Parental body size affects meal size in the chinstrap penguin (Pygoscelis antarctica)

Meal size of chinstrap penguins (Pygoscelis antarctica) was estimated through weighing adult breeders before and after food transfer during the crèche phase. Mean meal size delivered by each parent was 630.2 ± 178.3 g and was independent of breeding date, number of chicks and sex. Only body size as estimated by flipper length had a significant effect on meal size, suggesting an advantage of large body size which allows maximization of food carried per visit. Received: 22 September 1997 / Accepted: 16 December 1997     

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

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

Absence of extra-pair fertilisations in the Chinstrap Penguin Pygoscelis antarctica

We analysed genetic relatedness through DNA-fingerprinting in 38 two-chick families of Chinstrap Penguins Pygoscelis antarctica in the Vapour Col colony on Deception Island, Antarctica. Band-sharing coefficients between males and chicks in their nests, females and chicks in their nests and brood mates were very close to 0.50 and were not significantly different. Presumably unrelated pair mates showed much lower band-sharing coefficients (0.12), which differed significantly from those between putative parents and their chicks and between presumed siblings. No case of extra-pair paternity was detected according to the presence of novel bands or to band-sharing coefficients. Monogamy in the study population is not only social but also genetic.     

Experimental assessment of the effects of gastrointestinal parasites on offspring quality in chinstrap penguins (Pygoscelis antarctica)

Parasites reduce host fitness and consequently impose strong selection pressures on their hosts. It has been hypothesized that parasites are scarcer and their overall effect on hosts is weaker at higher latitudes. Although Antarctic birds have relatively low numbers of parasites, their effect on host fitness has rarely been investigated. The effect of helminth parasitism on growth rate was experimentally studied in chinstrap penguin (Pygoscelis antarctica) nestlings. In a total of 22 two-nestling broods, 1 nestling was treated with anthelminthics (for cestodes and nematodes) while its sibling was left as a control. Increased growth rate was predicted in de-wormed nestlings compared to their siblings. As expected, 15 days after treatment, the experimental nestlings had increased body mass more than their siblings. These results show a non-negligible negative effect of helminth parasites on nestling body condition that would presumably affect future survival and thus fitness, and it has been suggested there is a strong relationship between body mass and mortality in chinstrap penguins.     

Absence of haematozoa in a wild chinstrap penguin Pygoscelis antarctica population

No haematoza were detected in 40 adult and 58 nestling wild chinstrap penguins (Pygoscelis antarctica) from Antarctica examined by blood smear. Received: 27 November 1996 / Accepted: 24 March 1997     

Adult aggression during the post-guard phase in the chinstrap penguin Pygoscelis antarctica

We report an observational study of aggressive behaviour by adults during the post-guard phase in the chinstrap penguin. The study was carried out in one subcolony where all the individuals (breeders, chicks and failed breeders) were banded, as well as in nine other subcolonies where individuals were not identifiable. Breeding adults were more aggressive towards unrelated chicks when they were feeding their own chicks than in other contexts. Chicks were also attacked at high rates by adults that did not belong to their own subcolonies. The evidence suggests that some aggressions to chicks by adults may function to avoid interference from unrelated chicks during food transfers, but our data also show that aggressive interactions are common in contexts other than chick feeding. However, aggression by adults did not show any clear spatial directionality or purpose of shepherding the chicks, and these tended not to move in any particular direction when attacked.     

Breeding time, health and immune response in the chinstrap penguin Pygoscelis antarctica

Health status and immunocompetence have been proposed as important factors affecting individual variation in the attainment of breeding condition in birds. We studied individual variation in serological variables indicating health status (blood sedimentation rate, haematocrit, `buffy coat' layer, proportions of different types of leucocytes) in two groups of breeding chinstrap penguins Pygoscelis antarctica with breeding dates 9 days apart. We sampled these individuals shortly after hatching of their young and at the end of the chick-raising period. A group of failed breeders was also sampled. Birds of both sexes were included. We also measured the T-cell-mediated immune response as indicated by an in vivo hypersensitivity response to an intradermal injection of a mitogen (phytohaemagglutinin) in early and late breeders. Sex had no significant effect on most variables. Late breeders had poorer health (more leucocytes, especially heterophils and lymphocytes) and a lower T-cell-mediated immune response than early breeders. Failed breeders were more similar to late than to early breeders. Early breeders suffered a decline in health status throughout the chick-raising period. The impact of pathogens on variation in life history traits in avian populations may be important even in extreme Antarctic environments. Received: 18 December 1997 / Accepted: 1 March 1998     

Heat loss and site-dependent fecundity in chinstrap penguins (Pygoscelis antarctica)

We study the effects of heat loss and nest site quality on fecundity in a chinstrap penguin (Pygoscelis antarctica) colony on Deception Island, Antarctica. During the austral summers of 1990–1991, 1993–1994, and 1995–1996, 441 randomly selected nests were analyzed. Penguins breeding in the center of subcolonies hatched significantly earlier and had larger broods and than those nesting near or at the edge of the subcolonies. These differences, however, were significantly affected by interaction between year and nest location, being highly significant in colder years, when peripheral nests produce fewer young. Analysis of the rate of heat loss showed that penguins breeding at the edge of subcolonies lost heat twice as rapidly as those breeding in the interior of the subcolony. In a re-occupation experiment, evacuated center nests were re-occupied almost ten times as rapidly as edge nests. An analysis of mean fecundity in the period 1991–1996 and mean wind chill suggested that most of the variability in fecundity among years was related to differences in the rate of heat loss. Subcolonies tend to be as circular as possible, thereby decreasing the proportion of edge nests as the size of subcolony increases. Our results support the site-dependent fecundity hypothesis.     

