Beak colour dynamically signals changes in fasting status and parasite loads in king penguins

This research was funded by the French Polar Institute (IPEV–Research Program 119) and the French National Centre for Scientific Research (CNRS-INEE). Field logistic support was provided by Terres Australes et Antarctiques Françaises. Q.S. was funded by a doctoral fellowship from the Ministère Français de l’Education Supérieur et de la Recherche. We thank all over-wintering assistants: Benoit Gineste, Sylvia Pardonnet, Laureline Durand, Emilie Lefol and Hédi Saadaoui for field work and Emilio Rojas for helpful discussion on the analyses. We apologize to our stick insect (Carausius morosus) for bearing with VAV’s inquisitive curiosity during our debates on color ornaments in king penguins. We sincerely thank the editor and 2 anonymous reviewers for their helpful comments on a previous version of the paper.Peer reviewedPostprin

The oxidative debt of fasting : evidence for short to medium-term costs of advanced fasting in adult king penguins

This research was funded by the French Polar Institute (IPEV–Research Program 119) and the French National Centre for Scientific Research (CNRS-INEE). We are especially grateful to Dominic L. Cram and one anonymous reviewer for helpful comments on the paper. Field logistic support was provided by Terres Australes et Antarctiques Françaises. QS was funded by a doctoral fellowship from the Ministère Français de l’Education Supérieur et de la Recherche.Peer reviewedPublisher PD

Effects of Individual Pre-Fledging Traits and Environmental Conditions on Return Patterns in Juvenile King Penguins

Despite the importance of early life stages in individuals' life history and population dynamics, very few studies have focused on the constraints to which these juvenile traits are subjected. Based on 10 years of automatic monitoring of over 2500 individuals, we present the first study on the effects of environmental conditions and individual pre-fledging traits on the post-fledging return of non-banded king penguins to their natal colony. Juvenile king penguins returned exclusively within one of the three austral summers following their departure. A key finding is that return rates (range 68–87%) were much higher than previously assumed for this species, importantly meaning that juvenile survival is very close to that of adults. Such high figures suggest little juvenile dispersal, and selection occurring mostly prior to fledging in king penguins. Pre-fledging conditions had a strong quadratic impact on juvenile return rates. As expected, cohorts reared under very unfavourable years (as inferred by the breeding success of the colony) exhibited low return rates but surprisingly, so did those fledged under very favourable conditions. Juvenile sojourns away from the colony were shorter under warm conditions and subsequent return rates higher, suggesting a positive effect of climate warming. The longer the post-fledging trip (1, 2 or 3 years), the earlier in the summer birds returned to their natal colony and the longer they stayed before leaving for the winter journey. The presence of juveniles in the colony was more than twice the duration required for moulting purposes, yet none attempted breeding in the year of their first return. Juvenile presence in the colony may be important for acquiring knowledge on the social and physical colonial environment and may play an important part in the learning process of mating behaviour. Further studies are required to investigate its potential implications on other life-history traits such as recruitment age

Experimental stress during molt suggests the evolution of condition-dependent and condition-independent ornaments in the king penguin

Acknowledgments We thank Emilie Lefol for her help in the field, and two anonymous reviewers for their comments on the manuscript. This research was funded by the French Polar Institute (IPEV– Research Program 119) and the French National Centre for Scientific Research (CNRS- INEE). Field logistic support was provided by Terres Australes et Antarctiques Françaises. Q.S. was funded by a doctoral fellowship from the Ministère Français de l’Education Supérieur et de la Recherche.Peer reviewedPublisher PD

Male reproductive tactics to increase paternity in the polygynandrous Columbian ground squirrel ( Urocitellus columbianus )

In polyandrous and polygynandrous species where females mate with multiple partners, males are expected to maximize their fitness by exhibiting an array of reproductive behaviors to ensure fertilization success, such as competing for the best mating order within a mating sequence, optimizing their investment in copulation, and mate guarding. Though there is genetic evidence of a first-male precedence in siring success for many mammalian species, the causes of this effect are poorly understood. We studied influences on first-male precedence in Columbian ground squirrels (Urocitellus columbianus). We found that the time a male spent consorting and mate guarding declined with his mating order (both the highest for the first male to mate). Mate guarding by the first male significantly reduced, but did not exclude, the number of additional males a female accepted. Later mating males reduced the time spent consorting, suggesting a perceived decreased chance of fertilization success. Consortship and mate guarding durations were positively related to the male's siring success and to each other, suggesting that males adjusted these behaviors strategically to increase their chances of fertilization success. Our results suggest that besides being the first male to consort, first-male sperm precedence is further enhanced through longer mating bouts and by suppressing the chances and/or efforts of later mating male

