Matthieu Authier

Evidence for an age-dependent influence of environmental variations on a long-lived seabird's life-history traits.

Theoretical and empirical studies have highlighted the effects of age on several life-history traits in wild populations. There is also increasing evidence for environmental effects on their demographic traits. However, quantifying how individuals differentially respond to environmental variations according to their age remains a challenge in ecology. In a population of Black-browed Albatrosses monitored during 43 years, we analyzed how life-history traits varied according to age, and whether individuals of different ages responded in different ways to environmental conditions. To do so, we: (1) examined how age affected seven life-history traits, (2) investigated differences in temporal variance of demographic traits between age classes, and (3) tested for age-dependent effects of climate and fisheries covariates on demographic traits. Overall, there was a tendency for traits to improve during the first years of life (5-10 years), to peak and remain stable at middle age (10-30 years), and decline at old ages. At young ages, survival and reproductive parameters increased, except offspring body condition at fledging, suggesting that younger parents had already acquired good foraging capacities. However, they suffered from inexperience in breeding as suggested by their higher breeding failures during incubation. There was evidence for reproductive and actuarial senescence. In particular, breeding success and offspring body condition declined abruptly, suggesting altered foraging capacities of old individuals. Middle-aged individuals had the lowest temporal variance of demographic traits. Although this is predicted by the theory of environmental canalization, it could also results from a higher susceptibility of young and old birds due to their respective inexperience and senescence. The highest temporal variances were found in old individuals. Survival was significantly influenced by sea surface temperatures in the foraging zone of this albatross population during breeding. During warm events survival of young and old individuals improved, whereas a decrease was observed for middle-aged individuals. Presumably, during cold years with poor environmental conditions, young and old breeding birds may suffer more from intraspecific competition for resources than middle-aged individuals. This study showed that age, known as a major factor structuring demography in long-lived species, can also potentially influence the response of populations to global change.

Windscape and tortuosity shape the flight costs of northern gannets

When animals move across a landscape, they alternate between active searching phases in areas with high prey density and commuting phases towards and in-between profitable feeding patches. Such active searching movements are more sinuous than travelling movements, and supposedly more costly in energy. Here we provide an empirical validation of this long-lasting assumption. To this end, we evaluated simultaneously energy expenditure and trajectory in northern gannets (Morus bassanus) using GPS loggers, dive recorders and three-dimensional accelerometers. Three behavioural states were determined from GPS data: foraging, when birds actively searched for prey (high tortuosity, medium speed); travelling, when birds were commuting (straight trajectory, high speed); and resting (straight trajectory, low speed). Overall dynamic body acceleration, calculated from acceleration data, was used as a proxy for energy expenditure during flight. The impact of windscape characteristics (wind force and direction) upon flight costs was also tested. Energy expenditure of northern gannets was higher during sinuous foraging flight than during more rectilinear travelling flight, demonstrating that turns are indeed costly. Yet wind force and direction also strongly shaped flight energy expenditure; within any behavioural state it was less costly to fly with the wind than against it, and less costly to fly with strong winds. Despite the major flight costs of wind action, birds did not fully optimize their flight track relative to wind direction, probably because of prey distributions relative to the coastline and wind predictability. Our study illustrates how both tortuosity and windscape shape the foraging costs of marine predators such as northern gannets.

Antarctic Climate Change: Extreme Events Disrupt Plastic Phenotypic Response in Adélie Penguins

In the context of predicted alteration of sea ice cover and increased frequency of extreme events, it is especially timely to investigate plasticity within Antarctic species responding to a key environmental aspect of their ecology: sea ice variability. Using 13 years of longitudinal data, we investigated the effect of sea ice concentration (SIC) on the foraging efficiency of Adélie penguins (Pygoscelis adeliae) breeding in the Ross Sea. A ‘natural experiment’ brought by the exceptional presence of giant icebergs during 5 consecutive years provided unprecedented habitat variation for testing the effects of extreme events on the relationship between SIC and foraging efficiency in this sea-ice dependent species. Significant levels of phenotypic plasticity were evident in response to changes in SIC in normal environmental conditions. Maximum foraging efficiency occurred at relatively low SIC, peaking at 6.1% and decreasing with higher SIC. The ‘natural experiment’ uncoupled efficiency levels from SIC variations. Our study suggests that lower summer SIC than currently observed would benefit the foraging performance of Adélie penguins in their southernmost breeding area. Importantly, it also provides evidence that extreme climatic events can disrupt response plasticity in a wild seabird population. This questions the predictive power of relationships built on past observations, when not only the average climatic conditions are changing but the frequency of extreme climatic anomalies is also on the rise.

