Measuring the food left in experimental trays when study organisms cease feeding on them [so-called giving-up densities (GUDs)] is an accepted technique for assessing predation risk and disturbance. However, in natural settings, accessibility and energetic harvest costs may vary spatially, and GUDs may be confounded. In this study, we assessed whether at a heterogeneous site, non-experimental GUDs could reveal the effect of disturbance. We measured initial and GUDs of tubers of Fennel Pondweed Potamogeton pectinatus, which form here the exclusive food source of Bewick's Swans Cygnus columbianus bewickii during their migratory stopover. We calculated giving-up net energy intake rates (GUNs) by correcting for biomass accessibility and foraging costs. The study area was a shallow lake consisting of nine creeks, three of which were open to the public (i.e. disturbed). GUDs in creeks open or closed to the public were not significantly different. In contrast, GUNs were generally higher in creeks open to the public, after correcting for initial net energy intake rate. The results suggest that natural GUDs may not reflect the effects of disturbance in heterogeneous habitats. When environmental differences are large within a site, GUNs may be a useful alternative as a behavioural indicator.
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