in the experimental nests were respectively 34 and 37% lower than in the control nests. Thus, the quality and post‐fledging survival of chicks from the replacement clutches were probably lower compared to the chicks hatched from the first‐laid eggs.     

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.     

Growth and its relationship to fledging success of African black oystercatcher Haematopus moquini chicks

We investigated the growth of African black oystercatcher Haematopus moquini chicks on Robben Island, South Africa, over three austral summers, 2001-2004. Using a robust regression analysis to determine the growth parameters of chicks of known and unknown age we found that oystercatchers from our study population had a Gompertz growth rate coefficient that was 2% less than predicted for body mass based on the equation for waders. Leg growth lagged initially, then increased and slowed again as the chicks became older, whereas wing growth was slow initially but increased with age. Chicks with small growth rate coefficients for body mass exhibited retarded growth of all body measures except wing length. This enabled these chicks to fledge in a shorter period of time than their slow growth would otherwise allow. The growth rate of body mass was observed to vary greatly between chicks. Fast-growing African black oystercatchers had a shorter pre-fledging period; were larger at fledging and were more likely to fledge successfully. African black oystercatchers display sibling rivalry, and once a dominance relationship is established, the larger chick remains so during the pre-fledging period. Larger siblings fledged earlier and at a heavier mass than the smaller siblings and this may improve their chances of survival. Neither hatching date nor brood size influenced the growth rate coefficients.     

Provisioning strategies and growth patterns of Light-mantled Sooty Albatrosses Phoebetria palpebrata on Macquarie Island

Provisioning regimes and growth of Light-mantled Sooty Albatrosses (LMSA) (Phoebetria palpebrata) were investigated on subantarctic Macquarie Island using an automatic tracking system and automatic weighing nests. The nests were deployed under five chicks in the post-brood provisioning period in 2000 and 2001. Adults typically utilised a cyclical foraging strategy consisting of a long foraging trip followed by three to four shorter trips. Chicks received an average (±SE) of 37.5±2.3 kg of food in the post-brood provisioning period and were fed every 1.6±0.1 days with a mean meal size of 520±10 g. Chicks grew at a rate of 61.3±1.0 g day^-1 to a peak mass of 4.4±0.1 kg. Mean chick fledging mass was 3.0±0.1 kg. LMSA on Macquarie Island fed their chicks more frequently and showed a lower mean trip duration than conspecifics at South Georgia, which is likely related to proximity of productive Antarctic shelf waters, differences in prey availability and competition with other Procellariiformes.     

Growth, fledging success and post-fledging survival of juvenile Oystercatchers Haematopus ostralegus

We studied the consequences of differences in growth rate on the subsequent survival of Oystercatcher Haematopus ostralegus chicks. Fledging success increased sharply with growth rate, from zero in chicks growing at less than 6 g per day to about 85% in chicks growing at more than 10 g per day. The age at which chicks fledged varied from 27 to 52 days. Chicks which fledged at an early age displayed a much faster growth rate than later fledging chicks. Although slow growth resulted in a considerable prolongation of the period before fledging, slow-growing chicks fledged at a smaller size and with a lower body-weight than fast-growing chicks. After fledging, all chicks remained almost completely dependent on their parents up to an age of 3 months and often longer.
Almost 40% of the fledglings eventually returned to the breeding area. This figure probably reflects post-fledging survival. Age and size at fledging had no effect on a chick's probability of return. Body-weight at fledging had a small positive correlation with the return probability, but this was not statistically significant. We conclude that although slow growth severely reduces a chick's chance of fledging, it probably does not result in irreversible damage causing an increased risk of mortality during the first years after fledging. Apparently, any possible disadvantage associated with small size or low body-weight could be compensated for after fledging.     

Effects of food variability on growth rates, fledging sizes and reproductive success in the Yellow-eyed Penguin Megadyptes antipodes

Effects of a change of diet on growth rates and fledging sizes of Yellow-eyed Penguins Megadyptes antipodes were examined at two breeding areas on South Island, New Zealand, during two breeding seasons. An adverse change in diet was observed in the second season. Evidence for this included depressed growth rates of weight, differential growth of weight and most morphometric parameters between one- and two-chick nests in the second season, lower fledging weights, lower adult body weights, delayed moult, higher chick mortality and higher adult mortality during moult. The change in diet is suggested as being from one including oil-rich prey species, to one of oil-poor species.
Growth rates of first- and second-hatched chicks, and of survivors and non-survivors within a brood were not significantly different in either season, and growth rates of two-chick broods were only slightly slower than one-chick broods for some parameters in the second season. This, and synchronous hatching of chicks, equal egg-size and lack of sibling competition during feeding sessions, suggests that brood reduction is not an option available to Yellow-eyed Penguins, and that food supply may not be a limiting factor in the majority of breeding seasons.
Few changes in growth rates of morphometric parameters at either breeding area, and similar absolute sizes at fledging, indicate that slowing of growth rates of morphometric parameters only occurs when feeding conditions are so bad as to result in mortality and that, although fledging periods may be longer, patterns of development remain essentially unchanged.     

