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We test the predictive value of the main energetic currencies used in foraging theory using starlings that choose between two foraging modes (walking versus flying). Walking is low-cost, low-yield, whereas flying is the opposite. We fixed experimentally, at 11 different values, the amount of flight required to get one food reward, and for each flight cost value, we titrated the amount of walking until the birds showed indifference between foraging modes. We then compared the indifference points to those predicted by gross rate of gain over time, net rate of gain over time, and the ratio of gain to expenditure (efficiency). The results for the choice between modes show strong qualitative and quantitative support for net rate of gain over time over the alternatives. However, the birds foraged for only a fraction of the available time, indicating that the choice between foraging and resting could not be explained by any of these currencies. We suggest that this discrepancy could be accounted for functionally because nonenergetic factors such as predation risk may differ between resting and foraging in any mode but may not differ much between foraging modes, hence releasing the choice between foraging modes from the influence of such factors. Alternatively, the discrepancy may be attributable to the use of predictable (rather than stochastic) ratios of effort per prey in our experiment, and it may thus be better understood with mechanistic rather than functional arguments.

Original publication




Journal article


Proc Natl Acad Sci U S A

Publication Date





1089 - 1094


Animals, Animals, Wild, Basal Metabolism, Body Weight, Choice Behavior, Decision Making, Flight, Animal, Locomotion, Models, Biological, Predatory Behavior, Reward, Songbirds, Walking