Stephanie Jenouvrier

Wavelet analysis of ecological time series

Wavelet analysis is a powerful tool that is already in use throughout science and engineering. The versatility and attractiveness of the wavelet approach lie in its decomposition properties, principally its time-scale localization. It is especially relevant to the analysis of non-stationary systems, i.e., systems with short-lived transient components, like those observed in ecological systems. Here, we review the basic properties of the wavelet approach for time-series analysis from an ecological perspective. Wavelet decomposition offers several advantages that are discussed in this paper and illustrated by appropriate synthetic and ecological examples. Wavelet analysis is notably free from the assumption of stationarity that makes most methods unsuitable for many ecological time series. Wavelet analysis also permits analysis of the relationships between two signals, and it is especially appropriate for following gradual change in forcing by exogenous variables.

Demographic models and IPCC climate projections predict the decline of an emperor penguin population

Studies have reported important effects of recent climate change on Antarctic species, but there has been to our knowledge no attempt to explicitly link those results to forecasted population responses to climate change. Antarctic sea ice extent (SIE) is projected to shrink as concentrations of atmospheric greenhouse gases (GHGs) increase, and emperor penguins (Aptenodytes forsteri) are extremely sensitive to these changes because they use sea ice as a breeding, foraging and molting habitat. We project emperor penguin population responses to future sea ice changes, using a stochastic population model that combines a unique long-term demographic dataset (1962–2005) from a colony in Terre Adélie, Antarctica and projections of SIE from General Circulation Models (GCM) of Earths climate included in the most recent Intergovernmental Panel on Climate Change (IPCC) assessment report. We show that the increased frequency of warm events associated with projected decreases in SIE will reduce the population viability. The probability of quasi-extinction (a decline of 95% or more) is at least 36% by 2100. The median population size is projected to decline from ≈6,000 to ≈400 breeding pairs over this period. To avoid extinction, emperor penguins will have to adapt, migrate or change the timing of their growth stages. However, given the future projected increases in GHGs and its effect on Antarctic climate, evolution or migration seem unlikely for such long lived species at the remote southern end of the Earth.

LONG-TERM CONTRASTED RESPONSES TO CLIMATE OF TWO ANTARCTIC SEABIRD SPECIES

We examined the population dynamics of two Antarctic seabirds and the influence of environmental variability over a 40-year period by coupling the estimation of demographic parameters, based on capture-recapture data, and modeling, using Leslie ma- trix population models. We demonstrated that the demographic parameters showing the greatest contribution to the variance of population growth rate were adult survival for both species. Breeding success showed the same contribution as adult survival for Emperor Penguins, whereas the proportion of breeders had the next stronger contribution for Snow Petrels. The sensitivity of population growth rate to adult survival was very high and the adult survival variability was weak for both species. Snow Petrel males survived better than females, whereas Emperor Penguin males had lower survival than females. These differ- ences may be explained by the different investment in breeding. Emperor Penguin adult survival was negatively affected by air temperature during summer and winter for both sexes; male survival was negatively affected by sea ice concentration during summer, autumn, and winter. On the other hand, there was no effect of environmental covariates on Snow Petrel adult survival. The Emperor Penguin population has declined by 50% because of a decrease in adult survival related to a warming event during a regime shift in the late 1970s, whereas Snow Petrels showed their lowest numbers in 1976, but were able to skip reproduction. Indeed, the retrospective analysis of projection population matrix entries indicated that breeding abstention played a critical role in the population dynamics of Snow Petrels but not Emperor Penguins. Snow Petrels did not breed either when air temperature decreased during spring (probably reducing nest attendance and laying) or when sea ice decreased during autumn (reducing food availability). Emperor Penguin and Snow Petrel breeding population sizes were positively influenced by sea ice through its effect on adult survival for Emperor Penguins and on the proportion of breeders for Snow Petrels. Therefore, we hypothesize that the population sizes of the two species could be negatively affected by reduced sea ice in the context of global warming.

