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.

Author information


  1. Department of Ecology, Evolution, and Behavior, University of Minnesota, St Paul, Minnesota 55108, USA

    • Elizabeth T. Borer
    • , Eric W. Seabloom
    • , Eric M. Lind
    • , Yann Hautier
    •  & Peter D. Wragg
  2. Department of Entomology, University of Maryland, College Park, Maryland 20742, USA

    • Daniel S. Gruner
  3. Department of Ecology, Evolution, and Organismal Biology, Iowa State University, Ames, Iowa 50011, USA

    • W. Stanley Harpole
    • , Lori Biederman
    • , Wei Li
    • , Brent Mortensen
    • , Lauren L. Sullivan
    •  & Ryan J. Williams
  4. Institute for Chemistry and Biology of the Marine Environment, Carl-von- Ossietzky University, 26382 Wilhelmshaven, Oldenburg, Germany

    • Helmut Hillebrand
  5. Department of Wildland Resources and the Ecology Center, Utah State University, Logan, Utah 84322, USA

    • Peter B. Adler
  6. Instituto de Investigaciones Marinas y Costeras (IIMyC), Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Mar del Plata 7600 , Argentina

    • Juan Alberti
    • , Pedro Daleo
    • , Oscar Iribarne
    •  & Jesús Pascual
  7. Department of Biology, Wake Forest University, Winston-Salem, North Carolina 27109, USA

    • T. Michael Anderson
  8. School of Environmental and Forest Sciences, University of Washington, Seattle, Washington 98195, USA

    • Jonathan D. Bakker
  9. Agricultural Research Service (ARS), United States Department of Agriculture, Fort Collins, Colorado 80526, USA

    • Dana Blumenthal
  10. Deptartment of Forest, Rangeland and Watershed Stewardship, Colorado State University, Fort Collins, Colorado 80523, USA

    • Cynthia S. Brown
    • , Julia A. Klein
    •  & Melinda D. Smith
  11. Department of Plant Biology, Michigan State University, East Lansing, Michigan 48824, USA

    • Lars A. Brudvig
  12. ARC Centre of Excellence for Environmental Decisions, School of Biological Sciences, The University of Queensland, Queensland 4072, Australia

    • Yvonne M. Buckley
  13. School of Natural Sciences, Trinity College Dublin, Dublin 2, Ireland

    • Yvonne M. Buckley
  14. Department of Ecology and Evolutionary Biology, University of Toronto Scarborough, Ontario M1C 1A4, Canada

    • Marc Cadotte
  15. State Key Laboratory of Grassland and Agro-Ecosystems, Research Station of Alpine Meadow and Wetland Ecosystems, School of Life Sciences, Lanzhou University, Lanzhou, 730000 Gansu, China

    • Chengjin Chu
    •  & Guozhen Du
  16. Division of Biological Sciences, University of California, San Diego, California 92093, USA

    • Elsa E. Cleland
  17. Department of Biology, Imperial College at Silwood Park, Ascot, Berkshire SL5 7PY, UK

    • Michael J. Crawley
  18. Department of Zoology, University of Wisconsin, Madison, Wisconsin 53706, USA

    • Ellen I. Damschen
    •  & John L. Orrock
  19. Department of Ecology and Evolutionary Biology, University of Colorado, Boulder Colorado 80309, USA

    • Kendi F. Davies
    •  & Brett A. Melbourne
  20. US Geological Survey, Forest and Rangeland Ecosystem Science Center, Corvallis, Oregon 97331, USA

    • Nicole M. DeCrappeo
    •  & David A. Pyke
  21. Queensland University of Technology, Biogeosciences, Brisbane, Queensland 4001, Australia

    • Jennifer Firn
  22. Department of Biology, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina 27599, USA

    • Robert W. Heckman
    •  & Charles E. Mitchell
  23. Department of Plant Sciences, University of Oxford, Oxford OX1 3RB, UK

