Prairie forb response to timing of vole herbivory.

The timing of herbivory can be an important factor in the strength and direction of plant response to herbivore damage. To determine the effect of vole herbivory timing within a growing season on tallgrass prairie forbs, we used individual plant enclosures to limit vole access to three species, Desmanthus illinoensis, Echinacea purpurea, and Heliopsis helianthoides, in an experimental restoration in northern Illinois, USA. As part of a long-term experiment, we implemented five vole access treatments in 2003: (1) vole access for the entire growing season, (2) early-season access, (3) mid-season access, (4) late-season access, and (5) no vole access. We protected all plants from herbivory in the following growing season (2004) to test whether the effects of herbivory in one growing season carried over to the next. We also tested how restoration planting design, including seeding time (June or December) and density (35 or 350 seeds/m2 of each species) affected patterns of herbivory and plant recovery. Vole access for the entire growing season was most detrimental for the growth and reproduction of all three species. In contrast, vole access for a portion of the growing season had different effects on the three species: Desmanthus growth and reproduction was negatively affected by early-season access, Echinacea reproductive output was reduced by late-season access, and Heliopsis was not affected by early-, mid-, or late-season vole access. Negative effects of continual vole access carried over to the following growing season for Desmanthus and Heliopsis, but not for Echinacea. Effects of herbivory did not carry over to the next season for Echinacea and Heliopsis when plants were accessible to voles for only part of the growing season. In contrast, Desmanthus plants exposed to early-season herbivory in one year continued to produce fewer seeds per plant after being protected from vole herbivory for a growing season. Planting density and planting season had mixed effects. Echinacea and Desmanthus were larger in plots planted in June, showing that restoration design continued to affect plant population dynamics seven years after seeding. However, there was no interaction between plant response to vole herbivory and restoration design.