Factors that affect hatching asynchrony in the chinstrap penguin (Pygoscelis antarctica)

We examined if laying intervals and hatching asynchronies are related in a chinstrap penguin (Pygoscelis antarctica) population in the South Orkneys. The lack of association between the two variables, as well as data on brood patch development, indicated that incubation did not begin immediately after the first egg was laid. This suggests that longer laying intervals may be compensated by longer delays in the onset of full incubation. Hatching asynchrony increased with within-clutch egg size asymmetry, decreased with breeding date, and was related to the laying order of the eggs according to size. However, only egg-size asymmetry remained significant when controlling for the other variables. We conclude that more asymmetric clutches were more asynchronous, although a large part of the variation in hatching asynchrony remains unexplained.     

Parental physiological condition and reproductive success in chinstrap penguins (Pygoscelis antarctica)

Recent studies suggest that parental resource allocation may be the most important factor explaining differences in reproductive output among parents. That said at least two different hypotheses of balance between parental foraging effort and resource allocation have been proposed. First, parents with high foraging effort have high reproductive success. Second, parents with higher allocation of resources to offspring have high reproductive success. We tested the second hypothesis using chinstrap penguins (Pygoscelis antarctica) as a model. We evaluated nutritional condition of the parents using blood urea, uric acid, creatine kinase, and cholesterol levels. We evaluated reproductive success according to total mass of the brood and asymmetries inside the brood. We measured the degree of asymmetry using weight and culmen length. Generalized linear models were used to examine relationships between adult plasma urea levels with year, nest position, and degree of asymmetry in chicks. Our results demonstrate that lighter broods were more asymmetric and associated with lower values of adult plasma urea, uric acid, and creatine kinase. We interpret these findings as evidence that the birds allocate fewer resources to their chicks than adults with more symmetric broods are.     

Haematocrit values in chinstrap penguins (Pygoscelis antarctica): variation with age and reproductive status

We studied haematocrit variation with age and reproductive status in chinstrap penguins (Pygoscelis antarctica) at Deception Island, South Shetlands. There was a significant difference in haematocrit values between different age groups (adults vs nestlings and juveniles). Adults had a higher haematocrit value than juveniles and nestlings and juveniles had higher haematocrit values than nestlings. There was also a significant difference in haematocrit values between adults of different status (breeding vs moulting). Breeding adults had higher haematocrit values than moulting adults. The observed differences between age groups were thought to be the result of different oxygen demands due to exercise. The observed differences between adults of different status could reflect the different demands associated with reproduction and moult fasting.     

Population decline of chinstrap penguins (Pygoscelis antarctica) on Deception Island, South Shetlands, Antarctica

Population changes of top predators can provide key indications of environmental quality. In the Antarctic ecosystem, population dynamics of top predators like penguins may yield important information about how the environment is changing. From 1991–1992 to 2008–2009 censuses of the chinstrap penguin (Pygoscelis antarctica), breeding population of Vapour Col in Deception Island (South Shetlands Islands) were carried out. Censuses were conducted on 19 sub-colonies, mainly in December around the time of peak hatching. Nest counts were taken from photographs that were recorded from a standard location. Some censuses were also made in mid-January in 1999, 2006, 2007 and 2008. Our results reveal that the population has declined by 36?% when comparing data from 1991 with those from 2008. The strongest decline occurred since 2000. No temporal trends were detected in reproductive success. Declines of Chinstrap penguin populations constitute a general pattern in the South Shetlands, and it has been suggested to be related to climate change through effects of reduction in sea-ice extent during winter and a consequent decline in the abundance of krill.     

Phenotypic selection on morphology at independence in the Chinstrap penguin Pygoscelis antarctica

Every year, shortly after the emancipation of chicks at our study colony (Deception Island, South Shetlands), hundreds of carcasses of presumably starved Chinstrap penguin Pygoscelis antarctica chicks are washed on the shore. In 1997 we measured the flippers of fresh carcasses and compared their lengths with those of live chicks about to become independent. There was a highly significant difference of 6.5 mm between both distributions, which suggests strong directional phenotypic selection on skeletal size operating through its association with body reserves at independence. Given that heritabilities of flipper length and body weight measured on 36 families are 0.73(± 0.32) and 0.075(± 0.081), and that both characters show a genetic correlation of 0.44(± 0.14), we can expect an evolutionary response to this selection episode. Assuming that the target of selection is weight at emancipation (heavier chicks carry proportionally larger reserves), and that flipper length changes as a consequence of its genetic correlation with weight, we can predict a response of 1.32–2.87 mm or 0.23–0.51 standard deviation units for flipper length. This substantial evolutionary response may be countered by other selective pressures affecting other life stages of these birds. Selection on reserve storage capacity at independence may affect morphological traits also in other species.     

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     

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

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

Limited male incubation ability and the evolution of egg size in shorebirds

In bird species where males incubate but are smaller than females, egg size may be constrained by male body size, and hence ability to incubate the eggs. Using data from 71 such shorebird species, we show that egg size decreases as the degree of female-biased sexual size dimorphism increases, after controlling for female body mass. Relative egg size was not related to mean clutch size. However, when controlling for mating system, the relationship between female-biased sexual size dimorphism and relative egg size was only significant in polyandrous species. The relatively small eggs of socially polyandrous shorebirds have previously been explained as an energy-saving strategy associated with the production of multiple clutches. Our findings suggest that egg size evolution is better explained by male incubation limitation in these birds.