    • Andy Hector
  24. School of Environmental and Forest Sciences, University of Washington, Seattle, Washington 98195, USA

    • Janneke HilleRisLambers
  25. School of Biological Sciences, University of Nebraska, Lincoln, Nebraska 68588, USA

    • Johannes M. H. Knops
  26. Berkeley Initiative for Global Change Biology, University of California, Berkeley 94704, USA

    • Kimberly J. La Pierre
  27. Department of Plant Biology, University of Illinois at Urbana-Champaign, llinois 61820, USA

    • Andrew D. B. Leakey
  28. Department of Integrative Biology, University of Guelph, Guelph, Ontario N1G 2W1, Canada

    • Andrew S. MacDougall
  29. Department of Plant & Soil Sciences, University of Kentucky, Lexington, Kentucky 40546, USA

    • Rebecca L. McCulley
  30. Australian Research Center for Urban Ecology, c/o School of Botany, University of Melbourne, Victoria 3010, Australia, and School of Biological Sciences, Monash University, Victoria 3800, Australia

    • Joslin L. Moore
  31. Department of Zoology, Oregon State University, Corvallis, Oregon 97331, USA

    • Lydia R. O'Halloran
  32. CSIRO Ecosystem Sciences, Wembley, West Australia 6913, Australia

    • Suzanne M. Prober
  33. Swiss Federal Institute for Forest, Snow and Landscape Research, Birmensdorf 8903, Switzerland

    • Anita C. Risch
    •  & Martin Schuetz
  34. Lancaster Environment Center, Lancaster University, Lancaster LA1 4YQ, UK

    • Carly J. Stevens
  35. Department of Biology, Duke University, Durham, North Carolina 27708, USA

    • Justin P. Wright
  36. Department of Entomology, University of California, Davis, California 95616, USA

    • Louie H. Yang


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E.T.B., E.W.S., W.S.H. and E.M.L. are Nutrient Network coordinators. E.T.B., W.S.H., H.H. and D.S.G. developed and framed the research questions in this paper. All authors contributed data from this experiment. E.T.B. and E.W.S. analysed the data. D.S.G., W.S.H. and E.M.L. contributed to data analyses. E.T.B. wrote the paper with input from all authors. Supplementary Information Appendix S2 provides further information on author contributions.

Competing interests

The authors declare no competing financial interests.

Corresponding author

Correspondence to Elizabeth T. Borer.

Extended data

Extended data tables

  1. 1.

    Sites contributing experimental data

  2. 2.

    Statistical model for treatment effects on richness after 3 years of treatment (n = 29) as a function of fertilization by N, P and K and micronutrients, herbivore exclusion by fencing, and their interaction

  3. 3.

    Statistical model for treatment effects on biomass after 3 years of treatment (n = 29) as a function of fertilization by N, P and K and micronutrients, herbivore exclusion by fencing, and their interaction

  4. 4.

    Statistical model for treatment effects on proportion of photosynthetically active radiation (PAR) reaching ground level after 3 years of treatment (n = 29) as a function of fertilization by N, P and K and micronutrients, herbivore exclusion by fencing, and their interaction

  5. 5.

    Statistical model for biomass effects on ground-level proportion of photosynthetically active radiation (PAR) after 3 years of treatment (n = 29) as a function of total plot-scale biomass

  6. 6.

    Effects of climate, nitrogen deposition, soil nitrogen and site productivity on change in ground-level light across experimental fencing treatments after 3 years of treatment

  7. 7.

    Effects of climate, nitrogen deposition, soil nitrogen and site productivity on site-level mean biomass change across experimental fencing treatments after 3 years of treatment

  8. 8.

    Effects of climate, nitrogen deposition, soil nitrogen, site productivity, and change in light on change in site-level mean plant species richness across experimental fencing treatments after three years of treatments

Supplementary information

PDF files

  1. 1.

    Supplementary Information

    This file contains Supplementary Appendices S1-S2.


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