Individual competitiveness conditions access to resources when they are limited. Immature individuals that are less skilled than adults have to adapt their foraging strategies to survive. Among ...strategies to reduce competition, spatial segregation has been widely demonstrated. However, the use of spatial segregation by immatures to limit intra‐specific competition with adults has rarely been tested.
In this study, we investigated and compared habitat preferences and distributions of free‐ranging immature and breeding adult northern gannets Morus bassanus in order to determine whether they compete for similar habitats during the year, and if this results in a spatial segregation between birds of different age groups.
Based on > 66 000 km of aerial surveys conducted in the North‐East Atlantic Ocean during winter and summer 2012, habitats selected by immatures and adult birds were modelled independently, linking gannet density to a set of oceanographic and physiographic predictors. Their large‐scale seasonal distribution was then predicted.
We found that gannets displayed a strong season‐dependent competition between immatures and adults, as a consequence of immatures and adults using similar habitats in both summer and winter. During summer, when adults are constrained by reproduction, both groups were spatially highly segregated despite similar habitat preferences (thermal fronts), with youngest individuals selecting habitats out of range of central‐place foragers, highlighting intra‐specific competition. Contrastingly during winter, when reproductive constraints disappear, immature and adult distributions largely overlapped.
Our study provides new insights into the role played by age, foraging experience and reproductive constraints on the distribution of marine predators. More specifically, these results highlight in seabirds how the youngest fraction mitigates, through spatial segregation, the competition with experienced adults, and suggest a progressive strategy along the maturation process.
Overfishing and ocean warming are drastically altering the community composition and size structure of marine ecosystems, eliminating large bodied species 1. Against a backdrop of such environmental ...change, the heaviest of all bony fish, the ocean sunfish (Mola mola), seems an improbable survivor. Indeed this indolent giant is killed globally as bycatch, and is listed as ‘Vulnerable’2. We undertook the most extensive aerial surveys of sunfish ever conducted and found surprisingly high abundances off the Atlantic and Mediterranean coasts of Western Europe. With up to 475 individuals per 100 km2, these figures are one order of magnitude higher than abundance estimates for other areas 3–5. Using bioenergetic modelling, we estimate that each sunfish requires 71 kg day–1 of jellyfish, a biomass intake more than an order of magnitude greater than predicted for a similarly sized teleost. Scaled up to the population level, this equates to a remarkable 20,774 tonnes day–1 of predated jellyfish across our study area in summer. Sunfish abundance may be facilitated by overfishing and ocean warming, which together cause reduced predation of sunfish by sharks and elevated jellyfish biomass. Our combined survey and bioenergetic data provide the first-ever estimate of spatialized ocean sunfish daily food requirements, and stress the importance of this species as a global indicator for the ‘rise of slime’. This hypothesis posits that, in an overfished world ocean exposed to global warming, gelatinous zooplankton should flourish, to the detriment of other mesotrophic species such as small pelagic fish, causing irreversible trophic cascades as well as a series of other environmental and economic issues.
The Ocean Sunfish is the heaviest of all bony fish, and is listed as vulnerable by the IUCN. Grémillet et al. conducted aerial surveys off the Atlantic and Mediterranean coasts of Western Europe, finding up to 475 sunfish per 100 km2. The sunfish population in this region is estimated to consume over 20,000 tonnes of jellyfish per day in summer.
Species Distribution Models are commonly used with surface dynamic environmental variables as proxies for prey distribution to characterise marine top predator habitats. For oceanic species that ...spend lot of time at depth, surface variables might not be relevant to predict deep-dwelling prey distributions. We hypothesised that descriptors of deep-water layers would better predict the deep-diving cetacean distributions than surface variables. We combined static variables and dynamic variables integrated over different depth classes of the water column into Generalised Additive Models to predict the distribution of sperm whales Physeter macrocephalus and beaked whales Ziphiidae in the Bay of Biscay, eastern North Atlantic. We identified which variables best predicted their distribution. Although the highest densities of both taxa were predicted near the continental slope and canyons, the most important variables for beaked whales appeared to be static variables and surface to subsurface dynamic variables, while for sperm whales only surface and deep-water variables were selected. This could suggest differences in foraging strategies and in the prey targeted between the two taxa. Increasing the use of variables describing the deep-water layers would provide a better understanding of the oceanic species distribution and better assist in the planning of human activities in these habitats.
