Potential for redistribution of post-moult habitat for Eudyptes penguins in the Southern Ocean under future climate conditions.

Anthropogenic climate change is resulting in spatial redistributions of many species. We assessed the potential effects of climate change on an abundant and widely distributed group of diving birds, Eudyptes penguins, which are the main avian consumers in the Southern Ocean in terms of biomass consumption. Despite their abundance, several of these species have undergone population declines over the past century, potentially due to changing oceanography and prey availability over the important winter months. We used light-based geolocation tracking data for 485 individuals deployed between 2006 and 2020 across 10 of the major breeding locations for five taxa of Eudyptes penguins. We used boosted regression tree modelling to quantify post-moult habitat preference for southern rockhopper (E. chrysocome), eastern rockhopper (E. filholi), northern rockhopper (E. moseleyi) and macaroni/royal (E. chrysolophus and E. schlegeli) penguins. We then modelled their redistribution under two climate change scenarios, representative concentration pathways RCP4.5 and RCP8.5 (for the end of the century, 2071-2100). As climate forcings differ regionally, we quantified redistribution in the Atlantic, Central Indian, East Indian, West Pacific and East Pacific regions. We found sea surface temperature and sea surface height to be the most important predictors of current habitat for these penguins; physical features that are changing rapidly in the Southern Ocean. Our results indicated that the less severe RCP4.5 would lead to less habitat loss than the more severe RCP8.5. The five taxa of penguin may experience a general poleward redistribution of their preferred habitat, but with contrasting effects in the (i) change in total area of preferred habitat under climate change (ii) according to geographic region and (iii) the species (macaroni/royal vs. rockhopper populations). Our results provide further understanding on the regional impacts and vulnerability of species to climate change.

Phylogeography, Population Structure, and Species Delimitation in Rockhopper Penguins (Eudyptes chrysocome and Eudyptes moseleyi).

Rockhopper penguins are delimited as 2 species, the northern rockhopper (Eudyptes moseleyi) and the southern rockhopper (Eudyptes chrysocome), with the latter comprising 2 subspecies, the western rockhopper (Eudyptes chrysocome chrysocome) and the eastern rockhopper (Eudyptes chrysocome filholi). We conducted a phylogeographic study using multilocus data from 114 individuals sampled across 12 colonies from the entire range of the northern/southern rockhopper complex to assess potential population structure, gene flow, and species limits. Bayesian and likelihood methods with nuclear and mitochondrial DNA, including model testing and heuristic approaches, support E. moseleyi and E. chrysocome as distinct species lineages with a divergence time of 0.97 Ma. However, these analyses also indicated the presence of gene flow between these species. Among southern rockhopper subspecies, we found evidence of significant gene flow and heuristic approaches to species delimitation based on the genealogical diversity index failed to delimit them as species. The best-supported population models for the southern rockhoppers were those where E. c. chrysocome and E. c. filholi were combined into a single lineage or 2 lineages with bidirectional gene flow. Additionally, we found that E. c. filholi has the highest effective population size while E. c. chrysocome showed similar effective population size to that of the endangered E. moseleyi. We suggest that the current taxonomic definitions within rockhopper penguins be upheld and that E. chrysocome populations, all found south of the subtropical front, should be treated as a single taxon with distinct management units for E. c. chrysocome and E. c. filholi.

Extreme intra-clutch egg size dimorphism is not coupled with corresponding differences in antioxidant capacity and stable isotopes between eggs.

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.

Mates but not sexes differ in migratory niche in a monogamous penguin species.

Strong pair bonds generally increase fitness in monogamous organisms, but may also underlie the risk of hampering it when re-pairing fails after the winter season. We investigated whether partners would either maintain contact or offset this risk by exploiting sex-specific favourable niches during winter in a migratory monogamous seabird, the southern rockhopper penguin Eudyptes chrysocome. Using light-based geolocation, we show that although the spatial distribution of both sexes largely overlapped, pair-wise mates were located on average 595 ± 260 km (and up to 2500 km) apart during winter. Stable isotope data also indicated a marked overlap between sex-specific isotopic niches (δ¹³C and δ¹5N values) but a segregation of the feeding habitats (δ¹³C values) within pairs. Importantly, the tracked females remained longer (12 days) at sea than males, but all re-mated with their previous partners after winter. Our study provides multiple evidence that migratory species may well demonstrate pair-wise segregation even in the absence of sex-specific winter niches (spatial and isotopic). We suggest that dispersive migration patterns with sex-biased timings may be a sufficient proximal cause for generating such a situation in migratory animals.

Migratory constraints on yolk precursors limit yolk androgen deposition and underlie a brood reduction strategy in rockhopper penguins.

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.

Yolk androgen deposition without an energetic cost for female rockhopper penguins: a compensatory strategy to accelerate brood reduction?

