Reproduction and survival of Glaucous Gulls breeding in an Arctic seabird colony

ABSTRACT.   Despite being widespread and easily observed, little is known about the life history of Glaucous Gulls ( Larus hyperboreus ). From 1984 to 2007, we examined their breeding biology and demography at Coats Island, Nunavut, Canada, where they nest alongside a colony of 30,000 pairs of Thick-billed Murres ( Uria lomvia ). The gulls fed mainly on murre eggs and chicks and by scavenging adult carcasses. The median age at first breeding was 5 yr, and the mean age was 4.8 ± 0.9 yr. Adult survival was estimated as 0.84 ± 0.03 (SE). The mean clutch size was 2.56 eggs and the mean number of young reared per year was 1.6 (range = 0.9–2.2). Birds reared at the colony provided 40% of recruits. Assuming that survival of locally reared chicks that emigrated was similar to that of chicks that returned to the colony, about 22% of the young gulls survived to breeding age. The timing of breeding by Glaucous Gulls appeared related to the timing of laying by murres. Although the demographic characteristics of Glaucous Gulls in our study were similar to those of populations of other large gulls, adult survival was at the lower end of the range for populations of large Larus gulls. There is some evidence that Glaucous Gulls exhibit lower survival than large gulls breeding in temperate areas, possibly because of contaminant burdens. In general, however, the demographic characteristics of large gulls show little variation and are probably a product of their common phylogeny.     

Male chicks are more costly to rear than females in a monogamous seabird, the Common Murre

Feeding rates and mass loss during chick rearing were comparedfor individually marked parents of male and female Common Murre(Uria aalge) chicks at Great Island, Newfoundland, Canada, from1997–2001. Both parents in this socially monogamous seabirdspecies share parental care duties until colony departure, afterwhich the single chick is fed only by its father. Because murresprovision their single chicks with one clearly visible fishper trip, it is possible to accurately determine whether parentsdifferentially feed male and female chicks. Based on slightlygreater mass of males in adulthood, possibly favored by sexdifferences in breeding roles, we predicted that male nestlingswould be fed more than females. Fathers' feeding rate to sons,but not daughters, increased with chick age, whereas maternalfeeding rate increased with chick age for both sexes. When year-correctedfeeding rates of pairs rearing both sexes were compared, bothmothers and fathers fed their sons significantly more than theirdaughters in the later part of the chick-rearing period. Moreover,parents rearing male chicks lost mass at a significantly higherrate than those rearing females. There was no difference infledging age for sons and daughters. These results indicatethat differential parental allocation occurs and has measurablecosts even in a species with only slight adult sexual dimorphism.     

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.     

Timing of fledging is influenced by glucocorticoid physiology in Laysan Albatross chicks

Fledging is a major life transition for birds, when juveniles move from the safety of a nest into an environment where they must find food and avoid predators. The timing of fledging within a season can have significant effects on future survival and breeding success. Proximate triggers of fledging are unknown: though wing development is likely a primary factor, other physiological changes, such as elevated plasma corticosterone (CORT), may affect fledging behavior. Laysan Albatross (Phoebastria immutabilis) chicks have an extended post−hatching period during which they reach 150% of adult mass. However, approaching fledging, chicks fast for days to weeks and lose mass while still putting energy into feather growth. We evaluated chick morphology and physiology to elucidate proximate triggers of fledging. As in some other species, CORT increased as chicks fasted and lost body mass. At the same time, corticosteroid binding globulin (CBG) declined, thus amplifying free CORT prior to fledging. Once chicks reached a morphological threshold, free CORT levels predicted how long they stayed at the colony: chicks with higher free CORT fledged sooner. To perturb the relationship between body condition, endocrine physiology, and fledging behavior, we supplementally fed chicks for the month before fledging. Fed birds had a slower decrease in body mass, slower decrease in CBG, slower increase in free CORT, and stayed at the colony longer after reaching a morphological threshold. Our study suggests that as chicks lose mass, free CORT acts as a signal of energetic or nutritional state to adjust the timing of fledging.     

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).     

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.     

Seasonal declines in reproductive success of the common tern Sterna hirundo: timing or parental quality?