In habitat modelling, environmental variables are assumed to be proxies of lower trophic levels distribution and by extension, of marine top predator distributions. More proximal variables, such as ...potential prey fields, could refine relationships between top predator distributions and their environment. In situ data on prey distributions are not available over large spatial scales but, a numerical model, the Spatial Ecosystem And POpulation DYnamics Model (SEAPODYM), provides simulations of the biomass and production of zooplankton and six functional groups of micronekton at the global scale. Here, we explored whether generalised additive models fitted to simulated prey distribution data better predicted deep-diver densities (here beaked whales Ziphiidae and sperm whales Physeter macrocephalus) than models fitted to environmental variables. We assessed whether the combination of environmental and prey distribution data would further improve model fit by comparing their explanatory power. For both taxa, results were suggestive of a preference for habitats associated with topographic features and thermal fronts but also for habitats with an extended euphotic zone and with large prey of the lower mesopelagic layer. For beaked whales, no SEAPODYM variable was selected in the best model that combined the two types of variables, possibly because SEAPODYM does not accurately simulate the organisms on which beaked whales feed on. For sperm whales, the increase model performance was only marginal. SEAPODYM outputs were at best weakly correlated with sightings of deep-diving cetaceans, suggesting SEAPODYM may not accurately predict the prey fields of these taxa. This study was a first investigation and mostly highlighted the importance of the physiographic variables to understand mechanisms that influence the distribution of deep-diving cetaceans. A more systematic use of SEAPODYM could allow to better define the limits of its use and a development of the model that would simulate larger prey beyond 1,000 m would probably better characterise the prey of deep-diving cetaceans.
At the onset of winter, warm‐blooded animals inhabiting seasonal environments may remain resident and face poorer climatic conditions, or migrate towards more favourable habitats. While the origins ...and evolution of migratory choices have been extensively studied, their consequences on avian energy balance and winter survival are poorly understood, especially in species difficult to observe such as seabirds. Using miniaturized geolocators, time‐depth recorders and a mechanistic model, we investigated the migratory strategies, the activity levels and the energy expenditure of the closely‐related, sympatrically breeding Brünnich's guillemots Uria lomvia and common guillemots Uria aalge from Bjørnøya, Svalbard. The two guillemot species from this region present contrasting migratory strategies and wintering quarters: Brünnich's guillemots migrate across the North Atlantic to overwinter off southeast Greenland and Faroe Islands, while common guillemots remain resident in the Barents, the Norwegian and the White Seas. Results show that both species display a marked behavioural plasticity to respond to environmental constraint, notably modulating their foraging effort and diving behaviour. Nevertheless, we provide evidence that the migratory strategy adopted by guillemots can have important consequences for their energy balance. Overall energy expenditure estimated for the non‐breeding season is relatively similar between both species, suggesting that both southward migration and high‐arctic winter residency are energetically equivalent and suitable strategies. However, we also demonstrate that the migratory strategy adopted by Brünnich's guillemots allows them to have reduced daily energy expenditures during the challenging winter period. We therefore speculate that ‘resident’ common guillemots are more vulnerable than ‘migrating’ Brünnich's guillemots to harsh winter environmental conditions.
From the Habitat Directive to the recent Marine Strategy Framework Directive, the conservation status of cetaceans in European water has been of concern for over two decades. In this study, a ...seasonal comparison of the abundance and distribution of cetaceans was carried out in two contrasted regions of the Eastern North Atlantic, the Bay of Biscay and the English Channel. Estimates were obtained in the two sub-regions (375,000km²) from large aerial surveys conducted in the winter (November 2011 to February 2012) and in the summer (May to August 2012). The most abundant species encountered in the Channel, the harbour porpoise, displayed strong seasonal variations in its distribution but a stable abundance (18,000 individuals, CV=30%). In the Bay of Biscay, abundance and distribution patterns of common / striped dolphins varied from 285,000 individuals (95% CI: 174,000–481,000) in the winter, preferentially distributed close to the shelf break, to 494,000 individuals (95% CI: 342,000−719,000) distributed beyond the shelf break in summer. Baleen whales also exhibited an increase of their density in summer. Seasonal abundances of bottlenose dolphins were quite stable, with a large number of ‘pelagic’ encounters offshore in winter. No significant seasonal difference was estimated for pilot whales and sperm whale. These surveys provided baseline estimates to inform policies to be developed, or for existing conservation instruments such as the Habitats Directive. In addition, our results supported the hypothesis of a shift in the summer distributions of some species such as harbour porpoise and minke whale in European waters.
The biodiversity of the Mediterranean Sea is undergoing important changes. Cetaceans, as top predators, are an important component of marine ecosystems. The seasonal distribution and abundance of ...several cetacean species were studied with a large aerial survey over the North-Western Mediterranean Sea, including the international Pelagos sanctuary, the largest Marine Protected Area (MPA) designed for marine mammals in the Mediterranean. A total of 8 distinct species of cetaceans were identified, and their occurrence within the sanctuary was investigated. Abundance estimates were obtained for three groups of species: the small delphinids (striped dolphins mainly), the bottlenose dolphin and the fin whale. There was a seasonal variation in striped dolphin abundance between winter (57,300 individuals, 95% CI: 34,500–102,000) and summer (130,000, 95% CI: 76,800–222,100). In contrast, bottlenose dolphin winter abundance was thrice that of summer. It was also the only species to exhibit any preference for the Pelagos sanctuary. Fin whale abundance had the reverse pattern with winter abundance (1000 individuals, 95% CI: 500–2500) and summer (2500 individuals, 95% CI: 1500–4300), without any preference for the sanctuary. Risso's dolphins, pilot whales and sperm whales did not exhibit strong seasonal pattern in their abundance. These results provide baseline estimates which can be used to inform conservation policies and instruments such as the Habitats Directive or the recent European Marine Strategy Framework Directive.
