Intra-Clutch Ratio of Yolk Progesterone Level Changes with Laying Date in Rockhopper Penguins: A Strategy to Influence Brood Reduction?

Hatching asynchrony in avian species generally leads to a size hierarchy among siblings, favouring the first-hatched chicks. Maternally deposited hormones affect the embryo and chick's physiology and behaviour. It has been observed that progesterone, a hormone present at higher levels than other steroid hormones in egg yolks, is negatively related to body mass in embryos, chicks and adults. A differential within-clutch progesterone deposition could therefore be linked to the size hierarchy between siblings and to the resulting brood reduction. We tested whether yolk progesterone levels differed between eggs according to future parental ability to feed the entire clutch in wild rockhopper penguins Eudyptes chrysocome. This species presents a unique reversed egg-size dimorphism and hatching asynchrony, with the larger second-laid egg (B-egg) hatching before the smaller first-laid egg (A-egg). Yolk progesterone levels increased only slightly with female body mass at laying. However, intra-clutch ratios were not related to female body mass. On the other hand, yolk progesterone levels increased significantly with the date of laying onset for A-eggs while they decreased for B-eggs. Early clutches therefore had proportionally more progesterone in the B-egg compared to the A-egg while late clutches had proportionally less progesterone in the B-egg. We propose that females could strategically regulate yolk progesterone deposition within clutches according to the expected food availability during chick growth, an adaptive strategy to adjust brood reduction to conditions. We also discuss these results, relating to yolk progesterone, in the broader context of other yolk steroids

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

Interpreting ELISA analyses from wild animal samples: some recurrent issues and solutions

Many studies in disease and ecological immunology rely on the use of assays that quantify the amount of specific antibodies (immunoglobulin) in samples. Enzyme‐linked immunosorbent assays (ELISAs) are increasingly used in ecology due to their availability for a broad array of antigens and the limited amount of sampling material they require. Two recurrent methodological issues are nevertheless faced by researchers: (1) the limited availability of immunological assays and reagents developed for non‐model species, and (2) the statistical determination of the cut‐off threshold used to distinguish individual samples that are likely to have or not to have antibodies against a specific antigen. Here, we outline two solutions to deal with these issues. First, we show that implementing two assays with differing detection methods can help validate the use of reagents, such as antibodies, in species different from their intended target. We illustrate this by comparing the quantification of specific vaccinal antibodies against Newcastle disease virus (NDV) using two ELISA approaches in four seabird species (Cory's shearwater, European shag, European storm petrel and Southern rockhopper penguin). Second, we provide a simple way to determine from the distribution of ELISA values whether the assayed samples are likely to be made of a single group of individuals (likely negative) or of two groups of individuals (negative and positive). We illustrate the use of this approach with two independent datasets: NDV antibody levels following vaccination and anti‐Borrelia antibody levels following natural exposure. The practical implementation of these methodological approaches could provide a way to efficiently apply ELISAs and other immune‐based assays to address questions in the growing fields of ecological immunology and disease ecology

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

Are acoustical parameters of begging call elements of thin-billed prions related to chick condition?

Chicks of burrowing petrels use begging calls to advertise their hunger levels when parents arrived at the nest. In a previous study, adult thin-billed prions Pachyptila belcheri responded to higher begging call rates of their single chick by regurgitating larger meals. We tested whether acoustic parameters of begging call elements may also be involved in signalling. To describe variation in begging, we determined begging session parameters, namely the duration, number of calls and the mean and maximum rate of calling. We then digitised calls and carried out a semi-automatic extraction of six acoustic parameters of call elements, including mean and maximum acoustic frequency, the length of call elements and the location of the maximum frequency and amplitude within calls. Chicks showed strong individual differences in all parameters. While the session parameters were correlated with body condition and with the meal size the chick received, none of the acoustic parameters were related to body condition and provisioning. A cross-fostering experiment showed the same pattern, as only session parameters changed related to an experimentally altered body condition, while acoustical cues appear to play no role in signalling hunger levels. We suggest that this may be explained by the absence of sibling competition in these birds. As parents do not need to decide which chick to feed, immediate information on condition at the time of adult arrival may not be required

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. © 2019, The Author(s)

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.Peer reviewe

Parental behavior controls asynchronous hatching, but not incubation period, in the Magellanic Penguin : a commentary on Rebstock and Boersma (2011)

Rebstock and Boersma (2011) recently explored variation in the temperature and incubation period of eggs within clutches of the Magellanic Penguin (Spheniscus magellanicus). They defined incubation period as “the time from the laying of an egg to its hatching” and concluded that parental behavior explained why the incubation period of second eggs was shorter than that of first eggs and controlled asynchronous hatching. While we agree that parents influenced asynchronous hatching by delaying the onset of incubation, we argue that their conclusions are based on an unconventional definition of incubation period. They included the period before the delayed onset of incubation in the incubation period, which leads to confusion. We state that the incubation period cannot include the time before the (delayed) onset of incubation when parents are not warming the eggs. As regards this latter and widely accepted definition, Rebstock and Boersma (2011) provided a large dataset showing that incubation consistently lasted for 39 days, whenever the onset of incubation, for both first and second eggs. The divergence in the definition of “incubation period” and the failure to consider the “onset of incubation” for a species in which the first egg is not incubated immediately after laying led to confusion in the interpretation of the results and conclusions from Rebstock and Boersma (2011)