OIKOS 93: 488 – 496. Copenhagen 2001
Shape and sources of variations of the functional response of
wildfowl: an experiment with mallards, Anas platyrhynchos
Herve ´ Fritz, Daphne ´ Durant and Matthieu Guillemain
Fritz, H., Durant, D. and Guillemain, M. 2001. Shape and sources of variations of
the functional response of wildfowl: an experiment with mallards, Anas platyrhyn -
chos. – Oikos 93: 488–496.
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 interfer-
ence-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.
H. Fritz, D. Durant and M. Guillemain, CNRS -UPR 1934, Centre d’Etudes Bi -
ologiques de Chize ´ , 79360 Beauoir -sur -Niort, France ( fritzh@cebc.cnrs.fr).
Measuring and understanding the causes of variations
of the functional response of an organism, i.e. instan-
taneous predation rate in relation to prey density
(Holling 1959), have direct consequences for the un-
derstanding of ecological processes at the individual
level, such as the role of dominance and interference
on foraging efficiency (Goss-Custard et al. 1984, Still-
man et al. 1996, Norris and Johnstone 1998), and to
evaluate the role of these individual differences at the
population level (Rubenstein 1981, Goss-Custard et al.
1995, Piersma et al. 1995). There are several
mechanisms that can generate the various functional
responses, particularly the asymptotic part: the compe-
tition between handling and searching for a
prey, but also the interference between harvesting food
and the velocity of the animal, or the competition
between cropping and processing food (see Spalinger
and Hobbs 1992 for the theoretical approach on herbi-
vores). The understanding of the shape and parameters
of the functional response can thus allow for the test-
ing of hypotheses related to processes limiting intake
rate (Gross et al. 1993).
Accepted 19 January 2001
Copyright © OIKOS 2001
ISSN 0030-1299
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OIKOS 93:3 (2001) 488