Human alterations to nutrient cycles1,2 and herbivore communities3,4,5,6,7 are affecting global biodiversity dramatically2. Ecological theory predicts these changes should be strongly counteractive: nutrient addition drives plant species loss through intensified competition for light, whereas herbivores prevent competitive exclusion by increasing ground-level light, particularly in productive systems8,9. Here we use experimental data spanning a globally relevant range of conditions to test the hypothesis that herbaceous plant species losses caused by eutrophication may be offset by increased light availability due to herbivory. This experiment, replicated in 40 grasslands on 6 continents, demonstrates that nutrients and herbivores can serve as counteracting forces to control local plant diversity through light limitation, independent of site productivity, soil nitrogen, herbivore type and climate. Nutrient addition consistently reduced local diversity through light limitation, and herbivory rescued diversity at sites where it alleviated light limitation. Thus, species loss from anthropogenic eutrophication can be ameliorated in grasslands where herbivory increases ground-level light.
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This work uses data from the Nutrient Network (http://nutnet.org) experiment, funded at the site scale by individual researchers. Coordination and data management are supported by funding to E. Borer and E. Seabloom from the NSF Research Coordination Network (NSF-DEB-1042132) and Long Term Ecological Research (NSF-DEB-1234162 to Cedar Creek LTER) programs and the UMN Institute on the Environment (DG-0001-13). The Minnesota Supercomputer Institute hosts project data. We are grateful to F. Isbell for suggestions that improved the manuscript. Any use of trade names is for descriptive purposes only and does not imply endorsement by the US Government.
The authors declare no competing financial interests.
Extended data figures and tables
a–c, All available data are shown for richness (a), total biomass (b) and ambient light reaching ground level (c). Error bars represent ±2 s.e.m. Treatment years and their associated sample sizes are shown in each panel. One- and two-year models represent greater spatial extent and replication, but reduced temporal extent compared to Fig. 2 in the main text. Four-year models represent longer temporal effects, but reduced spatial extent, particularly for light measurements. All models were fitted as in Extended Data Tables 2, 3, 4 and described in the Methods.
Extended Data Figure 2 Fertilization does not alter the relationship between ‘fence’ effects on light and diversity.
The log response ratio (LRR) model of the effect of fences (herbivore exclusion) on richness and light (year 3 data) demonstrates no additional effect of nutrient addition on the relationship shown in Fig. 3. The grey region indicates the 95% confidence interval around the regression. The effect of fences on ground-level light predicts changes in plot-scale species richness (P = 0.00254), whereas fertilization is not included in the final statistical model of this relationship (P > 0.05). Thus, the magnitude of the effect of grazers on richness is dependent on the magnitude of their effect on light regardless of whether a plot has been fertilized.
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Borer, E., Seabloom, E., Gruner, D. et al. Herbivores and nutrients control grassland plant diversity via light limitation. Nature 508, 517–520 (2014). https://doi.org/10.1038/nature13144
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