Summary1. The functional response, i.e. the quantity of food consumed per unit of time as a function of food availability, is a central process in foraging ecology. The application of this concept to foraging by mammalian herbivores has led to major insights into the process of resource acquisition, but it has so far been little used to understand foraging in avian herbivores. 2. In this study, we describe the functional responses of three grazing Anatidae, the wigeon Anas penelope L. (mean body mass in this study 620 g), the barnacle goose Branta leucopsis B. (2000 g), and the greylag goose Anser anser L. (3500 g). We measured instantaneous intake rates on eight grass heights from 1 cm to 12 cm, as well as pecking rates and peck sizes, and used the Spalinger-Hobbs model developed for mammalian herbivores to explore the mechanisms limiting intake in these three species. 3. Greylag geese increased their intake rate with increasing food availability (a Type II response), wigeon showed a weak quadratic (Type IV) response, and intake rate by barnacle geese did not vary significantly across the range of variability we offered the birds. 4. Intake rates differed markedly between individuals, especially in greylag geese, where body mass explained much of the variation. For individuals in all three species, peck sizes strongly influenced instantaneous intake rates, and the size of the bill (width in particular) appeared to be an important determinant of peck size. 5. Peck sizes increased with sward height (although only very weakly in wigeon), but so did cropping time for wigeon and barnacle geese, which led to a significant decline of intake rates on the tallest grass, at least in the small wigeon. For these very selective small herbivores, the time to crop a mouthful was therefore a significant limiting factor for the birds' instantaneous intake rate (in addition to peck size and swallowing time). This differs markedly from the situation in mammalian herbivores where bite size (through chewing time) is the principal controlling process in food concentrated patches, a result that we found in greylag geese. We discuss the differences in foraging between the three species in relation to their principal food resources, body mass and bill morphology.

Although agriculture is pointed to as being one of the major causes of biodiversity loss, it is now recognised that some farming practices, grazing in particular, represent central issues in the on-running debate on wildlife conservation. This paper analyses the relationship between agricultural pasture management and bird preservation, by focusing on the illustrative case study of breeding waders in wet grasslands. After a brief account of the knowledge on research investigating the effect of grazing management on wader distribution or abundance, we reviewed (i) sward structure (i.e. mean sward height and frequency of tussocks) requirements, and (ii) breeding phenologies of five wader species commonly associated with wet grasslands for nesting in Europe. We found differences between species in these two aspects of their biology. Investigating grassland management thus underlines that the question of the timing of grazing should be considered. Moreover, the fragmented characteristic of agricultural landscapes (i.e. a mosaic of fields) in which waders breed emphasises the spatial nature of this management. We examine three spatial scales (i.e. field, set of adjacent fields, landscape) relevant to bird biology and influenced by agricultural activities. Based on these findings, we analyse which livestock system constraints are essential to take into account in order to minimise conflicts between livestock production and conservation aims in marshes. Finally, we highlight possible directions for future research.

Recent findings suggest that herbivores select feeding sites of intermediate biomass in order to maximise their digestible nutrient intake as the result of the trade-off between forage quality and quantity ('forage maturation hypothesis'). We propose a reformulation of this hypothesis which recognises this trade-off, but also underlines that constraints due to body mass (i.e. metabolism and digestive constraints, size of the feeding apparatus) can lead to variations in grazing patterns. We tested this latter hypothesis experimentally in three species of herbivorous Anatidae of different body mass: the wigeon Anas penelope (in our study c. 620 g), the barnacle goose Branta leucopsis (c. 2000 g), and the greylag goose Anser anser (c. 3500 g). Each species was tested separately from 0600 to 0930 hours, in an enclosure with a mosaic of patches of grass of three different heights: short, medium and tall. The behaviour, and the location (i.e. patch) of each individual were recorded every 5 minutes. Our results show important interspecific differences in intake rates resulting in different feeding site selection: wigeon and barnacle goose fed fastest on the shortest swards, and selected short grass which was also of higher quality. Tall grass provided the highest dry matter intake rate and digestible protein intake for greylag geese, and they preferred these swards. These choices allowed the birds to maximise their digestible nitrogen intake rate rather than dry matter intake rate and our results thus underline the importance of nitrogen as a major currency for foraging decisions in herbivorous Anatidae. Since the birds selected the two extreme sward heights (instead of the medium one), the results give support to our hypothesis and underline the role of body size as an important cause of variations in patch selection in herbivorous Anatidae. Foraging theory postulates that decisions made by a foraging animal optimise acquisition of some currency (Stephens and Krebs 1986) which determines its fitness (Schoener 1971). This currency has often been assumed to be the net rate of dry matter intake, which has led to the formulation of the 'intake rate maximisation theory' (Krebs and McCleery 1984). Selection of feeding sites by vertebrate herbivores has been studied frequently, in particular in relation to patchy food resources (Langvatn and Hanley 1993, Weckerly 1994, Wilmshurst et al. 1995, Hupp and Robertson 1998. In support of the 'intake rate maximisation theory', it has been shown that sheep prefer grazing in patches offering the highest dry matter intake rates (Kenney and Black 1984, Illius et al. 1992), as do goats (Illius et al. 1999) and cattle (Demment et al. 1993).However, as grasses mature, high biomass patches where the animals can achieve the highest dry matter intake rates, tend to be of poorer quality, i.e. to have higher fibre content, lower digestibility (Nehring and Nerge 1966, Prop andVulink 1992), and lower nutrient contents (Summers and Critchley 1990 (Palo and Robbins 1991). Herbivores should therefo...

Understanding the variations of the functional response of an organism, i.e. the predation rate in relation to prey density, is necessary to understand the interactions between the animal and its food supply. This has received little attention in dabbling ducks so we investigated experimentally the shape of the functional response of mallard feeding on poultry pellets, and assessed the influence of several factors such as the size of food items, sex or individual performance on this functional response. Individual differences in intake rate are of crucial importance in group or gregarious foraging species. We used two approaches of the functional response: 1) the relation between feeding rate (pellets/s) and pellet densities (pellets/m 2 ), and 2) the relationship between instantaneous intake rate (g/s) and biomass density (g/m 2 ). For both approaches, we found that the Type II functional response gave better estimates than a Type I linear functional response but explained only a third of the variance. Our results show that pellet size has a large effect on instantaneous intake rate. The comparison of the functional response parameters suggest that handling time per prey may not reflect the real constraints on intake rate, but that handling time per gram ingested may be more appropriate to integrate the effect of item size in the functional response. We then discuss the possible mechanisms involved. We also found individual variations in the functional response for each of the experiments, with some consistency in the hierarchy regarding feeding efficiency. We did not find any differences between males and females. Our results provide an evaluation of individual variations in intake rate in interference-free conditions, which has rarely been done, and call for more controlled experiments to allow a finer understanding of the mechanisms of food acquisition in dabbling ducks.