Oxidative stress and mitochondrial responses to stress exposure suggest that king penguins are naturally equipped to resist stress

We are grateful to four anonymous reviewers for their help in improving a previous draft of this manuscript and to the French Polar Institut (IPEV) for providing logistical support for this study through the programs 119 & 131. AS was self-funded during fieldwork, funded by the University of Angers during laboratory analyses, and was supported by a Marie Sklodowska-Curie Postdoctoral Fellowship (#658085) and a ‘Turku Collegium for Science and Medicine' Fellowship at the time of writing.Peer reviewedPublisher PD

Foster rather than biological parental telomere length predicts offspring survival and telomere length in king penguins

ACKNOWLEDGMENTS We are grateful to the field assistants who helped us with data collection in 2012-2013. This research was supported by the French Polar Research Institute (IPEV; program 119 ECONERGY), by the Centre National de la Recherche Scientifique (CNRS), by an International Emerging Action Grant (IEA n°203036) from the CNRS, and by the AXA Research Fund (post-doctoral fellowship to VA Viblanc). We are grateful to S Rogers and 5 anonymous reviewers for constructive and useful comments on previous drafts of the paper. DATA ACCESSIBILITY The data associated with this manuscript are available online at figshare doi: 10.6084/m9.figshare.12249902 (Viblanc et al. 2020).Peer reviewedPostprin

Foster rather than biological parental telomere length predicts offspring survival and telomere length in king penguins

Because telomere length and dynamics relate to individual growth, reproductive investment and survival, telomeres have emerged as possible markers of individual quality. Here, we tested the hypothesis that, in species with parental care, parental telomere length can be a marker of parental quality that predicts offspring phenotype and survival. In king penguins (Aptenodytes patagonicus), we experimentally swapped the single egg of 66 breeding pairs just after egg laying to disentangle the contribution of prelaying parental quality (e.g., genetics, investment in the egg) and/or postlaying parental quality (e.g., incubation, postnatal feeding rate) on offspring growth, telomere length and survival. Parental quality was estimated through the joint effects of biological and foster parent telomere length on offspring traits, both soon after hatching (day 10) and at the end of the prewinter growth period (day 105). We expected that offspring traits would be mostly related to the telomere lengths (i.e., quality) of biological parents at day 10 and to the telomere lengths of foster parents at day 105. Results show that chick survival up to 10 days was negatively related to biological fathers' telomere length, whereas survival up to 105 days was positively related to foster fathers' telomere lengths. Chick growth was not related to either biological or foster parents' telomere length. Chick telomere length was positively related to foster mothers' telomere length at both 10 and 105 days. Overall, our study shows that, in a species with biparental care, parents' telomere length is foremost a proxy of postlaying parental care quality, supporting the "telomere - parental quality hypothesis.

It Costs to Be Clean and Fit: Energetics of Comfort Behavior in Breeding-Fasting Penguins

), seabirds known to fast for up to one month during incubation shifts ashore.A time budget was estimated from focal and scan sampling field observations and the energy cost of comfort activities was calculated from the associated increase in heart rate (HR) during comfort episodes, using previously determined equations relating HR to energy expenditure. We show that incubating birds spent 22% of their daily time budget in comfort behavior (with no differences between day and night) mainly devoted to preening (73%) and head/body shaking (16%). During comfort behavior, energy expenditure averaged 1.24 times resting metabolic rate (RMR) and the corresponding energy cost (i.e., energy expended in excess to RMR) was 58 kJ/hr. Energy expenditure varied greatly among various types of comfort behavior, ranging from 1.03 (yawning) to 1.78 (stretching) times RMR. Comfort behavior contributed 8.8–9.3% to total daily energy expenditure and 69.4–73.5% to energy expended daily for activity. About half of this energy was expended caring for plumage.This study is the first to estimate the contribution of comfort behavior to overall energy budget in a free-living animal. It shows that although breeding on a tight energy budget, king penguins devote a substantial amount of time and energy to comfort behavior. Such findings underline the importance of comfort behavior for the fitness of colonial seabirds

Macrophysiology as a powerful tool for evaluating metapopulation stress and the effectiveness of conservation actions

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