In 2007, the Intergovernmental Panel on Climate Change highlighted an urgent need to assess the responses of marine ecosystems to climate change. Because they lie in a high-latitude region, the Southern Ocean ecosystems are expected to be strongly affected by global warming. Using top predators of this highly productive ocean (such as penguins) as integrative indicators may help us assess the impacts of climate change on marine ecosystems. Yet most available information on penguin population dynamics is based on the controversial use of flipper banding. Although some reports have found the effects of flipper bands to be deleterious, some short-term (one-year) studies have concluded otherwise, resulting in the continuation of extensive banding schemes and the use of data sets thus collected to predict climate impact on natural populations. Here we show that banding of free-ranging king penguins (Aptenodytes patagonicus) impairs both survival and reproduction, ultimately affecting population growth rate. Over the course of a 10-year longitudinal study, banded birds produced 41% [corrected] fewer chicks and had a survival rate 16 percentage points [corrected] lower than non-banded birds, demonstrating a massive long-term impact of banding and thus refuting the assumption that birds will ultimately adapt to being banded. Indeed, banded birds still arrived later for breeding at the study site and had longer foraging trips even after 10 years. One of our major findings is that responses of flipper-banded penguins to climate variability (that is, changes in sea surface temperature and in the Southern Oscillation index) differ from those of non-banded birds. We show that only long-term investigations may allow an evaluation of the impact of flipper bands and that every major life-history trait can be affected, calling into question the banding schemes still going on. In addition, our understanding of the effects of climate change on marine ecosystems based on flipper-band data should be reconsidered.

Since 2007, the ecosystem of the Gulf of Lions has shifted to a different regime, characterised by a low anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus) biomass and a remarkably high sprat (Sprattus sprattus) biomass. Surprisingly, the abundance and recruitment of anchovy and sardine remained high. To understand which processes (bottom-up or top-down control, etc.) could have caused this shift, we studied the changes in body condition, growth and size and age of anchovy, sardine and sprat over 1984-1985 and 1992-2012, using data from scientific surveys. The annual age structure of anchovy and sardine was estimated using Bayesian mixture models based on size frequency data with priors on the age-length relationship derived from independent otolith readings. The results indicated periods during which anchovy and sardine were in an average (1992-2004), good (2005-2007) or poor (2008-2012) overall state of condition. For sardine, the shift towards smaller fish observed during these past 4 years was explained by a combination of slower growth and the disappearance of older individuals (ages 2+). Despite the increase in biomass of sprat since 2008, indications were found that sprat was also smaller than in the past. As growth and condition decreased and overexploitation has not been documented or suspected for those three species in this area, we propose that the current decline in sardine and anchovy biomass could be due to qualitative and/or quantitative modifications in the planktonic production (i.e. a bottom-up control) or mass mortalities of adults due to an epidemic disease.

Around 2008, an ecosystem shift occurred in the Gulf of Lions, highlighted by considerable changes in biomass and fish mean weight of its two main small pelagic fish stocks (European anchovy, Engraulis encrasicolus; European sardine, Sardina pilchardus). Surprisingly these changes did not appear to be mediated by a decrease in fish recruitment rates (which remained high) or by a high fishing pressure (exploitation rates being extremely low). Here, we review the current knowledge on the population's dynamics and its potential causes. We used an integrative ecosystem approach exploring alternative hypotheses, ranging from bottom-up to top-down control, not forgetting epizootic diseases. First, the study of multiple population characteristics highlighted a decrease in body condition for both species as well as an important decrease in size resulting both from a slower growth and a progressive disappearance of older sardines. Interestingly, older sardines were more affected by the decrease in condition than younger ones, another sign of an unbalanced population structure. While top-down control by bluefin tuna or dolphins, emigration and disease were mostly discarded as important drivers, bottom-up control mediated by potential changes in the plankton community appeared to play an important role via a decrease in fish energy income and hence growth, condition and size. Isotopic and stomach content analyses indicated a dietary shift pre-and post-2008 and modeled mesozooplankton abundance was directly linked to fish condition. Despite low energy reserves from 2008 onwards, sardines and anchovies maintained if not increased their reproductive investment, likely altering the lifehistory trade-off between reproduction and survival and resulting in higher natural mortality. The current worrying situation might thus have resulted from changes in plankton availability/diversity, which remains to be thoroughly investigated together with fish phenotypic plasticity.