O' mother where wert thou? Maternal strategies in the southern elephant seal: a stable isotope investigation

Maternal effects are widespread in ecology and can alter the dynamics of a population. We investigated the impact of maternal foraging strategies on offspring weaning mass—a proxy of maternal foraging success and of offspring survival—in southern elephant seals on îles Kerguelen. Using 4 years of data, we modelled pup weaning mass as a two-component mixture and used blood stable isotope values to discriminate between maternal foraging strategies previously identified from bio-logging studies. Carbon isotope ratio was a strong predictor of weaning mass, but the relationship was non-monotonic in contrast to a priori expectations. Females foraging in the interfrontal zone weaned pups with a smaller mass compared with females foraging in Antarctic waters. Pup mass was positively correlated with a proxy of global primary production in the interfrontal zone for small weanlings. Maternal effects, via a poor foraging efficiency in the 1970s, may help explain the large population decrease observed at that time on îles Kerguelen because of an overall decrease in pup weaning mass, survival and subsequent recruitment.

Foraging fidelity as a recipe for a long life: foraging strategy and longevity in male Southern Elephant Seals.

Identifying individual factors affecting life-span has long been of interest for biologists and demographers: how do some individuals manage to dodge the forces of mortality when the vast majority does not? Answering this question is not straightforward, partly because of the arduous task of accurately estimating longevity in wild animals, and of the statistical difficulties in correlating time-varying ecological covariables with a single number (time-to-event). Here we investigated the relationship between foraging strategy and life-span in an elusive and large marine predator: the Southern Elephant Seal (Mirounga leonina). Using teeth recovered from dead males on îles Kerguelen, Southern Ocean, we first aged specimens. Then we used stable isotopic measurements of carbon () in dentin to study the effect of foraging location on individual life-span. Using a joint change-point/survival modelling approach which enabled us to describe the ontogenetic trajectory of foraging, we unveiled how a stable foraging strategy developed early in life positively covaried with longevity in male Southern Elephant Seals. Coupled with an appropriate statistical analysis, stable isotopes have the potential to tackle ecological questions of long standing interest but whose answer has been hampered by logistic constraints.

Designing observational biologging studies to assess the causal effect of instrumentation

1. Biologging has improved ecological knowledge on an increasing number of species for more than 2 decades. Most studies looking at the incidence of tags on behavioural, physiological or demographic parameters rely on ‘control’ individuals chosen randomly within the population, assuming that they will be comparable with equipped individuals. This assumption is usually untestable and untenable since biologging studies are more observational than experimental, and often involve small sample sizes. Notably, background characteristics of wild animals are, most of the time, unknown. Consequently, investigating any causal effect of instrumentation is a difficult task, subjected to hidden biases. 2. We describe the counterfactual model to causal inference which was implicit in early biologging studies. We adopted methods developed in social and political sciences to construct a posteriori an appropriate control group. Using biologging data collected on Scopoli’s shearwaters ( Calonectris diomedea ) from a small Mediterranean island, we used this method to achieve objective causal inference on the effect of instrumentation on breeding performance and divorce. 3. Our method revealed that the sample of instrumented birds was nonrandom. After identification of a relevant control group, we found no carry-over effects of instrumentation on breeding performance (taking into account imperfect detection probability) or divorce rate in Scopoli’s shearwaters. 4. Randomly chosen control groups can be both counterproductive and ethically dubious via unnecessary additional disturbance of populations. The counterfactual approach, which can correct for selection bias, has wide applicability to biologging within long-term studies.

Variable selection and accurate predictions in habitat modelling: a shrinkage approach

Habitat modelling is increasingly relevant in biodiversity and conservation studies. A typical application is to predict potential zones of specific conservation interest. With many environmental covariates, a large number of models can be investigated but multi-model inference may become impractical. Shrinkage regression overcomes this issue by dealing with the identification and accurate estimation of effect size for prediction. In a Bayesian framework we investigated the use of a shrinkage prior, the Horseshoe, for variable selection in spatial generalized linear models (GLM). As study cases, we considered 5 datasets on small pelagic fish abundance in the Gulf of Lion (Mediterranean Sea, France) and 9 environmental inputs. We compared the predictive performances of a simple kriging model, a full spatial GLM model with independent normal priors for regression coefficients, a full spatial GLM model with a Horseshoe prior for regression coefficients and 2 zero-inflated models (spatial and non-spatial) with a Horseshoe prior. Predictive performances were evaluated by cross-validation on a hold-out subset of the data: models with a Horseshoe prior performed best, and the full model with independent normal priors worst. With an increasing number of inputs, extrapolation quickly became pervasive as we tried to predict from novel combinations of covariate values. By shrinking regression coefficients with a Horseshoe prior, only one model needed to be fitted to the data in order to obtain reasonable and accurate predictions, including extrapolations.