Reproductive success declines over the course of the breeding season in many bird species. Two categories of hypothesis have been evoked to explain this decline. The “timing” hypothesis suggests that seasonal declines in breeding success are attributable to the date of laying. The “parental quality” hypothesis suggests that seasonal declines result from the fact that young, inexperienced, or low quality birds breed later in the season. To evaluate the relative importance of timing and parental quality, egg exchanges and removals were used to manipulate hatching dates of common terns Sterna hirundo. Indices of quality, attendance, provisioning rates, and reproductive success of birds in three experimental groups (delayed hatch pairs, advanced hatch pairs, and pairs induced to relay) were compared to those of date‐matched controls. Pairs that hatched chicks early raised more chicks than pairs hatching chicks late in the season, regardless of initial laying date. This suggests that hatching chicks early is advantageous in itself. Our results, however, also support the parental quality hypothesis. There was a significant negative relationship between natural laying date and fledging success, independent of hatching date. Differences in chick growth and survival suggest that higher quality adults may be able to compensate for the disadvantages of late hatching dates and achieve similar reproductive success to that of pairs hatching chicks early. We found that pairs hatching chicks late in the season were subject to more incidents of kleptoparasitism than those hatching chicks early. This may be a proximate factor contributing to seasonal declines in reproductive success for common terns, although such a mechanism would not be likely in non‐colonial species. Failure to control for egg quality may have biased the results of some prior egg exchange experiments. Additionally, altered cost of incubation may be an unavoidable confounding factor in studies designed to manipulate timing of breeding.     

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.     

Chick‐provisioning behaviour of two shearwaters breeding in south‐eastern Australia

The chick‐provisioning behaviour of the short‐tailed shearwater Puffinus tenuirostris and the wedge‐tailed shearwater Puffinus pacificus was investigated in a mixed colony on Montague Island, New South Wales, Australia, over two breeding seasons. This colony is located at opposite edges of the breeding distribution of the two species. Frequent weighing techniques were used to determine chick feeding frequency, feed timing, meal size, chick weight loss and indices of food conversion efficiency of the chicks. Short‐tailed shearwater parents fed their chicks larger more infrequent meals than wedge‐tailed shearwater parents. Short‐tailed shearwater chicks demonstrated higher food conversion efficiencies and lower weight loss than wedge‐tailed shearwater chicks, indicating either differences in diet or metabolic rates. The feeding frequency in wedge‐tailed shearwaters also fluctuated more widely than for short‐tailed shearwaters over the two breeding seasons. Despite the fact that the timing of the breeding cycle on Montague Island is almost identical for the two species, these differences in chick provisioning are probably a result of differences in prey type and location, so they may help explain variations in annual breeding success and limits to the distribution of the two species.     

Crop size as an index of chick provisioning in the Greater Flamingo Phoenicopterus roseus

After being fed by their parents, Greater Flamingo chicks store food in their crops, which protrude outwards. We allocated the crop profiles of chicks to four categories to assess the relationship between body mass and crop profile variation, and so determine whether crop size can be used as an accurate index of the amount of food ingested, and to determine the timing and frequency of provisioning. We registered changes in body mass and crop fullness in eight chicks captured with turgid crops and kept in captivity until constant mass was achieved. The meal mass ingested by the chicks during each parental feeding was around 18% of net chick mass and varied greatly with crop profile. Mean transition times between the four crop profile categories ranged from 6 to 14 h. Between 1998 and 2009, 34% of chicks caught for ringing in a breeding colony had empty crops. From crop profiles recorded during the handling of chicks, it was estimated that approximately one‐third of the chicks were fed in the evening and another third during the night. Our results have implications for the estimation of body condition indexes because body mass should be free of the influence of the mass of the food in the crop.     