Adult Brünnich's Guillemots Uria lomvia balance body condition and investment in chick growth

To investigate the covariation of adult body condition and nestling growth, we weighed adult Brünnich's Guillemots Uria lomvia rearing chicks at Coats Island, Nunavut, Canada, each year between 1988 and 2002. We estimated chick mass at 14 days for a sample of chicks reared in the same years. Adult mass and chick mass at 14 days were highly correlated, suggesting that, as feeding conditions deteriorate, adults compromise by reducing their own body reserves, while at the same time delivering less food to their offspring. We compared the prediction of the least-squares regression for the Coats Island data with observations made at Digges Island, a much larger colony about 300 km away, where birds are similar in linear body measurements to those at Coats Island and have a similar body mass while incubating. Adult mass at Digges Island averaged 11% less during chick-rearing than during incubation, compared with only a 5% difference at Coats Island. Mean chick mass at 14 days at Digges Island was lower in all years than was observed for chicks at Coats Island in any year. The observed 14-day chick masses at Digges Island in two years were close to values predicted by adult mass and somewhat lower in two other years (those when chick growth was slowest). At Digges Island, the distribution of mass for brooding adults was right skewed and suggested a lower threshold at 800–850 g, below which Brünnich's Guillemots terminate breeding. We conclude that the correlation between adult and chick mass represents a dynamic equilibrium in which adults simultaneously adjust their own energy reserves and their delivery rate to the chick. This compromise must be based on behavioural choices made by individual birds and is unlikely to be a passive consequence of fluctuating conditions.     

The breeding biology and factors affecting reproductive success in rockhopper penguins <Emphasis Type="Italic">Eudyptes chrysocome</Emphasis> at Macquarie Island

Adult mass changes, egg morphometrics, chick growth rates, fledging masses, reproductive success and reasons for reproductive failure were examined in rockhopper penguins at Macquarie Island from 1993/1994 to 1995/1996. Mean arrival masses, growth rates of chicks and fledging masses exhibited inter-annual variability, while egg morphometrics, hatching success (68.0±6.0%) and reproductive success (47.3±8.3%) were constant between years. Reproductive failures occurred primarily during incubation, with the majority of eggs lost to great skuas. Logistic regressions revealed that no variable significantly explained hatching success, and only in 1994/1995 was fledging success significantly correlated with the position of nest in the colony (those in the centre were more successful than those on the periphery). Reproductive success during this study was relatively high, and therefore an assessment during poor years would be instructive, particularly in relation to aspects of the penguins’ foraging ecology.     

Growth and energy expenditure of Wandering Albatross Diomedea exulans chicks

Wandering Albatross Diomedea exulans chicks require 9–10 months to achieve adult body size at fledging, at which time they are also sexually size dimorphic. Because the developmental period spans the winter season, chicks must endure severe winter conditions and variability in provisioning effort by their parents. Thus chicks may adjust their rate of energy utilization to accommodate variations in provisioning, but this has not previously been studied. We followed longitudinally the changes in growth, body composition and oxygen consumption of 10 chicks from the end of the brooding period until fledging on the Crozet Islands. Body mass, culmen length and wing length were measured every 10 days and total body water (TBW) and resting metabolic rate (RMR) were measured monthly. Overall growth followed a logistic curve for all chicks, and sexual dimorphism in body mass appeared as early as the second month of measurements (males being heavier than females). Absolute TBW followed a logistic increase like that of body mass and was significantly higher in males owing to the difference in body mass. Conversely, mass-specific TBW (i.e. the proportion of body mass made up of water) did not differ significantly between male and female chicks. Absolute RMR peaked at 1.5 × adult basal metabolism in midwinter when chicks achieved maximum body mass, but decreased to adult levels by the time chicks fledged. The decrease in absolute RMR following attainment of peak mass is atypical of most seabird chicks (Procellariiformes) and may be explained partly by a reduction in size of the gut when parents reduce provisioning effort. The changes in mass-specific RMR did not differ between sexes but male chicks, being heavier, had higher absolute oxygen consumption and therefore greater energy requirements.     

The effect of parental age,experience and historical reproductive success on wandering albatross (Diomedea exulans) chick growth and survival

Growth and survival of altricial young are influenced by their parents’ abilities to invest in a breeding attempt. As a result, chick growth and survival in one breeding season may be indicative of their parents’ long-term reproductive potential. To determine whether variation in long-term reproductive success is driven by differential breeding investment, parental care and chick growth in wandering albatrosses (Diomedea exulans) were correlated with parental historical reproductive success. Effects of age and breeding experience (determined from past breeding attempts) and pre-laying body condition (mass–size indices) on chick growth and survival also were tested. Longer brooding of chicks increased their survival, but length of chick brooding did not differ between historically unproductive and successful breeders. Past reproductive success also was not correlated with chick growth rates or fledging mass or size. Chick brooding period, chick growth rates, final mass and size were independent of parental body condition. Older and more experienced parents brooded chicks for longer and their chicks grew faster, supporting previous findings that breeding competence is a learnt skill. Chick care and growth characteristics differed more between than within pairs, suggesting that differences in these characteristics are driven by variation among pairs.     