Endogenous and environmental variables are fundamental in explaining variations in fish condition. Based on more than 20 yr of fish weight and length data, relative condition indices were computed for anchovy and sardine caught in the Gulf of Lions. Classification and regression trees (CART) were used to identify endogenous factors affecting fish condition, and to group years of similar condition. Both species showed a similar annual cycle with condition being minimal in February and maximal in July. CART identified 3 groups of years where the fish populations generally showed poor, average and good condition and within which condition differed between age classes but not according to sex. In particular, during the period of poor condition (mostly recent years), sardines older than 1 yr appeared to be more strongly affected than younger individuals. Time-series were analyzed using generalized linear models (GLMs) to examine the effects of oceanographic abiotic (temperature, Western Mediterranean Oscillation [WeMO] and Rhône outflow) and biotic (chlorophyll a and 6 plankton classes) factors on fish condition. The selected models explained 48 and 35% of the variance of anchovy and sardine condition, respectively. Sardine condition was negatively related to temperature but positively related to the WeMO and mesozooplankton and diatom concentrations. A positive effect of mesozooplankton and Rhône runoff on anchovy condition was detected. The importance of increasing temperatures and reduced water mixing in the NW Mediterranean Sea, affecting planktonic productivity and thus fish condition by bottom-up control processes, was highlighted by these results. Changes in plankton quality, quantity and phenology could lead to insufficient or inadequate food supply for both species.

Understanding the ecological and anthropogenic drivers of population dynamics requires detailed studies on habitat selection and spatial distribution. Although small pelagic fish aggregate in large shoals and usually exhibit important spatial structure, their dynamics in time and space remain unpredictable and challenging. In the Gulf of Lions (north-western Mediterranean), sardine and anchovy biomasses have declined over the past 5 years causing an important fishery crisis while sprat abundance rose. Applying geostatistical tools on scientific acoustic surveys conducted in the Gulf of Lions, we investigated anchovy, sardine and sprat spatial distributions and structures over 10 years. Our results show that sardines and sprats were more coastal than anchovies. The spatial structure of the three species was fairly stable over time according to variogram outputs, while year-to-year variations in kriged maps highlighted substantial changes in their location. Support for the McCall's basin hypothesis (covariation of both population density and presence area with biomass) was found only in sprats, the most variable of the three species. An innovative method to investigate species collocation at different scales revealed that globally the three species strongly overlap. Although species often co-occurred in terms of presence/absence, their biomass density differed at local scale, suggesting potential interspecific avoidance or different sensitivity to local environmental characteristics. Persistent favourable areas were finally detected, but their environmental characteristics remain to be determined.

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Abstract. Breeding animals face important time and energy constraints when caring for themselves and their offspring. For long-lived species, life-history theory predicts that parents should favor survival over current reproductive attempts, thus investing more into their own maintenance than the provisioning of their young. In seabirds, provisioning strategies may additionally be influenced by the distance between breeding sites and foraging areas, and offshore and inshore species should thus exhibit different strategies. Here, we examine the provisioning strategies of an inshore seabird using a long-term data set on more than 200 Little Penguins, Eudyptula minor. They alternated between two consecutive long and several short foraging trips all along chick rearing, a strategy almost never observed for inshore animals. Short trips allowed for regular provisioning of the chicks (high feeding frequency and larger meals), whereas long trips were performed when parent body mass was low and enabled them to rebuild their reserves, suggesting that adult body condition may be a key factor in initiating long trips. Inshore seabirds do use dual strategies of alternating short and long trips, but from our data, on a simpler and less flexible way than for offshore birds.

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