Phenological mismatch strongly affects individual fitness but not population demography in a woodland passerine

Populations are shifting their phenology in response to climate change, but these shifts are often asynchronous among interacting species. Resulting phenological mismatches can drive simultaneous changes in natural selection and population demography, but the links between these interacting processes are poorly understood. Here we analyse 37 years of data from an individual-based study of great tits (Parus major) in the Netherlands and use mixed-effects models to separate the within- and across-year effects of phenological mismatch between great tits and caterpillars (a key food source for developing nestlings) on components of fitness at the individual and population levels. Several components of individual fitness were affected by individual mismatch (i.e. late breeding relative to the caterpillar food peak date), including the probability of double-brooding, fledgling success, offspring recruitment probability and the number of recruits. Together these effects contributed to an overall negative relationship between relative fitness and laying dates, that is, selection for earlier laying on average. Directional selection for earlier laying was stronger in years where birds bred on average later than the food peak, but was weak or absent in years where the phenology of birds and caterpillars matched (i.e. no population mismatch). The mean number of fledglings per female was lower in years when population mismatch was high, in part because fewer second broods were produced. Population mismatch had a weak effect on the mean number of recruits per female, and no effect on mean adult survival, after controlling for the effects of breeding density and the quality of the autumnal beech (Fagus sylvatica) crop. These findings illustrate how climate change-induced mismatch can have strong effects on the relative fitness of phenotypes within years, but weak effects on mean demographic rates across years. We discuss various general mechanisms that influence the extent of coupling between breeding phenology, selection and population dynamics in open populations subject to strong density regulation and stochasticity.

Population growth in a wild bird Is buffered against phenological mismatch

Balancing Act Life cycles are strongly influenced by seasonal and interannual environmental and climate events. Such phenological timings are likely to shift as our climate changes, but species exist in communities, and not all species can be expected to shift in concert. Reed et al. (p. 488) used long-term data on European great tits to reveal how the negative consequences of phenological mismatch can be buffered: Lower fitness in individuals subject to mismatch-driven reductions of food availability was balanced by reductions in competition. Thus, overall, the population is resilient and has not declined. Warmer springtimes have induced great tits to lay their eggs earlier without affecting long-term fitness. Broad-scale environmental changes are altering patterns of natural selection in the wild, but few empirical studies have quantified the demographic cost of sustained directional selection in response to these changes. We tested whether population growth in a wild bird is negatively affected by climate change–induced phenological mismatch, using almost four decades of individual-level life-history data from a great tit population. In this population, warmer springs have generated a mismatch between the annual breeding time and the seasonal food peak, intensifying directional selection for earlier laying dates. Interannual variation in population mismatch has not, however, affected population growth. We demonstrated a mechanism contributing to this uncoupling, whereby fitness losses associated with mismatch are counteracted by fitness gains due to relaxed competition. These findings imply that natural populations may be able to tolerate considerable maladaptation driven by shifting climatic conditions without undergoing immediate declines.

Evidence of a shift in the cyclicity of Antarctic seabird dynamics linked to climate

Ecosystems and populations are known to be influenced not only by long-term climatic trends, but also by other short-term climatic modes, such as interannual and decadal-scale variabilities. Because interactions between climatic forcing, biotic and abiotic components of ecosystems are subtle and complex, analysis of long-term series of both biological and physical factors is essential to understanding these interactions. Here, we apply a wavelet analysis simultaneously to long-term datasets on the environment and on the populations and breeding success of three Antarctic seabirds (southern fulmar, snow petrel, emperor penguin) breeding in Terre Adélie, to study the effects of climate fluctuations on Antarctic marine ecosystems. We show that over the past 40 years, populations and demographic parameters of the three species fluctuate with a periodicity of 3–5 years that was also detected in sea-ice extent and the Southern Oscillation Index. Although the major periodicity of these interannual fluctuations is not common to different species and environmental variables, their cyclic characteristics reveal a significant change since 1980. Moreover, sliding-correlation analysis highlighted the relationships between environmental variables and the demography of the three species, with important change of correlation occurring between the end of the 1970s and the beginning of the 1980s. These results suggest that a regime shift has probably occurred during this period, significantly affecting the Antarctic ecosystem, but with contrasted effects on the three species.

Effects of climate change on an emperor penguin population: analysis of coupled demographic and climate models

Sea ice conditions in the Antarctic affect the life cycle of the emperor penguin (Aptenodytes forsteri). We present a population projection for the emperor penguin population of Terre Adélie, Antarctica, by linking demographic models (stage-structured, seasonal, nonlinear, two-sex matrix population models) to sea ice forecasts from an ensemble of IPCC climate models. Based on maximum likelihood capture-mark-recapture analysis, we find that seasonal sea ice concentration anomalies (SICa ) affect adult survival and breeding success. Demographic models show that both deterministic and stochastic population growth rates are maximized at intermediate values of annual SICa , because neither the complete absence of sea ice, nor heavy and persistent sea ice, would provide satisfactory conditions for the emperor penguin. We show that under some conditions the stochastic growth rate is positively affected by the variance in SICa . We identify an ensemble of five general circulation climate models whose output closely matches the historical record of sea ice concentration in Terre Adélie. The output of this ensemble is used to produce stochastic forecasts of SICa , which in turn drive the population model. Uncertainty is included by incorporating multiple climate models and by a parametric bootstrap procedure that includes parameter uncertainty due to both model selection and estimation error. The median of these simulations predicts a decline of the Terre Adélie emperor penguin population of 81% by the year 2100. We find a 43% chance of an even greater decline, of 90% or more. The uncertainty in population projections reflects large differences among climate models in their forecasts of future sea ice conditions. One such model predicts population increases over much of the century, but overall, the ensemble of models predicts that population declines are far more likely than population increases. We conclude that climate change is a significant risk for the emperor penguin. Our analytical approach, in which demographic models are linked to IPCC climate models, is powerful and generally applicable to other species and systems.

