Herbivores and nutrients control grassland plant diversity via light limitation

Human alterations to nutrient cycles and herbivore communities are affecting global biodiversity dramatically. 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 systems. 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.

Brett A. Melbourne | Peter B. Adler | Janneke HilleRisLambers | Ellen I. Damschen | John L. Orrock | Wei Li | Elizabeth T. Borer | W. Stanley Harpole | Eric M. Lind | Michael J. Crawley | Helmut Hillebrand | Andy Hector | Jennifer Firn | Eric W. Seabloom | Cynthia S. Brown | Lars A. Brudvig | Dana Blumenthal | Marc Cadotte | Yvonne M. Buckley | David A. Pyke | Andrew D. B. Leakey | Anita C. Risch | Jonathan D. Bakker | Kimberly J. La Pierre | Guozhen Du | Lauren L. Sullivan | Louie H. Yang | Carly J. Stevens | Chengjin Chu | E. Borer | T. M. Anderson | S. Prober | Wei Li | M. Crawley | J. HilleRisLambers | P. Adler | J. Klein | M. Cadotte | D. Pyke | J. Firn | Y. Buckley | A. Hector | Melinda D. Smith | H. Hillebrand | W. Harpole | B. Melbourne | Joslin L. Moore | C. Stevens | E. Cleland | Justin P. Wright | C. Mitchell | A. Leakey | D. Gruner | D. Blumenthal | R. McCulley | J. Knops | E. Seabloom | A. Risch | Ryan J. Williams | K. Davies | Y. Hautier | J. Orrock | O. Iribarne | L. Sullivan | P. Daleo | E. Lind | J. Bakker | E. Damschen | A. MacDougall | L. Brudvig | G. Du | L. Biederman | J. Alberti | P. D. Wragg | Chengjin Chu | N. DeCrappeo | R. Heckman | K. J. L. Pierre | B. Mortensen | Lydia R. O’Halloran | J. Pascual | M. Schuetz | Johannes M. H. Knops | Suzanne M. Prober | Elsa E. Cleland | Andrew S. MacDougall | Kendi F. Davies | T. Michael Anderson | Oscar Iribarne | Lydia R. O'Halloran | Peter D. Wragg | Daniel S. Gruner | Rebecca L. McCulley | Nicole M. DeCrappeo | Yann Hautier | Julia A. Klein | Martin Schuetz | Brent Mortensen | Charles E. Mitchell | Lori Biederman | Juan Alberti | Pedro Daleo | Robert W. Heckman | Jesús Pascual | A. Hector | J. H. Knops | J. Wright | J. M H Knops | Lauren L. Sullivan | K. J. Pierre | W. StanleyHarpole | HelmutHillebrand

[1]  A. Schrijver,et al.  Cumulative nitrogen input drives species loss in terrestrial ecosystems , 2011 .

[2]  D. Milchunas,et al.  Quantitative Effects of Grazing on Vegetation and Soils Over a Global Range of Environments , 1993 .

[3]  A. Mazumder,et al.  REVERSAL OF GRAZING IMPACT ON PLANT SPECIES RICHNESS IN NUTRIENT‐POOR VS. NUTRIENT‐RICH ECOSYSTEMS , 1998 .

[4]  H. Olff,et al.  Herbivore impact on grassland plant diversity depends on habitat productivity and herbivore size. , 2006, Ecology letters.

[5]  Andy Hector,et al.  Competition for Light Causes Plant Biodiversity Loss After Eutrophication , 2009, Science.

[6]  J. P. Grime,et al.  The Evolutionary Strategies that Shape Ecosystems: Grime/The Evolutionary Strategies that Shape Ecosystems , 2012 .

[7]  Helmut Hillebrand,et al.  Consumer versus resource control of producer diversity depends on ecosystem type and producer community structure , 2007, Proceedings of the National Academy of Sciences.

[8]  F. Stuart Chapin,et al.  Resource Availability and Plant Antiherbivore Defense , 1985, Science.

[9]  Navin Ramankutty,et al.  Our share of the planetary pie , 2007, Proceedings of the National Academy of Sciences.

[10]  J. P. Grime,et al.  The Evolutionary Strategies that Shape Ecosystems , 2012 .

[11]  I. Jamieson,et al.  Multimodel inference in ecology and evolution: challenges and solutions , 2011, Journal of evolutionary biology.

[12]  Peter B. Adler,et al.  Finding generality in ecology: a model for globally distributed experiments , 2014 .

[13]  S. Díaz,et al.  Biodiversity Loss Threatens Human Well-Being , 2006, PLoS biology.

[14]  Franz J. Weissing,et al.  Growth and competition in a light gradient , 1994 .

[15]  E. Newman,et al.  Competition and Diversity in Herbaceous Vegetation , 1973, Nature.

[16]  F. Chapin,et al.  A safe operating space for humanity , 2009, Nature.

[17]  D. Tilman,et al.  Resource Use Patterns Predict Long‐Term Outcomes of Plant Competition for Nutrients and Light , 2007, The American Naturalist.

[18]  James P. Grover,et al.  Simple Rules for Interspecific Dominance in Systems with Exploitative and Apparent Competition , 1994, The American Naturalist.

[19]  W. Stanley Harpole,et al.  Grassland species loss resulting from reduced niche dimension , 2007, Nature.

[20]  A. Wilkes,et al.  Review of evidence on drylands pastoral systems and climate change: implications and opportunities for mitigation and adaptation. , 2009 .

[21]  P. Chesson Mechanisms of Maintenance of Species Diversity , 2000 .

[22]  David J. Gibson,et al.  Grasses and Grassland Ecology , 2009 .

[23]  H. Olff,et al.  Effects of herbivores on grassland plant diversity. , 1998, Trends in ecology & evolution.

[24]  Helmut Hillebrand,et al.  Life-history constraints in grassland plant species: a growth-defence trade-off is the norm. , 2013, Ecology letters.

[25]  Rebecca E. Irwin,et al.  Direct and ecological costs of resistance to herbivory , 2002 .

[26]  Ulrich Sommer,et al.  Consumer versus resource control of species diversity and ecosystem functioning , 2002, Nature.

[27]  Franz J. Weissing,et al.  Light-limited growth and competition for light in well-mixed aquatic environments : An elementary model , 1994 .

[28]  J. L. Parra,et al.  Very high resolution interpolated climate surfaces for global land areas , 2005 .

[29]  S. Carpenter,et al.  Solutions for a cultivated planet , 2011, Nature.

[30]  Peter H. Verburg,et al.  Projecting land use changes in the Neotropics: the geography of pasture expansion into forest , 2007 .