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Concept: Rockhopper penguin


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.

Concepts: Egg, Androgen, Rockhopper penguin, Western Rockhopper Penguin, Penguins


Due to their restricted foraging range, flightless seabirds are ideal models to study the short-term variability in foraging success in response to environmentally driven food availability. Wind can be a driver of upwelling and food abundance in marine ecosystems such as the Southern Ocean, where wind regime changes due to global warming may have important ecological consequences. Southern rockhopper penguins (Eudyptes chrysocome) have undergone a dramatic population decline in the past decades, potentially due to changing environmental conditions. We used a weighbridge system to record daily foraging mass gain (the difference in mean mass of adults leaving the colony in the morning and returning to the colony in the evening) of adult penguins during the chick rearing in two breeding seasons. We related the day-to-day variability in foraging mass gain to ocean wind conditions (wind direction and wind speed) and tested for a relationship between wind speed and sea surface temperature anomaly (SSTA). Foraging mass gain was highly variable among days, but did not differ between breeding seasons, chick rearing stages (guard and crèche) and sexes. It was strongly correlated between males and females, indicating synchronous changes among days. There was a significant interaction of wind direction and wind speed on daily foraging mass gain. Foraging mass gain was highest under moderate to strong winds from westerly directions and under weak winds from easterly directions, while decreasing under stronger easterly winds and storm conditions. Ocean wind speed showed a negative correlation with daily SSTA, suggesting that winds particularly from westerly directions might enhance upwelling and consequently the prey availability in the penguins' foraging areas. Our data emphasize the importance of small-scale, wind-induced patterns in prey availability on foraging success, a widely neglected aspect in seabird foraging studies, which might become more important with increasing changes in climatic variability.

Concepts: Weather, Penguin, Wind, Rockhopper penguin, Western Rockhopper Penguin, Eastern Rockhopper Penguin, Crested penguin, Flightless birds


Although many studies have investigated organohalogenated contaminants (OHCs) in yolk, little is known about the mechanisms and timing of transfer of OHCs from the female to the egg. Vitellogenin, a yolk precursor, has been suggested to play a role in this transport. We here report for the first time the temporal changes in OHC and an index of vitellogenin concentrations in female plasma from the pre-laying period to clutch completion in free-living birds: the southern rockhopper penguin (Eudyptes chrysocome chrysocome) breeding in the Falkland/Malvinas Islands. In addition, OHC concentrations in the corresponding clutches were analysed. OHC concentrations in female plasma and in the yolk of both the first (A-) and the second (B-)eggs followed a similar pattern, with hexachlorobenzene (HCB) > Σpolychlorinated biphenyls (PCBs) > Σdichlorodiphenyltrichloroethanes (DDTs) > Σmethoxylated polybrominated diphenyl ethers (MeO-PBDEs) > Σchlordanes (CHLs) > Σpolybrominated diphenyl ethers (PBDEs) ≈ Σhexachlorocyclohexanes (HCHs). The higher concentrations of MeO-PBDEs compared to PBDEs indicate a diet containing naturally-produced MeO-PBDEs. All OHC compounds except for PBDEs increased from the pre-laying period to A-egg laying and subsequently declined from A-egg laying to B-egg laying, and female plasma vitellogenin showed the same pattern. For ΣPCBs and ΣMeO-PBDEs, we found positive correlations between female plasma during A-egg laying and both eggs, and for HCB between female plasma and A-eggs only. During pre-laying, only ΣMeO-PBDEs correlated between both eggs and female plasma, and no correlations between OHC concentrations in eggs and female plasma were found during B-egg laying, highlighting that maternal transfer of OHCs is time- and compound-specific. Finally, female vitellogenin concentrations did not significantly correlate with any OHC compounds in either female plasma or eggs, and our results therefore did not confirm the suggested role of vitellogenin in the maternal transfer of OHC molecules into their eggs.