Mating Behavior, Population Growth, and the Operational Sex Ratio: A Periodic Two‐Sex Model Approach

We present a new approach to modeling two‐sex populations, using periodic, nonlinear two‐sex matrix models. The models project the population growth rate, the population structure, and any ratio of interest (e.g., operational sex ratio). The periodic formulation permits inclusion of highly seasonal behavioral events. A periodic product of the seasonal matrices describes annual population dynamics. The model is nonlinear because mating probability depends on the structure of the population. To study how the vital rates influence population growth rate, population structure, and operational sex ratio, we used sensitivity analysis of frequency‐dependent nonlinear models. In nonlinear two‐sex models the vital rates affect growth rate directly and also indirectly through effects on the population structure. The indirect effects can sometimes overwhelm the direct effects and are revealed only by nonlinear analysis. We find that the sensitivity of the population growth rate to female survival is negative for the emperor penguin, a species with highly seasonal breeding behavior. This result could not occur in linear models because changes in population structure have no effect on per capita reproduction. Our approach is applicable to ecological and evolutionary studies of any species in which males and females interact in a seasonal environment.

Buying Years to Extinction: Is Compensatory Mitigation for Marine Bycatch a Sufficient Conservation Measure for Long-Lived Seabirds?

Along the lines of the ‘polluter pays principle’, it has recently been proposed that the local long-line fishing industry should fund eradication of terrestrial predators at seabird breeding colonies, as a compensatory measure for the bycatch caused by the fishing activity. The measure is economically sound, but a quantitative and reliable test of its biological efficacy has never been conducted. Here, we investigated the demographic consequences of predator eradication for Corys shearwater Calonectris diomedea, breeding in the Mediterranean, using a population model that integrates demographic rates estimated from individual life-history information with experimental measures of predation and habitat structure. We found that similar values of population growth rate can be obtained by different combinations of habitat characteristics, predator abundance and adult mortality, which explains the persistence of shearwater colonies in islands with introduced predators. Even so, given the empirically obtained values of survival, all combinations of predator abundance and habitat characteristics projected a decline in shearwater numbers. Perturbation analyses indicated that the value and the sensitivity of shearwater population growth rates were affected by all covariates considered and their interactions. A decrease in rat abundance delivered only a small increase in the population growth rate, whereas a change in adult survival (a parameter independent of rat abundance) had the strongest impact on population dynamics. When adult survival is low, rat eradication would allow us to “buy” years before extinction but does not reverse the process. Rat eradication can therefore be seen as an emergency measure if threats on adult survival are eliminated in the medium-term period. For species with low fecundity and long life expectancy, our results suggest that rat control campaigns are not a sufficient, self-standing measure to compensate the biological toll of long-line fisheries.

Avian cholera, a threat to the viability of an Arctic seabird colony?

Evidence that infectious diseases cause wildlife population extirpation or extinction remains anecdotal and it is unclear whether the impacts of a pathogen at the individual level can scale up to population level so drastically. Here, we quantify the response of a Common eider colony to emerging epidemics of avian cholera, one of the most important infectious diseases affecting wild waterfowl. We show that avian cholera has the potential to drive colony extinction, even over a very short period. Extinction depends on disease severity (the impact of the disease on adult female survival) and disease frequency (the number of annual epidemics per decade). In case of epidemics of high severity (i.e., causing >30% mortality of breeding females), more than one outbreak per decade will be unsustainable for the colony and will likely lead to extinction within the next century; more than four outbreaks per decade will drive extinction to within 20 years. Such severity and frequency of avian cholera are already observed, and avian cholera might thus represent a significant threat to viability of breeding populations. However, this will depend on the mechanisms underlying avian cholera transmission, maintenance, and spread, which are currently only poorly known.