Egg size, composition and offspring quality in some Alcidae (Aves: Charadriiformes)

The composition of fresh and fully developed (pipping) eggs of four alcids, Razorbill, Common guillemot, Brünnich's guillemot and Atlantic puffin was examined. There were no differences in egg composition between the semi-precocial Atlantic puffin and the three species with "intermediate" developmental patterns. The absolute amount of yolk increased with egg size in fresh eggs, and the relative amount remained constant with egg size for Common guillemot and Razorbill, but decreased in the Atlantic puffin. In fully developed eggs chick weight and egg weight were closely correlated, and this was due mainly to larger eggs producing chicks with larger yolk reserves. Under some conditions chicks from larger eggs do better than those from smaller eggs. Several factors influence egg size; a comparison of first and replacement eggs laid by the same females showed that a maternal effect accounted for 60–90% of the variance in egg-size and that laying date accounted for most of the remaining variance.     

Fitness consequences of egg-size variation in the lesser snow goose

We investigated the relationship between eggsize variation and (a) egg hatching success, (b) chick survival to fledging and recruitment, and (c) adult female survival, over 12 years in the lesser snow goose (Anser caerulescens caerulescens). By comparing the means and variances of egg size for successful and unsuccessful eggs, our aim was to assess the relative fitness of eggs of different sizes and to determine the type of selection operating on egg size in this species. As both egg size and reproductive success vary with age in the lesser snow goose we controlled for the effects of female age. Egg-size variation is very marked in this population, varying by up to 52% for eggs hatching successfully. However, there was no relationship between egg size and post-hatching survival of goslings to fledging or recruitment, either within or between broods, pooling across years. Egg size varied significantly between successful and unsuccessful clutches in only 2 of 33 individual year comparisons. First-laid eggs surviving to onset of incubation, and eggs hatching successfully, were on average larger than unsuccessful eggs, but this was probably due to the confounding effects of female age-specific and sequence-specific egg survival. Variance of egg size differed significantly between successful and unsuccessful eggs in only 3 of 24, and 0 of 21, individual year comparisons for pre- and post-hatching survival respectively. We therefore found little evidence for a relationship between egg-size variation and offspring fitness, or for strong directional, normalising or diversifying selection operating on egg size, in the lesser snow goose. In addition, there was only weak support for the hypothesis that egg-size variation is maintained by temporal variation in selection pressure (sensu Ankney and Bisset 1973). It is likely that egg-size variation represents the pleiotropic expression of alleles affecting more general physiological or metabolic processes. While this does not rule out the existence of alleles with more direct effects on egg size we suggest that their contribution to heritable egg size is small.     

Validating growth and development of a seabird as an indicator of food availability: captive‐reared Caspian Tern chicks fed ad libitum and restricted diets

ABSTRACT For seabirds raising young under conditions of limited food availability, reducing chick provisioning and chick growth rates are the primary means available to avoid abandonment of a breeding effort. For most seabirds, however, baseline data characterizing chick growth and development under known feeding conditions are unavailable, so it is difficult to evaluate chick nutritional status as it relates to foraging conditions near breeding colonies. To address this need, we examined the growth and development of young Caspian Terns (Hydroprogne caspia), a cosmopolitan, generalist piscivore, reared in captivity and fed ad libitum and restricted (ca. one‐third lower caloric intake) diets. Ad libitum‐fed chicks grew at similar rates and achieved a similar size at fledging as previously documented for chicks in the wild and had energetic demands that closely matched allometric predictions. We identified three general characteristics of food‐restricted Caspian Tern chicks compared to ad libitum chicks: (1) lower age‐specific body mass, (2) lower age‐specific skeletal and feather size, such as wing chord length, and (3) heightened levels of corticosterone in blood, both for baseline levels and in response to acute stress. Effects of diet restriction on feather growth (10–11% slower growth in diet‐restricted chicks) were less pronounced than effects on structural growth (37–52% slower growth) and body mass (24% lower at fledging age), apparently due to preferential allocation of food resources to maintain plumage growth. Our results suggest that measurements of chick body mass and feather development (e.g., wing chord or primary length) or measurement of corticosterone levels in the blood would allow useful evaluation of the nutritional status of chicks reared in the wild and of food availability in the foraging range of adults. Such evaluations could also inform demography studies (e.g., predict future recruitment) and assist in evaluating designated piscivorous waterbird conservation (colony) sites.