Predicting Effects of Water Regime Changes on Waterbirds: Insights from Staging Swans.

Predicting the environmental impact of a proposed development is notoriously difficult, especially when future conditions fall outside the current range of conditions. Individual-based approaches have been developed and applied to predict the impact of environmental changes on wintering and staging coastal bird populations. How many birds make use of staging sites is mostly determined by food availability and accessibility, which in the case of many waterbirds in turn is affected by water level. Many water systems are regulated and water levels are maintained at target levels, set by management authorities. We used an individual-based modelling framework (MORPH) to analyse how different target water levels affect the number of migratory Bewick's swans Cygnus columbianus bewickii staging at a shallow freshwater lake (Lauwersmeer, the Netherlands) in autumn. As an emerging property of the model, we found strong non-linear responses of swan usage to changes in water level, with a sudden drop in peak numbers as well as bird-days with a 0.20 m rise above the current target water level. Such strong non-linear responses are probably common and should be taken into account in environmental impact assessments.

Nonlinear effects of food aggregation on interference competition in mallards.

Previous studies of interference competition have shown an asymmetric effect on intake rate of foragers on clumped resources, with only subordinate individuals suffering. However, the food distributions in these studies were uniform or highly clumped, whereas in many field situations, food aggregation is intermediate. Here we investigated whether food distribution (i.e., uniform, slightly clumped, and highly clumped) affects the behavioral response of mallards foraging alone or competing with another. Although the amount of food was the same in all distributions, the mallards reached higher intake rates, visited fewer patches, and showed longer average feeding times in the highly clumped distribution. Competing mallards had lower intake rates on the slightly clumped than on the uniform or highly clumped food distributions. Subordinates generally visited more patches and had shorter feeding times per patch, but their intake rates were not significantly lower than those of dominants. Therefore, we propose that subordinates do not necessarily suffer from interference competition in terms of intake rate, but do suffer higher search costs. In addition, although dominants had significantly higher average feeding times on the best quality patches of the highly clumped food distribution, such an effect was not found in the slightly clumped distribution. These findings indicate that in environments where food is aggregated to a lesser extent, monopolization is not the best strategy for dominants. Our results suggest that interference experiments should use food distributions that resemble the natural situation animals are faced with in the field.

Avian herbivory: an experiment, a field test, and an allometric comparison with mammals.

Mechanistic studies on herbivore functional responses have largely taken place in mammals; very little has been done in herbivorous birds so far. Here we aim to fill that gap by experimentally quantifying the (short-term) functional response of a large avian herbivore, the Bewick's Swan (Cygnus columbianus bewickii). We explicitly distinguish between encounter-limited and handling-limited foraging by analyzing the results in the framework of the models of D. E. Spalinger and N. T. Hobbs, originally developed for mammalian herbivory. Bite size in captive swans was experimentally manipulated by varying sward height. The time interval between two bites increased with bite size, which supports the handling-limited model (process 3) and rejects the encounter-limited models (processes 1 and 2). Subsequently, we took the obtained functional response parameters into the field in order to predict, from measurements of sward height, (1) bite sizes, (2) handling times, and (3) short-term intake rates in free-ranging swans. Indeed, for all three variables, the observed values closely matched the experimentally based predictions. Finally, we review functional response parameters available in the literature on avian herbivores and scale them allometrically in relation to mammals. This analysis revealed that maximum bite sizes, and therefore maximum intake rates, in herbivorous birds are smaller than in herbivorous mammals. We hypothesize and provide evidence that birds compensate by longer daily foraging times.

Prediction of bird-day carrying capacity on a staging site: a test of depletion models.

1. The carrying capacity of a site for migratory water birds, expressed in bird-days, can be of particular conservation value. Several attempts have been made to model this carrying capacity using ideal free distribution models such as, for instance, depletion models, in which the distribution is fully determined by exploitative competition. 2. In the tests of depletion models carried out so far, no alternative models were compared; rather, one specific model was tested. We tested whether bird-days were more in accordance with birds depleting the food resource (a1) until a critical food density which just enabled survival or (a2) until a threshold food density which renders the site as profitable as an alternative site; and birds (b1) satisfying their daily requirements or (b2) maximizing daily intake. 3. We studied Bewick's swans feeding on below-ground tubers of fennel pondweed in one part of an autumn staging site. In most years between 1995 and 2005, we measured tuber biomass densities around September, November and March, and counted swans daily during their stopover in October. 4. The best fit between observed and predicted bird-days was obtained by assuming that the swans were maximizing their daily intake and depleting the tubers until a threshold biomass density (which yielded the same energetic return as the alternative food source after accounting for a small part of the initial tuber biomass being out of reach of the swans). Also in line with daily intake maximization, the daily feeding time did not differ from 10 h day(-1), the value predicted for Bewick's swans based on their feeding costs. 5. Our results suggests that the applicable model to calculate carrying capacity may depend strongly on whether birds use a site to stopover or to winter, because it determines whether the birds are more likely to use a threshold or critical food density, and to behave as energy maximizers or satisficers.