Abstract Perturbations to the density of a species can
be propagated to distant members of a food web via
shifts in the density or the traits (i.e. behavior) of inter-
mediary species. Predators with differing foraging
modes may have different effects on prey behavior, and
these effects may be transmitted differently through food
webs. Here we test the hypothesis that shifts in the type
of predator present in a food web indirectly affect the
prey’s resource independent of changes in the density of
prey. We assessed the importance of predator identity in
mediating the grazing effects of the freshwater snail
Physa integra on its periphyton resources using field and
mesocosm studies. Field observations showed that Physa
used covered habitats more in ponds containing fish than
in ponds containing crayfish or no predators at all. A
field experiment confirmed that snail behavior depended
on predator identity. Physa placed near caged pumpkin-
seed sunfish (Lepomis gibbosus) selected covered habi-
tats, but Physa placed near caged crayfish (Orconectes
rusticus) moved to the surface of the water. The effects
of predator identity on periphtyon were then examined in
a mesocosm experiment, using caged predators. Habitat
use of Physa was similar to their habitat use in the field
experiment. In the presence of caged sunfish, periphyton
standing crop in covered habitats was reduced to 34% of
the standing crop in the presence of crayfish. In contrast,
periphyton in near-surface habitats was 110% higher in
the presence of fish than in the presence of crayfish.
Thus, the effects of predator identity on Physa behavior
cascaded through the food web to affect the abundance
and spatial distribution of periphyton.
Keywords Predator identity · Trait-mediated indirect
effects · Physa · Littoral food webs · Predator avoidance
Introduction
Changes in the abundance of predators affect not only the
population size of foragers, but also their behavioral, mor-
phological, life-historical, and physiological traits (re-
views in Kerfoot 1987; Lima 1998). Because the per-capi-
ta effect of foragers on other species may depend on their
phenotypic traits, perturbations to the abundance of top
predators can be transmitted through a food web via shifts
in the traits of intermediary species, independent of any
density effects (Turner and Mittelbach 1990). Interactions
in which the per-capita effects of one species on another
depend on the abundance of a third species have been
called higher-order interactions (Abrams 1983; Werner
1992), interaction modifications (Wooton 1994), and trait-
mediated indirect interactions (Abrams 1995; Abrams et
al. 1996). Theory suggests that trait-mediated indirect ef-
fects may be common and important in structuring food
webs (Abrams 1992, 1995; Werner 1992), and a few em-
pirical studies demonstrate that the strength of trait-medi-
ated indirect effects can rival that of density-mediated in-
direct effects (Peacor and Werner 1997; Schmitz et al.
1997; Peacor and Werner 2000; Relyea 2000).
To predict the occurrence of trait-mediated indirect
effects, community ecologists must first consider the
specific features of predators that induce trait shifts in
foragers. Most studies on trait-mediated indirect interac-
tions have focused on predator-induced shifts in habitat
use and foraging activity (Turner and Mittelbach 1990;
Peacor and Werner 1997; Turner 1997; Kuhara et al.
1999). Because predators vary in their hunting mode and
distribution, predators with differing foraging modes
have different effects on prey behavior (Sih et al. 1998;
McIntosh and Peckarsky 1999; Turner et al. 1999). Thus,
different predators will likely vary in the food web ef-
fects they create due to differing effects on prey traits
(McIntosh and Townsend 1996).
R.J. Bernot (
✉
) · A.M. Turner
Department of Biology, Clarion University of Pennsylvania,
Clarion, PA 16412, USA
e-mail: rjb5578@ksu.edu
Tel.: +1-785-5326814, Fax: +1-785-5326653
Present address:
R.J. Bernot, Division of Biology, Kansas State University,
232 Ackert Hall, Manhattan, KS 66506, USA
Oecologia (2001) 129:139–146
DOI 10.1007/s004420100705
Randall J. Bernot · Andrew M. Turner
Predator identity and trait-mediated indirect effects
in a littoral food web
Received: 23 October 2000 / Accepted: 27 February 2001 / Published online: 23 May 2001
© Springer-Verlag 2001