How much are stranding records affected by variation in reporting rates? A case study of small delphinids in the Bay of Biscay

Marine vertebrate strandings offer an opportunistic sampling scheme that can provide abundant data over long periods. Because the stranding process involves biological, physical and sociological parameters, confounding complicates the interpretation of results. The statistical analysis of these data relies on generalized linear or additive models in order to infer long-term trends, but does not easily account for drift or variation in reporting rates. Here, we capitalized on county-level (administrative) variation following the passing of a law for compulsory reporting of stranded marine mammals in France to investigate how variation in reporting rates may affect the observed trend in stranded small delphinids in the Bay of Biscay. Using a time-series spanning more than 30 years across eight administrative counties, we built variance partitioning models for the analysis of count data. We discussed the choice of an appropriate likelihood to conclude the Negative Binomial useful and interpretable in the context of small delphinid strandings. We expanded the model with a recent methodology to detect structural breaks in the time series, focusing on overdispersion. We performed statistical robustness checks with respect to variations in reporting rates and discuss their causal interpretation in the context of observational data. Stranding frequencies increased on average 7-fold over 30 years. We conclude that reporting rates to the French stranding network have been stable since the early 1990s, and the average 3-fold increase in stranded small delphinids observed in the Bay of Biscay since 1990 is due to other factors, including bycatch. Codes and data are available to replicate the analysis to other national stranding networks.

Wolf in sheep’s clothing: Model misspecification undermines tests of the neutral theory for life histories

Abstract Understanding the processes behind change in reproductive state along life‐history trajectories is a salient research program in evolutionary ecology. Two processes, state dependence and heterogeneity, can drive the dynamics of change among states. Both processes can operate simultaneously, begging the difficult question of how to tease them apart in practice. The Neutral Theory for Life Histories (NTLH) holds that the bulk of variations in life‐history trajectories is due to state dependence and is hence neutral: Once previous (breeding) state is taken into account, variations are mostly random. Lifetime reproductive success (LRS), the number of descendants produced over an individuals reproductive life span, has been used to infer support for NTLH in natura. Support stemmed from accurate prediction of the population‐level distribution of LRS with parameters estimated from a state dependence model. We show with Monte Carlo simulations that the current reliance of NTLH on LRS prediction in a null hypothesis framework easily leads to selecting a misspecified model, biased estimates and flawed inferences. Support for the NTLH can be spurious because of a systematic positive bias in estimated state dependence when heterogeneity is present in the data but ignored in the analysis. This bias can lead to spurious positive covariance between fitness components when there is in fact an underlying trade‐off. Furthermore, neutrality implied by NTLH needs a clarification because of a probable disjunction between its common understanding by evolutionary ecologists and its translation into statistical models of life‐history trajectories. Irrespective of what neutrality entails, testing hypotheses about the dynamics of change among states in life histories requires a multimodel framework because state dependence and heterogeneity can easily be mistaken for each other.

How large is large: estimating ecologically meaningful isotopic differences in observational studies of wild animals

RATIONALE In ecological studies of wildlife movements and foraging, bio-logging and isotopic data are routinely collected and increasingly analyzed in tandem. Such analyses have two shortcomings: (1) small sample size linked with the number of telemetric tags that can be deployed, and (2) the observational nature of isotopic gradients. Wildlife ecologists are thus put in a statistical conundrum known as the small n, large p problem. METHODS Using shrinkage regression, which directly addresses the issue of accurately estimating effects from sparse data, we studied what counts as a biologically meaningful isotopic difference (a prerequisite to delineate isoscapes) in the southern elephant seal (Mirounga leonina), a large and elusive marine predator. RESULTS Seals foraging in Antarctic waters had a lower carbon isotopic value (by ≈ 2‰) than seals foraging either in the interfrontal zone or on the Kerguelen Plateau. The latter two foraging strategies were indistinguishable on the sole basis of δ(13) C values with our data. CONCLUSIONS Shrinkage regression is a conservative statistical technique that has wide applicability in isotopic ecology to help separate robust biological signals from noise.