Key features of intertidal food webs that support migratory shorebirds.

The migratory shorebirds of the East Atlantic flyway land in huge numbers during a migratory stopover or wintering on the French Atlantic coast. The Brouage bare mudflat (Marennes-Oléron Bay, NE Atlantic) is one of the major stopover sites in France. The particular structure and function of a food web affects the efficiency of carbon transfer. The structure and functioning of the Brouage food web is crucial for the conservation of species landing within this area because it provides sufficient food, which allows shorebirds to reach the north of Europe where they nest. The aim of this study was to describe and understand which food web characteristics support nutritional needs of birds. Two food-web models were constructed, based on in situ measurements that were made in February 2008 (the presence of birds) and July 2008 (absence of birds). To complete the models, allometric relationships and additional data from the literature were used. The missing flow values of the food web models were estimated by Monte Carlo Markov Chain--Linear Inverse Modelling. The flow solutions obtained were used to calculate the ecological network analysis indices, which estimate the emergent properties of the functioning of a food-web. The total activities of the Brouage ecosystem in February and July are significantly different. The specialisation of the trophic links within the ecosystem does not appear to differ between the two models. In spite of a large export of carbon from the primary producer and detritus in winter, the higher recycling leads to a similar retention of carbon for the two seasons. It can be concluded that in February, the higher activity of the ecosystem coupled with a higher cycling and a mean internal organization, ensure the sufficient feeding of the migratory shorebirds.

Scaling up ideals to freedom: are densities of red knots across western Europe consistent with ideal free distribution?

Local studies have shown that the distribution of red knots Calidris canutus across intertidal mudflats is consistent with the predictions of an ideal distribution, but not a free distribution. Here, we scale up the study of feeding distributions to their entire wintering area in western Europe. Densities of red knots were compared among seven wintering sites in The Netherlands, UK and France, where the available mollusc food stocks were also measured and from where diets were known. We tested between three different distribution models that respectively assumed (i) a uniform distribution of red knots over all areas, (ii) a uniform distribution across all suitable habitat (based on threshold densities of harvestable mollusc prey), and (iii) an ideal and free distribution (IFD) across all suitable habitats. Red knots were not homogeneously distributed across the different European wintering areas, also not when considering suitable habitats only. Their distribution was best explained by the IFD model, suggesting that the birds are exposed to interference and have good knowledge about their resource landscape at the spatial scale of NW Europe, and that the costs of movement between estuaries, at least when averaged over a whole winter, are negligible.

Diet selection in a molluscivore shorebird across Western Europe: does it show short- or long-term intake rate-maximization?

1. Studies of diet choice usually assume maximization of energy intake. The well-known 'contingency model' (CM) additionally assumes that foraging animals only spend time searching or handling prey. Despite considerable empirical support, there are many foraging contexts in which the CM fails, but such cases were considered exceptions rather than the rule. 2. For animals constrained by the rate at which food is digested, CM does not necessarily lead to maximal energy intake rates because the time for digestion is not part of the selection criteria. In the main model developed to explain diet choice under a digestive constraint, the 'digestive rate model' (DRM), time lost to digestive breaks is minimized so that energy intake over total time (searching, handling, digestive breaks) is maximized. 3. It is increasingly acknowledged that most animals may face digestive constraints as prey capture rates vary over time and as it would be a waste to carry around heavy digestive machinery that can rapidly process food under all circumstances: this is only needed in times of high demand, provided that enough food can be found. 4. In molluscivore shorebirds ingesting hard-shelled prey such as red knots (Calidris canutus), the predictions of DRM were held up so far, whereas those of CM were rejected. However, most tests were carried out under controlled experimental conditions. Red knots overwinter in coastal areas over much of Western Europe and we capitalized on this variation by comparing, during a single winter, observed diet composition with predictions of DRM, CM and a null model assuming no prey selection ('no-selection model', NSM). 5. The observed diets were best predicted by DRM followed by CM. NSM poorly predicted observed diet choice. Under the present conditions, diet choice based on DRM would on average have yielded an energy intake rate twice as large as one based on CM. By adjusting the size of their gizzard (held constant in the present simulations), red knots could have lifted their energy intake rate further. We suggest that application of the DRM can help many diet studies forward, especially those previously seen as exceptions to the classical CM-based rule.