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      Early diving behaviour in juvenile penguins: improvement or selection processes

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          Abstract

          The early life stage of long-lived species is critical to the viability of population, but is poorly understood. Longitudinal studies are needed to test whether juveniles are less efficient foragers than adults as has been hypothesized. We measured changes in the diving behaviour of 17 one-year-old king penguins Aptenodytes patagonicus at Crozet Islands (subantartic archipelago) during their first months at sea, using miniaturized tags that transmitted diving activity in real time. We also equipped five non-breeder adults with the same tags for comparison. The data on foraging performance revealed two groups of juveniles. The first group made shallower and shorter dives that may be indicative of early mortality while the second group progressively increased their diving depths and durations, and survived the first months at sea. This surviving group of juveniles required the same recovery durations as adults, but typically performed shallower and shorter dives. There is thereby a relationship between improved diving behaviour and survival in young penguins. This long period of improving diving performance in the juvenile life stage is potentially a critical period for the survival of deep avian divers and may have implications for their ability to adapt to environmental change.

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          How life history influences population dynamics in fluctuating environments.

          A major question in ecology is how age-specific variation in demographic parameters influences population dynamics. Based on long-term studies of growing populations of birds and mammals, we analyze population dynamics by using fluctuations in the total reproductive value of the population. This enables us to account for random fluctuations in age distribution. The influence of demographic and environmental stochasticity on the population dynamics of a species decreased with generation time. Variation in age-specific contributions to total reproductive value and to stochastic components of population dynamics was correlated with the position of the species along the slow-fast continuum of life-history variation. Younger age classes relative to the generation time accounted for larger contributions to the total reproductive value and to demographic stochasticity in "slow" than in "fast" species, in which many age classes contributed more equally. In contrast, fluctuations in population growth rate attributable to stochastic environmental variation involved a larger proportion of all age classes independent of life history. Thus, changes in population growth rates can be surprisingly well explained by basic species-specific life-history characteristics.
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            From cradle to early grave: juvenile mortality in European shags Phalacrocorax aristotelis results from inadequate development of foraging proficiency.

            In most long-lived animal species, juveniles survive less well than adults. A potential mechanism is inferior foraging skills but longitudinal studies that follow the development of juvenile foraging are needed to test this. We used miniaturized activity loggers to record daily foraging times of juvenile and adult European shags Phalacrocorax aristotelis from fledging to the following spring. Juveniles became independent from their parents 40 days post-fledging. They compensated for poor foraging proficiency by foraging for approximately 3 h d(-1) longer than adults until constrained by day length in early November. Thereafter, juvenile foraging time tracked shortening day length up to the winter solstice, when foraging time of the two age classes converged and continued to track day length until early February. Few individuals died until midwinter and mortality peaked in January-February, with juvenile mortality (including some of the study birds) five times that of adults. In their last two weeks of life, juveniles showed a marked decline in foraging time consistent with individuals becoming moribund. Our results provide compelling evidence that juveniles compensate for poor foraging proficiency by increasing foraging time, a strategy that is limited by day length resulting in high winter mortality.
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              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.
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                Author and article information

                Journal
                Biol Lett
                Biol. Lett
                RSBL
                roybiolett
                Biology Letters
                The Royal Society
                1744-9561
                1744-957X
                August 2016
                August 2016
                : 12
                : 8
                : 20160490
                Affiliations
                Centre d'Etudes Biologiques de Chize UMR 7372 du CNRS-Universite de LaRochelle, 79360 Villiers-en-Bois, France
                Author notes
                Author information
                http://orcid.org/0000-0002-1940-7797
                Article
                rsbl20160490
                10.1098/rsbl.2016.0490
                5014042
                27484650
                2764ca51-86e3-43c1-8b4a-69541621ea97
                © 2016 The Authors.

                Published by the Royal Society under the terms of the Creative Commons Attribution License http://creativecommons.org/licenses/by/4.0/, which permits unrestricted use, provided the original author and source are credited.

                History
                : 8 June 2016
                : 8 July 2016
                Funding
                Funded by: European Community's Seven Framework;
                Award ID: ERC-2012-ADG_20120314
                Categories
                1001
                14
                60
                Animal Behaviour
                Custom metadata
                August, 2016

                Life sciences
                juveniles,ontogeny,diving behaviour,penguins,bio-logging,tracking
                Life sciences
                juveniles, ontogeny, diving behaviour, penguins, bio-logging, tracking

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