Oj. Schmitz, DIRECT AND INDIRECT EFFECTS OF PREDATION AND PREDATION RISK IN OLD-FIELD INTERACTION WEBS, The American naturalist, 151(4), 1998, pp. 327-342
Indirect effects emerge when a change in the: abundance of one species
indirectly affects another by changing the abundances of intermediate
species-called density-mediated indirect effects-or they arise when o
ne species modifies how two other species interact-called trait-mediat
ed indirect effects. I report on field experiments that evaluated how
grass and herb biomass in old-field interaction webs was influenced in
directly by a spider carnivore through its interactions with a general
ist and a grass-specialist grasshopper species. I manipulated interact
ion pathways between the spider and the plants using different combina
tions of the grasshopper species. I changed the modality of predator-p
rey interactions to isolate density-mediated from trait-mediated effec
ts using natural spiders (predation spiders) or spiders that were prev
ented from subduing prey by mouthpart manipulation (risk spiders). I f
ound that indirect effects were stronger in speciose, reticulate food
webs than in linear food chains owing to a trait-mediated effect, a di
et shift by herbivores in response to predation risk. Spiders alone di
d not have significant effects on grasshopper densities in the field e
xperiments, removing any possibility of density-mediated indirect effe
cts. The study illustrates that ecologists should not underestimate th
e importance of behavioral ecology in determining community-level inte
ractions.