Defoliation effects on reproductive biomass: Importance of scale and timing

Community-level (per unit area) and individual tiller reproductive biomass inside and outside of long-term exclosures on the northern winter range of Yellowstone National Park, USA were compared. Grazed areas had twice the number of reproductive tillers m^-2 (186 compared to 88 tillers m ^-2), and greater total reproductive biomass m^-2 than ungrazed plots (13 compared to 7 g m^-2). In contrast, seed number tiller^-1 was greater for grasses in exclosures. Because of these offsetting responses, seed production (no. m^-2) was unaffected by herbivores. On an area basis, grazed grasses allocated proportionally more biomass to reproduction (reproductive biomass/above-ground biomass) than ungrazed grasses. We propose that altered plant demography and morphology following defoliation explain how grazers might increase the allocation of biomass to reproduction in Yellowstone grasslands. To understand these results in light of ecological and agronomic studies, we reviewed literature from 118 sources that reported the effects of defoliation on the production of reproductive biomass. The review suggested that the results of herbivory or defoliation on plant reproductive biomass depended on the scale of measurement (community vs. plant). In addition, timing of grazing or defoliation emerged as a key factor that determined whether sexual reproduction was inhibited. Like the early season grazing that is typical of Yellowstone's northern winter range, studies often showed that early season defoliation stimulated production of community-level reproductive biomass. Our results rectify disagreements in the literature that ultimately derive from differences in either timing of defoliation or measurement scale.