Whether androgen deposition in eggs is physiologically costly for female birds has remained a crucial but unsolved question, despite a broad use of this assumption in functional studies. We tested whether females depositing high androgen concentrations experienced higher mass losses than females depositing low androgen concentrations. Analysing female body mass change during egg formation in rockhopper penguins (Eudyptes chrysocome chrysocome), we observed no energetic cost to androgen deposition. Nevertheless, lighter females laid eggs with higher yolk androgen concentrations. This relationship existed only for the second-laid egg (B-egg), but not for the first-laid egg (A-egg). Since the B-egg is usually the first to hatch and the only one to produce a fledging chick, we hypothesize that differential yolk androgen deposition may be an adaptive strategy for females to affect brood reduction.

Measuring corticosterone in seabird egg yolk and the presence of high yolk gestagen concentrations.

Large inter-species differences have been found in yolk corticosterone amounts in avian eggs. While some studies have failed to detect significant amounts of corticosterone, in other species high amounts have been recorded, such as in a recent study of southern rockhopper penguins Eudyptes chrysocome chrysocome. However, attention has been drawn recently to the fact that many antibodies for corticosterone measurement cross-react with other steroids present in the yolk. In particular, progesterone and related substances can occur in yolk in high concentrations, such that also low cross-reactions of corticosterone assays may lead to measurement errors. We thus performed high-performance liquid chromatographic (HPLC) analyses of yolk extracts and determined the concentration of immunoreactive corticosterone, as well as cross-reacting progesterone and cortisol in egg yolks of southern rockhopper penguins and imperial shags Phalacrocorax atriceps albiventer. We found that high gestagen concentrations in the yolk result in large measurement errors for yolk corticosterone, even when the cross-reactivity seems small. This was observed for both species. We further found species-specific differences in the actual corticosterone amounts present in the egg yolks.

Reversed hatching order, body condition and corticosterone levels in chicks of southern rockhopper penguins (Eudyptes chrysocome chrysocome).

In altricial and semi-altricial species, asynchronous hatching gives the first chicks to hatch an initial advantage over other siblings and often leads to the elimination of the smallest chicks. Both baseline corticosterone and acute stress-induced corticosterone levels have been shown to be higher in food deprived chicks than in chicks fed ad libitum. However, first-hatched chicks have also been shown to exhibit higher corticosterone levels than last-hatched chicks, suggesting an influence of the initial differences between eggs on corticosterone levels. We subjected single-chicks of southern rockhopper penguins Eudyptes chrysocome chrysocome to a standardised capture-stress protocol. In this species having very dimorphic two-egg clutches, we examined whether corticosterone levels were different between the two chick categories and tested for the effect of body condition controlled by the chick category. Neither body sizes, nor corticosterone levels differed between A- and B-chicks at 18 days. In contrast to baseline corticosterone levels, acute stress-induced levels of corticosterone were negatively correlated to body condition: chicks with a good body condition had lower acute stress-induced levels of corticosterone than chicks with a poor condition, whatever the chick category. Our results do not support the idea that initial differences in egg characteristics could drive the difference in corticosterone levels between siblings. On the contrary, they show that the A-egg of rockhopper penguins has, when reared alone, the same intrinsic potential to develop into a fledged chick as the B-egg. Later differences in body condition appear to lead to variation in the acute stress-induced levels of corticosterone.

Acute stress hyporesponsive period in nestling Thin-billed prions Pachyptila belcheri.

When confronted with acute stressors, vertebrates show a highly conserved evolved sequence of physiological, hormonal and behavioural responses, including the activation of the hypothalamic-pituitary-adrenal axis. Many young vertebrates show a stress hyporesponsive period, where they exhibit a reduced glucocorticoid response. Here, we analyzed the stress response of nestling Thin-billed prions Pachyptila belcheri and compared chicks with different previous experience with capture and handling. We found that chicks had a stress response, but baseline and peak levels were below those measured in adults. The stress response of the chicks was rapid and followed by fast recovery, such that the total amount of corticosterone released in response to handling was very much lower in chicks than adults. These results indicate that nestling Thin-billed prions exhibit a stress hyporesponsive period. This was not due to habituation, as CORT measurements at baseline and elevated levels were similar in chicks handled daily and naïve chicks. The comparison with other published studies showed that the stress response of chicks usually peaks earlier and lower than in adults, and researchers should take care to measure stress-induced levels at an appropriate sampling time.

Maternal deposition of yolk corticosterone in clutches of southern rockhopper penguins (Eudyptes chrysocome chrysocome).

High corticosterone levels can have deleterious effects in developing avian embryos and chicks. Therefore, it may be adaptive for avian mothers to reduce corticosterone transfer to their eggs. However, until now, data about the active or/and passive role of mothers in corticosterone transfer to eggs are inconclusive. Here, we study maternal investment into A- and B-eggs of southern rockhopper penguins (Eudyptes chrysocome chrysocome). This species exhibits reversed hatching asynchrony and provides a unique model to test whether there is a strategic investment in corticosterone favoring the B-chick, which is most likely to survive. We found that rockhopper penguins had the highest yolk concentrations of any wild bird species studied so far. Contrary to our expectations, B-eggs had more yolk corticosterone both in concentration and in quantity than A-eggs, independently of the laying period and the level of human disturbance. Additionally, females deposited more yolk corticosterone in their eggs when they were disturbed. However, this disturbance effect was particularly strong for A-eggs and for late-laid eggs. The present data support neither the predictions for an active regulation nor for a passive deposition, and hormone deposition mechanisms still need to be explored. The adaptive value, if any, of high yolk corticosterone is presently unknown.

