Energyscapes and prey fields shape a North Atlantic seabird wintering hotspot under climate change.

There is an urgent need for a better understanding of animal migratory ecology under the influence of climate change. Most current analyses require long-term monitoring of populations on the move, and shorter-term approaches are needed. Here, we analysed the ecological drivers of seabird migration within the framework of the energyscape concept, which we defined as the variations in the energy requirements of an organism across geographical space as a function of environmental conditions. We compared the winter location of seabirds with their modelled energy requirements and prey fields throughout the North Atlantic. Across six winters, we tracked the migration of 94 little auks (Alle alle), a key sentinel Arctic species, between their East Greenland breeding site and wintering areas off Newfoundland. Winter energyscapes were modelled with Niche Mapper™, a mechanistic tool which takes into account local climate and bird ecophysiology. Subsequently, we used a resource selection function to explain seabird distributions through modelled energyscapes and winter surface distribution of one of their main prey, Calanus finmarchicus. Finally, future energyscapes were calculated according to IPCC climate change scenarios. We found that little auks targeted areas with high prey densities and moderately elevated energyscapes. Predicted energyscapes for 2050 and 2095 showed a decrease in winter energy requirements under the high emission scenario, which may be beneficial if prey availability is maintained. Overall, our study demonstrates the great potential of the energyscape concept for the study of animal spatial ecology, in particular in the context of global change.

Microplastic pollution in the Greenland Sea: Background levels and selective contamination of planktivorous diving seabirds.

Microplastics have been reported everywhere around the globe. With very limited human activities, the Arctic is distant from major sources of microplastics. However, microplastic ingestions have been found in several Arctic marine predators, confirming their presence in this region. Nonetheless, existing information for this area remains scarce, thus there is an urgent need to quantify the contamination of Arctic marine waters. In this context, we studied microplastic abundance and composition within the zooplankton community off East Greenland. For the same area, we concurrently evaluated microplastic contamination of little auks (Alle alle), an Arctic seabird feeding on zooplankton while diving between 0 and 50 m. The study took place off East Greenland in July 2005 and 2014, under strongly contrasted sea-ice conditions. Among all samples, 97.2% of the debris found were filaments. Despite the remoteness of our study area, microplastic abundances were comparable to those of other oceans, with 0.99 ± 0.62 m-3 in the presence of sea-ice (2005), and 2.38 ± 1.11 m-3 in the nearby absence of sea-ice (2014). Microplastic rise between 2005 and 2014 might be linked to an increase in plastic production worldwide or to lower sea-ice extents in 2014, as sea-ice can represent a sink for microplastic particles, which are subsequently released to the water column upon melting. Crucially, all birds had eaten plastic filaments, and they collected high levels of microplastics compared to background levels with 9.99 and 8.99 pieces per chick meal in 2005 and 2014, respectively. Importantly, we also demonstrated that little auks took more often light colored microplastics, rather than darker ones, strongly suggesting an active contamination with birds mistaking microplastics for their natural prey. Overall, our study stresses the great vulnerability of Arctic marine species to microplastic pollution in a warming Arctic, where sea-ice melting is expected to release vast volumes of trapped debris.

Assessing the validity of the accelerometry technique for estimating the energy expenditure of diving double-crested cormorants Phalacrocorax auritus.

Over the past few years, acceleration-data loggers have been used to provide calibrated proxies of energy expenditure: the accelerometry technique. Relationships between rate of oxygen consumption and a derivation of acceleration data termed "overall dynamic body acceleration" (ODBA) have now been generated for a range of species, including birds, mammals, and amphibians. In this study, we examine the utility of the accelerometry technique for estimating the energy expended by double-crested cormorants Phalacrocorax auritus to undertake a dive cycle (i.e., a dive and the subsequent pause at the surface before another dive). The results show that ODBA does not calibrate with energy expenditure in diving cormorants, where energy expenditure is calculated from measures of oxygen uptake during surface periods between dives. The possible explanations include reasons why energy expenditure may not relate to ODBA but also reasons why oxygen uptake between dives may not accurately represent energy expenditure during a dive cycle.

Northern gannets anticipate the spatio-temporal occurrence of their prey.

Seabirds, as other marine top predators, are often assumed to forage in an unpredictable environment. We challenge this concept and test the hypothesis that breeding Northern gannets (Morus bassanus) anticipate the spatio-temporal occurrence of their prey in the English Channel. We analyzed 23 foraging tracks of Northern gannets breeding on Rouzic Island (Brittany) that were recorded using GPS loggers during 2 consecutive years. All birds commuted between the breeding colony and foraging areas located at a mean distance of 85 km and 72 km (in 2005 and 2006, respectively) from the colony. Mean linearity indices of the outbound and inbound trips were between 0.83 and 0.87, approaching a beeline path to and from the foraging area. Additional parameters (flight speed, and number and duration of stopovers at sea) for the outbound and inbound trip were not statistically different, indicating that birds are capable of locating these feeding areas in the absence of visual clues, and to pin-point their breeding site when returning from the sea. Our bearing choice analysis also revealed that gannets anticipate the general direction of their foraging area during the first 30 min and the first 10 km of the trip. These results strongly suggest that birds anticipate prey location, rather than head into a random direction until encountering a profitable area. Further investigations are necessary to identify the mechanisms involved in seabird resource localization, such as sensorial abilities, memory effects, public information or a combination of these factors.

