Joël M. Durant

Ecosystem oceanography for global change in fisheries

Overexploitation and climate change are increasingly causing unanticipated changes in marine ecosystems, such as higher variability in fish recruitment and shifts in species dominance. An ecosystem-based approach to fisheries attempts to address these effects by integrating populations, food webs and fish habitats at different scales. Ecosystem models represent indispensable tools to achieve this objective. However, a balanced research strategy is needed to avoid overly complex models. Ecosystem oceanography represents such a balanced strategy that relates ecosystem components and their interactions to climate change and exploitation. It aims at developing realistic and robust models at different levels of organisation and addressing specific questions in a global change context while systematically exploring the ever-increasing amount of biological and environmental data.

Reliability of flipper-banded penguins as indicators of climate change

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 39% fewer chicks and had a survival rate 16% 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.

Phenological changes in the southern hemisphere.

Current evidence of phenological responses to recent climate change is substantially biased towards northern hemisphere temperate regions. Given regional differences in climate change, shifts in phenology will not be uniform across the globe, and conclusions drawn from temperate systems in the northern hemisphere might not be applicable to other regions on the planet. We conduct the largest meta-analysis to date of phenological drivers and trends among southern hemisphere species, assessing 1208 long-term datasets from 89 studies on 347 species. Data were mostly from Australasia (Australia and New Zealand), South America and the Antarctic/subantarctic, and focused primarily on plants and birds. This meta-analysis shows an advance in the timing of spring events (with a strong Australian data bias), although substantial differences in trends were apparent among taxonomic groups and regions. When only statistically significant trends were considered, 82% of terrestrial datasets and 42% of marine datasets demonstrated an advance in phenology. Temperature was most frequently identified as the primary driver of phenological changes; however, in many studies it was the only climate variable considered. When precipitation was examined, it often played a key role but, in contrast with temperature, the direction of phenological shifts in response to precipitation variation was difficult to predict a priori. We discuss how phenological information can inform the adaptive capacity of species, their resilience, and constraints on autonomous adaptation. We also highlight serious weaknesses in past and current data collection and analyses at large regional scales (with very few studies in the tropics or from Africa) and dramatic taxonomic biases. If accurate predictions regarding the general effects of climate change on the biology of organisms are to be made, data collection policies focussing on targeting data-deficient regions and taxa need to be financially and logistically supported.

King penguin population threatened by Southern Ocean warming

Seabirds are sensitive indicators of changes in marine ecosystems and might integrate and/or amplify the effects of climate forcing on lower levels in food chains. Current knowledge on the impact of climate changes on penguins is primarily based on Antarctic birds identified by using flipper bands. Although flipper bands have helped to answer many questions about penguin biology, they were shown in some penguin species to have a detrimental effect. Here, we present for a Subantarctic species, king penguin (Aptenodytes patagonicus), reliable results on the effect of climate on survival and breeding based on unbanded birds but instead marked by subcutaneous electronic tags. We show that warm events negatively affect both breeding success and adult survival of this seabird. However, the observed effect is complex because it affects penguins at several spatio/temporal levels. Breeding reveals an immediate response to forcing during warm phases of El Niño Southern Oscillation affecting food availability close to the colony. Conversely, adult survival decreases with a remote sea-surface temperature forcing (i.e., a 2-year lag warming taking place at the northern boundary of pack ice, their winter foraging place). We suggest that this time lag may be explained by the delay between the recruitment and abundance of their prey, adjusted to the particular 1-year breeding cycle of the king penguin. The derived population dynamic model suggests a 9% decline in adult survival for a 0.26°C warming. Our findings suggest that king penguin populations are at heavy extinction risk under the current global warming predictions.

Collapse of South Africa's penguins in the early 21st century

The number of African penguins Spheniscus demersus breeding in South Africa collapsed from about 56 000 pairs in 2001 to some 21 000 pairs in 2009, a loss of 35 000 pairs (>60%) in eight years. This reduced the global population to 26 000 pairs, when including Namibian breeders, and led to classification of the species as Endangered. In South Africa, penguins breed in two regions, the Western Cape and Algoa Bay (Eastern Cape), their breeding localities in these regions being separated by c. 600 km. Their main food is anchovy Engraulis encrasicolus and sardine Sardinops sagax, which are also the target of purse-seine fisheries. In Algoa Bay, numbers of African penguins halved from 21 000 pairs in 2001 to 10 000 pairs in 2003. In the Western Cape, numbers decreased from a mean of 35 000 pairs in 2001–2005 to 11 000 pairs in 2009. At Dassen Island, the annual survival rate of adult penguins decreased from 0.70 in 2002/2003 to 0.46 in 2006/2007; at Robben Island it decreased from 0.77 to 0.55 in the same period. In both the Western and Eastern Cape provinces, long-term trends in numbers of penguins breeding were significantly related to the combined biomass of anchovy and sardine off South Africa. However, recent decreases in the Western Cape were greater than expected given a continuing high abundance of anchovy. In this province, there was a south-east displacement of prey around 2000, which led to a mismatch in the distributions of prey and the western breeding localities of penguins.