Differences in the stable isotope signatures of seabird egg membrane and albumen--implications for non-invasive studies.

In many bird species, egg membranes can be obtained non-invasively after the chicks have hatched, and stable isotope analysis of egg membranes can be used to study the diet and foraging distribution of these birds during egg formation. It has been suggested that the enrichment factors of albumen and egg membranes differ for 13C, but are similar for 15N. In this study, we compared carbon and nitrogen stable isotopes of the membranes and albumen of individual eggs of three wild seabird species, the Southern Rockhopper penguin Eudyptes chrysocome, the Imperial shag Phalacrocorax atriceps albiventer, and the Thin-billed prion Pachyptila belcheri. We also included chicken eggs for comparison. Egg membranes were generally enriched in 13C, compared with albumen. The difference varied between species, with 2.1 per thousand in Rockhopper penguins, 1.6 per thousand in Imperial shags, but only 0.5 per thousand in Thin-billed prions and 0.4 per thousand in chicken eggs. Egg membranes were slightly enriched in 15N in Imperial shags (0.9 per thousand) and chickens (0.5 per thousand), compared with albumen, while there was no difference for Thin-billed prions and Rockhopper penguins. The isotopic values of carbon and nitrogen were correlated between albumen and egg membranes of individual eggs, suggesting that egg membranes can be used reliably to investigate trophic differences between individuals, seasons or colonies. Species-specific mathematical corrections could be used to compare results across studies that use different egg components.

Adaptive responses of animals to climate change are most likely insufficient.

Biological responses to climate change have been widely documented across taxa and regions, but it remains unclear whether species are maintaining a good match between phenotype and environment, i.e. whether observed trait changes are adaptive. Here we reviewed 10,090 abstracts and extracted data from 71 studies reported in 58 relevant publications, to assess quantitatively whether phenotypic trait changes associated with climate change are adaptive in animals. A meta-analysis focussing on birds, the taxon best represented in our dataset, suggests that global warming has not systematically affected morphological traits, but has advanced phenological traits. We demonstrate that these advances are adaptive for some species, but imperfect as evidenced by the observed consistent selection for earlier timing. Application of a theoretical model indicates that the evolutionary load imposed by incomplete adaptive responses to ongoing climate change may already be threatening the persistence of species.

Organohalogenated contaminants in plasma and eggs of rockhopper penguins: Does vitellogenin affect maternal transfer?

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.

Is individual consistency in body mass and reproductive decisions linked to individual specialization in foraging behavior in a long-lived seabird?

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.

Two eggs, two different constraints: a potential explanation for the puzzling intraclutch egg size dimorphism in Eudyptes penguins.

Phenotypic plasticity and phenotypic stability are major components of the adaptive evolution of organisms to environmental variation. The invariant two-egg clutch size of Eudyptes penguins has recently been proposed to be a unique example of a maladaptive phenotypic stability, while their egg mass is a plastic trait. We tested whether this phenotypic plasticity during reproduction might result from constraints imposed by migration (migratory carry-over effect) and breeding (due to the depletion of female body reserves). For the first time, we examined whether these constraints differ between eggs within clutches and between egg components (yolk and albumen). The interval between colony return and clutch initiation positively influenced the yolk mass, the albumen mass, and the subsequent total egg mass of first-laid eggs. This time interval had only a slight negative influence on the yolk mass of second-laid eggs and no influence on their albumen and subsequent total masses. For both eggs, female body mass at laying positively influenced albumen and total egg masses. Female investment into the entire clutch was not related to the time in the colony before laying but increased with female body mass. These novel results suggest that the unique intraclutch egg size dimorphism exhibited in Eudyptes penguins, with first-laid eggs being consistently smaller than second-laid eggs, might be due to a combination of constraints: a migratory carry-over effect on the first-laid egg and a body reserve depletion effect on the second-laid egg. Both these constraints might explain why the timing of reproduction, especially egg formation, is narrow in migratory capital breeders.

Individual consistency and phenotypic plasticity in rockhopper penguins: female but not male body mass links environmental conditions to reproductive investment.

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.

Good days, bad days: wind as a driver of foraging success in a flightless seabird, the southern rockhopper penguin.

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.

Females paired with new and heavy mates reduce intra-clutch differences in resource allocation.

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.

Evaluating the impact of handling and logger attachment on foraging parameters and physiology in southern rockhopper 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.

Penguin chicks benefit from elevated yolk androgen levels under sibling competition.

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