Provisioning and growth in chicks of Wilson's storm-petrels (Oceanites oceanicus) on King George Island, South Shetland Islands

We present data on chick growth and chick feeding in Wilson's storm-petrel (Oceanites oceanicus) in a colony on King George Island, South Shetland Islands. Chicks were repeatedly weighed and the weight differences over 24 h were corrected for metabolic loss in order to obtain an estimation of meal sizes. Chicks were fed on 93% of the nights (n=688 nights). The average meal size for a single feeding was 8.5 g. Chicks received on average 1.2 feedings per night. These results are compared with data for this species from other locations. There was a trend for increased meal sizes from northern to southern populations, parallel to an increase in the adult mass, indicating that Wilson's storm-petrels carry optimal meal sizes according to their body size and may take advantage of increased food abundance by increasing feeding frequencies. We describe chick growth and discuss the influence of egg size, hatching date and feeding frequency on chick growth. The egg size had a positive influence on tarsus growth and body mass of chicks. Later-hatched chicks started wing growth and finished mass growth at a younger age and reached lower peak masses, indicating that late chicks may adapt to the restricted breeding season in their Antarctic breeding grounds by a more rapid development, but will fledge with a lower degree of development and less resources. Accepted: 22 May 2000     

Inter-annual variation in provisioning behaviour of Southern Rockhopper Penguins Eudyptes chrysocome chrysocome at Staten Island, Argentina

Understanding the relationship between reproductive performance and food availability requires knowledge about many different variables, including such factors as the length of incubation shifts, provisioning rates and patterns, as well as how variability in these factors affects reproductive output. To examine some of the most important aspects of parental investment, we studied the provisioning behaviour and patterns of adult Southern Rockhopper Penguins Eudyptes chrysocome chrysocome breeding at Staten Island, Argentina. We investigated foraging trip duration, provisioning rates and chick survival using adult foraging patterns. Our results show that Rockhopper Penguins had clear sex-specific differences in their provisioning behaviour. Females provision chicks throughout chick rearing. By contrast, males provision chicks only during the crèche stage and at a slightly lower rate than females during this period. Foraging trips increased in length as the breeding season progressed. Rockhopper Penguins from Staten Island performed longer trips throughout the breeding season than do other species of Eudyptes at several other locations. Our results also show differences in parental investment between years that were related to differences in chick survival. We suggest that this was most likely to be related to female rather than male foraging behaviour as only females showed inter-annual differences in their provisioning rates.     

Post-fledging body mass as a determinant of subadult survival in Common Terns Sterna hirundo

Once a bird has fledged it becomes hardly accessible for researchers and consequently knowledge about post-fledging ontogeny is scarce. In this study on juvenile Common Terns (Sterna hirundo) we used an automated transponder-based detection and weighing system at Banter See colony, Northern Germany, which enabled us to investigate body mass growth of post-fledglings and its consequences for their survival until first return to the natal colony when 2 years old. We analysed data from two contrasting breeding seasons, 2000 and 2001, in order to determine inter-year and inter-sex variation of post-fledging parameters assumed to potentially affect subadult survival, such as the period a juvenile is still present at colony surroundings (departure age), its fledging mass and last recorded post-fledging body mass, and hatch date. Using an information-theoretic model selection approach, neither the date of hatching nor the departure age was found to affect survival. The only predictor of survival was last post-fledging body mass whereas fledging mass itself was of minor importance. Although there was weak evidence for an interaction with year, individuals of the cohort 2000, which left the colony area on average 5 g lighter than those reared under the more favourable conditions in 2001, did not exhibit lower return probability. We suggest that under unfavourable conditions selection had eliminated weak individuals prior to fledging or during the post-fledging period. This study underlines the importance of the post-fledging period and its consequences for survival, especially in species with prolonged parental care post-fledging.     

Tracking day and night provides insights into the relative importance of different wader chick predators