Influence of wing loading on the trade-off between pursuit-diving and flight in common guillemots and razorbills.

Species of bird that use their wings for underwater propulsion are thought to face evolutionary trade-offs between flight and diving, leading to the prediction that species with different wing areas relative to body mass (i.e. different wing loadings) also differ in the relative importance of flight and diving activity during foraging trips. We tested this hypothesis for two similarly sized species of Alcidae (common guillemots and razorbills) by using bird-borne devices to examine three-dimensional foraging behaviour at a single colony. Guillemots have 30% higher wing loading than razorbills and, in keeping with this difference, razorbills spent twice as long in flight as a proportion of trip duration whereas guillemots spent twice as long in diving activity. Razorbills made a large number of short, relatively shallow dives and spent little time in the bottom phase of the dive whereas guillemots made fewer dives but frequently attained depths suggesting that they were near the seabed (ca. 35-70 m). The bottom phase of dives by guillemots was relatively long, indicating that they spent considerable time searching for and pursuing prey. Guillemots also spent a greater proportion of each dive bout underwater and had faster rates of descent, indicating that they were more adept at maximising time for pursuit and capture of prey. These differences in foraging behaviour may partly reflect guillemots feeding their chicks single large prey obtained near the bottom and razorbills feeding their chicks multiple prey from the water column. Nonetheless, our data support the notion that interspecific differences in wing loadings of auks reflect an evolutionary trade-off between aerial and underwater locomotion.

Marine no-take zone rapidly benefits endangered penguin.

No-take zones may protect populations of targeted marine species and restore the integrity of marine ecosystems, but it is unclear whether they benefit top predators that rely on mobile pelagic fishes. In South Africa, foraging effort of breeding African penguins decreased by 30 per cent within three months of closing a 20 km zone to the competing purse-seine fisheries around their largest colony. After the fishing ban, most of the penguins from this island had shifted their feeding effort inside the closed area. Birds breeding at another colony situated 50 km away, whose fishing grounds remained open to fishing, increased their foraging effort during the same period. This demonstrates the immediate benefit of a relatively small no-take zone for a marine top predator relying on pelagic prey. Selecting such small protected areas may be an important first conservation step, minimizing stakeholder conflicts and easing compliance, while ensuring benefit for the ecosystems within these habitats.

Fine-scale foraging behaviour of a medium-ranging marine predator.

1. Movement patterns of predators should allow them to detect and respond to prey patches at different spatial scales, particularly through the adoption of area-restricted search (ARS) behaviour. Here we use fine-scale movement and activity data combined with first-passage time (FPT) analysis to examine the foraging strategy of northern gannets Morus bassanus in the western North Sea, and to test the following hypotheses: (i) birds adopt a hierarchical foraging strategy characterized by nested ARS behaviour; (ii) the locations and characteristics of ARS zones are strongly influenced by physical oceanography; (iii) the initiation of ARS behaviour is triggered by the detection and pursuit of prey; (iv) ARS behaviour is strongly linked to increased foraging effort, particularly within nested ARS areas. 2. Birds on 13 of 15 foraging trips adopted ARS behaviour at a scale of 9.1 +/- 1.9 km, and birds on 10 of these 13 trips adopted a second, nested ARS scale of 1.5 +/- 0.8 km, supporting hypothesis 1 above. ARS zones were located 117 +/- 55 km from the colony and over half were within 5 km of a tidal mixing front ~50 km offshore, supporting hypothesis 2 above. 3. The initiation of ARS behaviour was usually followed after only a short time interval (typically ~5 min) by the commencement of diving. Gannets do not dive until after they have located prey, and so this pattern strongly suggests that ARS behaviour was triggered by prey detection, supporting hypothesis 3 above. However, ~33% of dives in mixed coastal water and 16% of dives in stratified water were not associated with any detectable ARS behaviour. Hence, while ARS behaviour resulted from the detection and pursuit of prey, encounters with prey species did not inevitably induce ARS behaviour. 4. Following the initiation of ARS behaviour, dive rates were almost four times higher within ARS zones than elsewhere and almost three times higher in zones with nested ARS behaviour than in those without, supporting hypothesis 4 above and suggesting that the foraging success of birds was linked to their ability to match the hierarchical distribution of prey.

Body temperatures of free-living African penguins (Spheniscus demersus) and bank cormorants (Phalacrocorax neglectus).

Two free-living seabirds (the African penguin Spheniscus demersus and the bank cormorant Phalacrocorax neglectus) were equipped with stomach temperature-loggers to study body temperature changes during foraging. Body temperature in these endotherms was environmentally and activity-dependent and varied in the case of the cormorant by over 5 degrees C. Considerations of heat flux show that such flexibility confers considerable energetic advantages: by allowing body temperature to drop when the heat loss to the environment is high, such as in water, birds may save the energy that would normally be necessary to compensate for this drop. It appears that, in cormorants, low body temperature resulting from extended time in water can subsequently be elevated using solar energy when birds return to land in a manner similar to that of ectotherms. In the better-insulated penguins, muscle-generated heat during swimming is used to re-elevate low body temperature. Continued swimming eventually causes body temperature to rise above normal resting levels so that metabolic rate could theoretically be dramatically reduced immediately post-exercise when the temperature drops to some critical level before any increase in metabolism is necessary to correct it.