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

P>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.