Poor reproductive success driven by nest and chick predation severely limits the population recovery of waders breeding on lowland wet grassland. Managing predation requires knowledge of the predators and because these can be grouped into nocturnal or diurnal hunters, detecting the timing of predation can help assess their relative impacts. Wader nest studies investigating the timing of egg predation have identified nocturnal mammals, primarily Red Foxes Vulpes vulpes, as the most important nest predators, but quantifying predator importance for highly mobile wader chicks is more difficult. Manual radiotelemetry can detect whether chicks are alive but cannot detect the time of predation, and predator identity can be determined only in the few cases where remains are recovered. As an alternative we used automatic radio tracking stations (ARTS) to constantly record the signals and predation timing of 179 radiotagged Lapwing Vanellus vanellus chicks, combining this with manual telemetry, inference about predator identity from predated remains and site‐level Fox, mustelid and avian predator activity monitoring. This approach succeeded in detecting the time of predation for 60% of the 155 chicks that were predated. Diurnal chick predation accounted for a larger number of predation events, but nocturnal predation was more intensive in terms of predation likelihood per hour. Mammalian predation during both day and night had a larger impact on chick survival than did avian predation. Raptors were primarily responsible for predation by birds and Foxes for predation by mammals, with Foxes also having a larger influence on daily chick predation rates than other predators. Chick predation increased seasonally, implying that earlier‐hatching breeding attempts are more likely to be successful. Higher Fox, raptor and mustelid activity resulted in higher proportions of chicks being predated by those predators, so quantifying the activity of those three predator groups on a site could be a quicker alternative to studying chicks when investigating which predator species to target with site‐specific predation management.     

Effects of age and body mass on development of diving capabilities of gray seal pups: costs and benefits of the postweaning fast

Development of adequate diving capabilities is crucial for survival of seal pups and may depend on age and body size. We tracked the diving behavior of 20 gray seal pups during their first 3 mo at sea using satellite relay data loggers. We employed quantile analysis to track upper limits of dive duration and percentage time spent diving, and lower limits of surface intervals. When pups first left the breeding colony, extreme (ninety-fifth percentile) dive duration and percentage time spent diving were positively correlated with age, but not mass, at departure. Extreme dive durations and percentage time spent diving peaked at [Formula: see text] d of age at values comparable with those of adults, but were not sustained. Greater peaks in extreme percentage time spent diving occurred in pups that had higher initial values, were older at their peak, and were heavier at departure. Pups that were smaller and less capable divers when they left the colony improved extreme dive durations and percentage time spent diving more rapidly, once they were at sea. Minimum survival time correlated positively with departure mass. Pups that were heavier at weaning thus benefitted from being both larger and older at departure, but smaller pups faced a trade-off. While age at departure had a positive effect on early dive performance, departure mass impacted on peak percentage time spent diving and longer-term survival. We speculate that once small pups have attained a minimum degree of physiological development to support diving, they would benefit by leaving the colony when younger but larger to maximize limited fuel reserves, rather than undergoing further maturation on land away from potential food resources, because poor divers may be able to "catch up" once at sea.     

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.     

Hitting the buffers: conspecific aggression undermines benefits of colonial breeding under adverse conditions

Colonial breeding in birds is widely considered to benefit individuals through enhanced protection against predators or transfer of information about foraging sites. This view, however, is largely based on studies of seabirds carried out under favourable conditions. Recent breeding failures at many seabird colonies in the UK provide an opportunity to re-examine costs and benefits of coloniality under adverse conditions. Common guillemots Uria aalge are highly colonial cliff-nesting seabirds with very flexible parental care. Although the single chick is normally never left alone, more than 50 per cent of offspring were left unattended at a North Sea colony in 2007, apparently because poor conditions forced both parents to forage simultaneously. Contrary to expectation, unattended chicks were not killed by avian predators. Rather, although non-breeders and failed breeders sometimes provided alloparental care, unattended chicks were frequently attacked by breeding guillemots at neighbouring sites, often with fatal consequences. These results highlight a previously unsuspected trade-off between provisioning chicks and avoiding conspecific attacks, and indicate that understanding how environmental conditions affect social dynamics is crucial to interpreting costs and benefits of colonial breeding.     