Ocean climate prior to breeding affects the duration of the nestling period in the Atlantic puffin

Time-series covering 23 years for a long-lived seabird, the Atlantic puffin (Fratercula arctica L.) at Røst, northern Norway, was used to explore any indirect effects of climatic variations on chick production. By fitting statistical models on the duration of the nestling period, we found that it may be estimated using the average sea temperature and salinity at 0–20 m depth in March (having a positive and a negative effect, respectively). We propose that when the phytoplankton bloom occurs in early spring, adverse oceanographic conditions, i.e. low temperature and high salinity in March, have a negative effect on puffin reproduction by degradation of the prey availability (mainly Clupea harengus) for chick-feeding adults three months later.

Body mass and clutch size may modulate prolactin and corticosterone levels in eiders.

Altered body condition, increased incubation costs, and egg loss are important proximate factors modulating bird parental behavior, since they inform the adult about its remaining chances of survival or about the expected current reproductive success. Hormonal changes should reflect internal or external stimuli, since corticosterone levels (inducing nest abandonment) are known to increase while body condition deteriorates, and prolactin levels (stimulating incubation) decrease following egg predation. However, in a capital incubator that based its investment on available body reserves and naturally lost about half of its body mass during incubation, corticosterone should be maintained at a low threshold to avoid protein mobilization for energy supply. This study focused on the regulation of corticosterone and prolactin release in such birds during incubation, when facing egg manipulation (control, reduced, or increased) or a stressful event. Blood samples were taken before and after clutch manipulation and at hatching. Corticosterone levels were determined before and after 30 min of captivity. Female eiders exhibited a high hypothalamic‐pituitary‐adrenal sensitivity, plasma concentration of corticosterone being increased by four‐ to fivefold following 30 min of captivity. The adrenocortical response was not modified by body mass loss but was higher in birds for which clutch size was increased. In the same way, females did not show different prolactin levels among the experimental groups. However, when incubation started, prolactin levels were correlated to body mass, suggesting that nest attendance is programmed in relation to the female initial body condition. Moreover, due to an artifactual impact of bird manipulation, increased baseline corticosterone was associated with a prolactin decrease in the control group. These data suggest that, in eiders, body mass and clutch size modification can modulate prolactin and corticosterone levels, which cross‐regulate each other in order to finely control incubation behavior.

Body reserves and nutritional needs during laying preparation in barn owls

Abstract To investigate the composition of the body reserves made during pre-laying by breeding European barn owls (Tyto alba), we have analysed the body composition of captive breeding and non-breeding females sacrificed during the laying period. The data obtained were compared to the daily requirement for egg formation obtained by an egg composition analysis and the timing of egg synthesis. This study demonstrates that body mass gain observed in breeding females (+38.3 g after eggs in formation and gonadal tractus were removed) was not the consequence of an accumulation of body fuels like lipids but of mainly water and lean material. The lipidic reserves were found to be less important in breeding than in non-breeding females and their localisation modified; lipids were absent from medullar bones in breeding females which liberated room for other storage. The subcutaneous tissue, which was homogeneous in non-breeding females, was located principally under the brood patch in breeding females. Nutrients and energy required during egg formation could be obtained without modification of daily food intake. These results show that a laying event can be initiated in 14 days and that the onset of reproduction is not triggered by body condition in barn owls. The water accumulation observed is suggested to be the mere consequence of an increase of protein metabolism (egg and moult). These results intimate that the body mass increase observed in diurnal and nocturnal raptors during laying preparation, interpreted as an energy safety strategy, ought to be reconsidered.

Testing for effects of climate change on competitive relationships and coexistence between two bird species.

Climate change is expected to have profound ecological effects, yet shifts in competitive abilities among species are rarely studied in this context. Blue tits (Cyanistes caeruleus) and great tits (Parus major) compete for food and roosting sites, yet coexist across much of their range. Climate change might thus change the competitive relationships and coexistence between these two species. Analysing four of the highest-quality, long-term datasets available on these species across Europe, we extend the textbook example of coexistence between competing species to include the dynamic effects of long-term climate variation. Using threshold time-series statistical modelling, we demonstrate that long-term climate variation affects species demography through different influences on density-dependent and density-independent processes. The competitive interaction between blue tits and great tits has shifted in one of the studied sites, creating conditions that alter the relative equilibrium densities between the two species, potentially disrupting long-term coexistence. Our analyses show that long-term climate change can, but does not always, generate local differences in the equilibrium conditions of spatially structured species assemblages. We demonstrate how long-term data can be used to better understand whether (and how), for instance, climate change might change the relationships between coexisting species. However, the studied populations are rather robust against competitive exclusion.

MORE EGGS THE BETTER: EGG FORMATION IN CAPTIVE BARN OWLS (TYTO ALBA)

Abstract We studied rapid yolk deposition (RYD) in Barn Owls (Tyto alba) on the basis of the analysis of 26 eggs laid by “dye-dosed” captive female Barn Owls in five different broods. Pictures of yolks were examined to assess daily rates of yolk deposition. We used those data in combination with data from the dissection of ovaries of another five breeding females. We found that the total duration between initiation of RYD and laying of the corresponding egg was only 13.6 days, with an interval between yolk completion and oviposition of 2.4 days. The total number of follicles that may have given eggs was found to be 25. That high number of follicles and the short RYD period explain the particularly high reproductive potential of this nocturnal raptor species.