Concepts: Correlation and dependence, Pearson product-moment correlation coefficient, Rockhopper penguin, Western Rockhopper Penguin, Eastern Rockhopper Penguin, Crested penguin, Penguins, Eudyptes


Oviparous females need to allocate resources optimally to their eggs in order to maximize their fitness. Among these resources, dietary antioxidants, acquired by females and transferred to the eggs during egg formation, can greatly affect the development and survival of the embryo and chick. In crested penguins, incubation starts after the second and last egg is laid and, as opposed to many other bird species, this egg hatches first, thereby enhancing the survival of the chick. Here, we assessed whether antioxidant and isotopic composition could underlie these differences between eggs within clutches of southern rockhopper penguins (Eudyptes chrysocome chrysocome). The second-laid egg had higher total antioxidant capacity than the first-laid egg, although this was not due to higher antioxidant concentration but to its higher mass. This suggests that resources are allocated by females at a constant rate in both eggs within clutches. Accordingly, we found a strong correlation for isotopic compositions between eggs suggesting that resources were allocated similarly to each egg within the clutch. Overall, we found little evidence for a significant role of antioxidant and isotopic compositions to explain differences in terms of embryo/chick development between eggs in crested penguins. However, since our results suggest a constant rate of antioxidant transfer from females to eggs, limiting the mass of the first-laid egg might represent a strategy for females to spare antioxidant defences and preserve self-maintenance.

Concepts: Nutrition, Antioxidant, Bird, Isotope, Oxygen radical absorbance capacity, Resource allocation, Rockhopper penguin, Western Rockhopper Penguin


Individual specialization in diet or foraging behavior within apparently generalist populations has been described for many species, especially in polar and temperate marine environments, where resource distribution is relatively predictable. It is unclear, however, whether and how increased environmental variability - and thus reduced predictability of resources - due to global climate change will affect individual specialization. We determined the within- and among-individual components of the trophic niche and the within-individual repeatability of δ(13)C and δ(15)N in feathers and red blood cells of individual female southern rockhopper penguins (Eudyptes chrysocome) across 7 years. We also investigated the effect of environmental variables (Southern Annular Mode, Southern Oscillation Index, and local sea surface temperature anomaly) on the isotopic values, as well as the link between stable isotopes and female body mass, clutch initiation dates, and total clutch mass. We observed consistent red blood cell δ(13)C and δ(15)N values within individuals among years, suggesting a moderate degree of within-individual specialization in C and N during the prebreeding period. However, the total niche width was reduced and individual specialization not present during the premolt period. Despite significant interannual differences in isotope values of C and N and environmental conditions, none of the environmental variables were linked to stable isotope values and thus able to explain phenotypic plasticity. Furthermore, neither the within-individual nor among-individual effects of stable isotopes were found to be related to female body mass, clutch initiation date, or total clutch mass. In conclusion, our results emphasize that the degree of specialization within generalist populations can vary over the course of 1 year, even when being consistent within the same season across years. We were unable to confirm that environmental variability counteracts individual specialization in foraging behavior, as phenotypic plasticity in δ(13)C and δ(15)N was not linked to any of the environmental variables studied.

Concepts: Blood, Red blood cell, Climate, Climate change, Blood cells, Blood cell, Rockhopper penguin, Climate pattern


In marine habitats, increasing ocean temperatures due to global climate change may distinctly reduce nutrient and consequently food availability for seabirds. Food availability is a known driver of body mass and reproductive investment in birds, but these traits may also depend on individual effects. Penguins show extreme intra-annual body mass variation and rely on accumulated body reserves for successful breeding. However, no study so far has tested individual consistency and phenotypic responses in body mass and reproductive investment in this taxon. Using a unique dataset on individually marked female and male southern rockhopper penguins (Eudyptes chrysocome chrysocome) across six years, we investigated 1) the individual consistency in body mass (measured at egg laying), body condition and reproductive investment across years, subsequently 2) identified the best-explanatory temperature-related environmental variables for female and male body mass, and 3) tested the effect of female and male body mass on reproductive investment. Body mass, body condition and reproductive investment were all highly repeatable. As body condition should control for the structural size of the birds, the similarly high repeatability estimates for body mass and body condition suggested that the consistent between-individual body mass differences were independent of structural size. This supported the use of body mass for the subsequent analyses. Body mass was higher under colder environmental conditions (positive Southern Annular Mode), but the overall phenotypic response appeared limited. Reproductive investment increased with female but not male body mass. While environmental effects on body mass in our study period were rather small, one can expect that ongoing global climate change will lead to a deterioration of food availability and we might therefore in the long-term expect a phenotypical decline in body mass and reproductive investment.