Adverse effects of agricultural intensification and climate change on breeding habitat quality of Black‐tailed Godwits Limosa l. limosa in the Netherlands

Agricultural intensification is one of the main drivers of farmland bird declines, but effects on birds may be confounded with those of climate change. Here we examine the effects of intensification and climate change on a grassland breeding wader, the Black‐tailed Godwit Limosa l. limosa, in the Netherlands. Population decline has been linked to poor chick survival which, in turn, has been linked to available foraging habitat. Foraging habitat of the nidifugous chicks consists of uncut grasslands that provide cover and arthropod prey. Conservation measures such as agri‐environment schemes aim to increase the availability of chick foraging habitat but have not yet been successful in halting the decline. Field observations show that since the early 1980s, farmers advanced their first seasonal mowing or grazing date by 15 days, whereas Godwits did not advance their hatching date. Ringing data indicate that between 1945 and 1975 hatching dates advanced by about 2 weeks in parallel with the advancement of median mowing dates. Surprisingly, temperature sums at median mowing and hatching dates suggest that while the agricultural advancement before 1980 was largely due to agricultural intensification, after 1980 it was largely due to climate change. Examining arthropod abundance in a range of differently managed grasslands revealed that chick food abundance was little affected but that food accessibility in intensively used tall swards may be problematic for chicks. Our results suggest that, compared with 25 years ago, nowadays (1) a much higher proportion of clutches and chicks are exposed to agricultural activities, (2) there is little foraging habitat left when chicks hatch and (3) because of climate change, the vegetation in the remaining foraging habitat is taller and denser and therefore of lower quality. This indicates that for agri‐environment schemes to make a difference, they should not only be implemented in a larger percentage of the breeding area than the current maxima of 20–30% but they should also include measures that create more open, accessible swards.     

The breeding biology and aspects of the feeding ecology of brown noddies Anous stolidus nesting near Culebra, Puerto Rico, 1985–1989

We have studied brown noddies Anous stolidus breeding on Cayo Noroeste in the Culebra National Wildlife Refuge for the past five years (1985–1989). Daily visits during our residence there each year permitted collection of precise data on arrival chronology, egg-laying and incubation patterns, egg morphometries, chick growth rates, food delivered to chicks and breeding success. Over 150 adults are now individually colour-banded, and each annual chick cohort has been uniquely marked. Some features of the breeding biology of Culebran brown noddies were similar to those reported for pairs at Atlantic and Pacific Ocean colonies; these included duration of egg development, mass of newly hatched chicks, chick growth rates and fledging periods. Other features appear unique to Culebra. These include a tightly synchronous arrival (range of first egg dates: 29 April-4 May) and short egg-laying period (about six weeks), a consistently high breeding success each year (average 84% hatching success, 88% chicks fledged from eggs hatched, 0.79 chicks fledged per pair), and an unusually narrow range of food items (two fish species) taken by adults for their own needs and those of chicks. Observations of the direction taken by adults departing the colony suggest a predictable and productive foraging area along a prominent east-west shelf-break located c. 20 km to the north. We conclude that during these five years, brown noddies on Cayo Noroeste were not limited by food during the breeding season.     

Annual variation in breeding biology of gentoo penguins, Pygoscelis papua, at Bird Island, South Georgia

The breeding biology of the gentoo penguin, Pygoscelis papua , was studied over a three-year period (1986–1988) at Bird Island, South Georgia, with particular reference to birds of known age or breeding experience. Laying date varied significantly between all three years, being three weeks later in 1987, when the breeding population decreased markedly. Factors involved in the timing of breeding are discussed. Within years egg-laying was highly synchronous: 95% of clutches were initiated in 14·5 days or less. The incubation period was 35 days and the laying interval, between the two eggs, 3·3–3·4 days. Chicks creched when 25–30 days old, and this varied between years, possibly related to food supply and chick growth. Chicks left the colony for the first time between 75 and 85 days of age. The breeding population at Bird Island decreased by 20% and increased by 84% in successive years during the study period. Breeding success (chicks fledged per egg laid) varied between 0·33 and 0·65 within colonies, but for the whole island was very consistent over the three years: 0·45, 0·51 and 0·47. Overall, colony differences were not correlated between years. Disturbance from Antarctic fur seals, Arctocephalus gazella , is suggested as the cause of consistently lower breeding success at one colony. Mean egg weight varied annually, and with age of the breeding bird, nest location and, in one year, with laying date. Young, first-time breeders laid smaller eggs and had lower breeding success compared to older, experienced birds, similar to other seabirds. However, they differed from other species in laying on average earlier than older birds. The relationship between age, egg weight, laying date and breeding success is discussed in relation to predation and seasonal food supply.