Contrasting to the overall oligotrophic Mediterranean Sea, the north-western basin is characterised by high productivity and marked by seasonality, which induces spatiotemporal heterogeneity of ...habitat. Cetaceans and seabirds are expected to perceive this repetition of the seasonal cycle and to anticipate the recurrent variability of their environment. Because phenology imposes strong constraints over marine predators, especially through reproduction, we expected them to exhibit variations in their habitat preferences over seasons. Indeed, during reproductive period, marine predators have to face their own needs and those of their young, while out of this period, they can focus on maximising their own survival only. We therefore hypothesised that some species would change their habitat preferences to exploit the most favourable habitat during each season, while other species might accommodate the same habitat all year-round, for example thanks to the use of an habitat favourable all the year. To explore these hypotheses, we used aerial surveys data conducted over north-western Mediterranean Sea during winter 2011–2012 and summer 2012. Generalised Additive Models were used to link the species density to a set of 12 physiographic and oceanographic predictors describing their environment. Habitat models resulted in deviances from 12 to 47%. Our results provided the first assessment of habitat preferences for the winter season for most of our studied species. Small-sized delphinids (mostly stripped dolphins), fin whales, Globicephalinae (long-finned pilot whales and Risso׳s dolphins) and small-sized shearwaters (Yelkouan and Balearic shearwaters) exhibited no habitat variations between seasons, although for the first two, abundances were lower in winter. On the contrary, bottlenose dolphins switched from coastal habitat in summer to pelagic habitat in winter, while Cory׳s shearwaters and storm petrels exhibited the largest habitat variations between seasons with a complete absence during winter.
Seabird distributions and the associated seasonal variations remain challenging to investigate, especially in oceanic areas. Recent advances in telemetry have provided considerable information on ...seabird ecology, but still exclude small species, non-breeding birds and individuals from inaccessible colonies from any scientific survey. To overcome this issue and investigate seabird distribution and abundance in the eastern North Atlantic (ENA), large-scale aerial surveys were conducted in winter 2011-12 and summer 2012 over a 375,000km2 area encompassing the English Channel (EC) and the Bay of Biscay (BoB). Seabird sightings, from 15 taxonomic groups, added up to 17,506 and 8263 sightings in winter and summer respectively, along 66,307km. Using geostatistical methods, density maps were provided for both seasons. Abundance was estimated by strip transect sampling. Most taxa showed marked seasonal variations in their density and distribution. The highest densities were recorded during winter for most groups except shearwaters, storm-petrels, terns and large-sized gulls. Subsequently, the abundance in winter nearly reached one million individuals and was 2.5 times larger than in summer. The continental shelf and the slope in the BoB and the EC were identified as key areas for seabird conservation, especially during winter, as birds from northern Europe migrate southward after breeding. This large-scale study provided a synoptic view of the seabird community in the ENA, over two contrasting seasons. Our results highlight that oceanic areas harbour an abundant avifauna. Since most of the existing marine protected areas are restricted to the coastal fringe, the importance of oceanic areas in winter should be considered in future conservation plans. Our work will provide a baseline for the monitoring of seabird distribution at sea, and could inform the EU Marine Strategy Framework Directive.
Marine Protected Areas (MPAs) are being established across all marine regions but their validity for the conservation of highly mobile marine vertebrates has been questioned. We tested the hypothesis ...that French coastal MPAs primarily designed for coastal and benthic biota are also beneficial for the conservation of a pelagic seabird, the Vulnerable yelkouan shearwater (Puffinus yelkouan), an endemic species to the Mediterranean Sea. We used a vast spectrum of electronic devices (GPS, temperature-depth-recorders, satellite transmitters and geolocators) and stable isotopic analyses to study the year-round movements and the trophic status of yelkouan shearwaters from the Hyères archipelago (France). In addition we conducted large-scale ship and aircrafts observation surveys to investigate spatio-temporal density patterns of shearwaters (genus Puffinus) in the western Mediterranean Sea. This extensive investigation permitted the first comprehensive study of the at-sea ecology of yelkouan shearwaters showing strikingly coastal habits, partial migration, unsuspected diving capabilities (max dive depth of 30m), and a broad diet ranging from zooplankton to small pelagic fish. Importantly, 31% of yelkouan shearwaters GPS positions associated with foraging, 38% of diving positions, and 27% of resting positions were within the three French MPAs during the breeding season. These high scores confirmed by year-round distribution derived from GLS, PTTs, at-sea and aerial observations, validated our hypothesis of the major importance of coastal MPAs for the conservation of yelkouan shearwater. Our case-study is therefore a major contribution to research efforts aiming at linking the spatial ecology of highly mobile marine vertebrates with effective conservation of marine biodiversity.
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