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First published online August 25, 2003
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The Journal of Experimental Biology 206, 3361-3368 (2003)
doi: 10.1242/jeb.00545

Cost-benefit analysis of mollusc eating in a shorebird I. Foraging and processing costs estimated by the doubly labelled water method

Theunis Piersma1,2, Anne Dekinga1, Jan A. van Gils1,2,*, Bart Achterkamp2 and G. Henk Visser2,3

1 Department of Marine Ecology and Evolution, Royal Netherlands Institute for Sea Research (NIOZ), PO Box 59, 1790 AB Den Burg, Texel, The Netherlands
2 Animal Ecology Group, Centre for Ecological and Evolutionary Studies, University of Groningen, PO Box 14, 9750 AA Haren, The Netherlands
3 Centre for Isotope Research, University of Groningen, Nijenborgh 4, 9747 AG Groningen, The Netherlands

* Author for correspondence (e-mail: janvg{at}nioz.nl)

Accepted 8 June 2003

Although the energy costs of foraging and food processing in vertebrates may be considerable, they have rarely been quantified separately. Here we present estimates for both cost factors based on a series of trials with a shorebird, the red knot Calidris canutus, fed natural and artificial prey types under naturalistic but fully controlled indoor aviary conditions. During eight 1-day trials we successfully manipulated the extent to which the five red knots were (1) actively probing and walking (i.e. foraging) and (2) actually ingesting prey (i.e. processing food) that was (3) either hard-shelled or not (i.e. crushing). Energy expenditures, estimated by the doubly labelled water (DLW) method, calibrated for use in this particular condition, varied between 1.5 and 4 W. A hierarchical analysis of variance indicated that the crushing of hard-shelled prey entailed no extra cost. We arrived at the following breakdown of cost components under the thermoneutral conditions of the experiment: a cost of active rest/maintenance of 1.665 W, an additional cost of foraging of 0.602 W and an additional digestive processing cost of 1.082 W. These cost levels are all well within the range of expectation and are consistent with the results of a separate outdoor aviary experiment in which the thermostatic costs needed separate estimation. On the basis of the cost and performance functions of gizzards of different mass, it was shown that under the conditions of this experiment the red knots expended the bare minimum for a balanced budget, maintaining the smallest possible gizzard. Under field conditions a larger gizzard would be required.

Key words: aviary experiment, bivalve, Calidris canutus, cost-benefit analysis, digestion, doubly labelled water, energetics, food selection, foraging, prey quality


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