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    Foraging by fearful frugivores: combined effect of fruit ripening and predation risk

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    1. Plant defensive compounds and predation risk are main determinants of herbivore foraging, though empirical studies have seldom measured the combined effects of these two factors. By considering the interaction between the herb Helleborus foetidus and its main fruit and seed predator, the Wood Mouse Apodemus sylvaticus, we evaluated whether the defensive role against seed predators of compounds present in H. foetidus unripe fruits holds across a micro-landscape that differs in foraging costs (i.e. predation risk). 2. First, we used standardized food patches that simulated fruiting H. foetidus plants to ascertain fruit preferences of captive mice. Then, by means of field experiments, we assessed the combined effects of fruit ripening and predation risk on foraging by free- ranging mice. 3. Captive mice avoided plants with unripe fruit and avoided consuming unripe fruits within a particular plant. Free-ranging mice also avoided unripe fruits in safe micro- habitats (rocky substrate), but not in risky microhabitats (bare ground) where few fruits were consumed. This unexpected result may be driven by predation risk experienced by mice foraging on H. foetidus fruits, and/or plant defensive compounds acting in a dose- dependent manner. 4. Frugivorous mice responded to both chemical defences present in unripe H. foetidus fruits as well as to predation cost though such response was sequential. Plant defence compounds appeared to play a part in mouse foraging only after mice selected low predation risk microhabitats. 5. Our study indicates that both digestive and ecological factors influence foraging decisions, which in turn affects pressures exerted by herbivores on plant populationsPeer reviewe
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