Concepts: Male, Climate, Weather, Climate change, Solar variation, Rockhopper penguin, Western Rockhopper Penguin, Crested penguin



Reproductive investment affects both offspring and parental fitness and influences the evolution of life histories. Females may vary their overall primary reproductive effort in relation to the phenotypic characteristics of their mate. However, the effects of male quality on differential resource allocation within clutches have been largely neglected despite the potential implications for mate choice and population dynamics, especially in species exhibiting biparental care and brood reduction. Female southern rockhopper penguins Eudyptes chrysocome paired with heavy mates reduced intra-clutch variation in egg and albumen masses. Females paired with new mates also reduced intra-clutch variation in yolk androgen levels. Since both an increased mass and increased androgen concentrations positively influence chick survival under sibling competition, the chances of fledging the whole clutch are likely to be higher for newly formed pairs with heavy males than for previously formed pairs with light males. Interestingly, total clutch provisioning did not vary with male quality. We show for the first time that females vary intra-clutch variation in resource allocation according to male quality. In species with brood reduction, it may be more adaptive for females to modulate the distribution of resources within the clutch according to breeding conditions, than to change their total clutch provisioning.

Concepts: Human, Male, Reproduction, Evolution, Female, Sex, Rockhopper penguin, Western Rockhopper Penguin


Hormonally mediated maternal effects link maternal phenotype and environmental conditions to offspring phenotype. The production of lipid-rich maternal yolk precursors may provide a mechanism by which lipophilic steroid hormones can be transported to developing yolks, thus predicting a positive correlation between yolk precursors in mothers and androgen levels in eggs. Using rockhopper penguins (Eudyptes chrysocome), which produce a two-egg clutch characterized by extreme egg-size dimorphism, reversed hatching asynchrony and brood-reduction, we examined correlations between circulating concentrations of the primary yolk-precursor vitellogenin (VTG) and levels of yolk androgens. Previous work in Eudyptes penguins has shown that egg-size dimorphism is the product of migratory constraints on yolk precursor production. We predicted that if yolk precursors are constrained, androgen transport to developing yolks would be similarly constrained. We reveal positive linear relationships between maternal VTG and androgens in small A-eggs but not larger B-eggs, which is consistent with a migratory constraint operating on the A-egg. Results suggest that intra-clutch variation in total yolk androgen levels depends on the production and uptake of yolk precursors. The brood reduction strategy common to Eudyptes might thus be best described as the result of a migratory constraint.

Concepts: Cholesterol, Androgen, Correlation and dependence, Steroid, Constraint, Rockhopper penguin, Western Rockhopper Penguin, Penguins


Logger technology has revolutionised our knowledge of the behaviour and physiology of free-living animals but handling and logger attachments may have negative effects on the behaviour of the animals and their welfare. We studied southern rockhopper penguin (Eudyptes chrysocome) females during the guard stage in three consecutive breeding seasons (2008/09-2010/11) to evaluate the effects of handling and logger attachment on foraging trip duration, dive behaviour and physiological parameters. Smaller dive loggers (TDRs) were used in 2010/11 for comparison to larger GPS data loggers used in all three seasons and we included two categories of control birds: handled controls and PIT control birds that were previously marked with passive integrative transponders (PITs), but which had not been handled during this study. Increased foraging trip duration was only observed in GPS birds during 2010/11, the breeding season in which we also found GPS birds foraging further away from the colony and travelling longer distances. Compared to previous breeding seasons, 2010/11 may have been a period with less favourable environmental conditions, which would enhance the impact of logger attachments. A comparison between GPS and TDR birds showed a significant difference in dive depth frequencies with birds carrying larger GPS data loggers diving shallower. Mean and maximum dive depths were similar between GPS and TDR birds. We measured little impact of logger attachments on physiological parameters (corticosterone, protein, triglyceride levels and leucocyte counts). Overall, handling and short-term logger attachments (1-3 days) showed limited impact on the behaviour and physiology of the birds but care must be taken with the size of data loggers on diving seabirds. Increased drag may alter their diving behaviour substantially, thus constraining them in their ability to catch prey. Results obtained in this study indicate that data recorded may also not represent their normal dive behaviour.

Concepts: Penguin, Breeding season, Rockhopper penguin, Western Rockhopper Penguin, Eastern Rockhopper Penguin, Crested penguin, Penguins, Eudyptes