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Stan Harpole

Posted by | February 28, 2021


Stan Harpole

Job Title:



Phone Number:

+49 0341 973 3171


Google Scholar Profile


My research focuses on developing, testing, and applying biodiversity theory to understand the interactive effects of multiple global change drivers. My expertise bridges ecosystem, theoretical and community ecology. Specific research topics include ecological stoichiometry, niche theory and mechanisms of coexistence.

Research Areas:

  • Nutrient limitation
  • Biodiversity

Current N-related Projects:


Harpole, W.S., L. Sullivan, E. Lind, J. Firn, P. Adler, E. Borer, J. Chase, P. Fay, Y. Hautier, H. Hillebrand, A.S. MacDougall, E. Seabloom, J.D. Bakker, M. Cadotte, E. Chaneton, C. Chu, N. Hagenah, K. Kirkman, K.J. La Pierre, J.L. Moore, J.W. Morgan, S. Prober, A.C. Risch, M. Schuetz, C. Stevens. 2017. Out of the shadows: multiple nutrient limitation drives relationships between biomass, light, and plant diversity. Functional Ecology. 31:1839-1846. doi 10.1111/1365-2435.12967

Borer, E.T., J.B. Grace, W.S. Harpole, A.S. MacDougall, E.W. Seabloom, 2017. A decade of insights into grassland ecosystem responses to global environmental change. Nature Ecology & Evolution, 1, 0118.

Harpole, W.S., L. Sullivan, E. Lind, J. Firn, P. Adler, E. Borer, J. Chase, P. Fay, Y. Hautier, H. Hillebrand, A.S. MacDougall, E. Seabloom, R. Williams, J.D. Bakker, M. Cadotte, E. Chaneton, C. Chu, E.E. Cleland, C. D’Antonio, K.F. Davies, D.S. Gruner, N. Hagenah, K. Kirkman, J.M.H. Knops, K.J. La Pierre, R.L. McCulley, J.L. Moore, J.W. Morgan, S. Prober, A.C. Risch, M. Schuetz, C. Stevens, P.D. Wragg. 2016. Addition of multiple limiting resources reduces grassland diversity. Nature. 537: 93-96.

Fay, P., S. Prober, W.S. Harpole, J. Knops, J. Bakker, E. Borer, E. Lind, A.S. MacDougall, E. Seabloom, P. Wragg, P. Adler, D. Blumenthal, Y. Buckley, C. Chu, E. Cleland, S. Collins, K. Davies, G. Du, X. Feng, J. Firn, D. Gruner, N. Hagenah, Y. Hautier, R. Heckman, V. Jin, K. Kirkman, J. Klein, L.M. Ladwig, Q. Li, R. McCulley, B. Melbourne, C. Mitchell, J. Moore, J. Morgan, A. Risch, M. Schuetz, C. Stevens, D.A. Wedin, L. Yang. 2015. Grassland Productivity is Limited by Multiple Nutrients. Nature Plant. 1:15080.  Commentary by Peter Vitousek:

Stevens C.J., Eric M. Lind E.L., Hautier Y., Harpole W.S., Borer E.T., Hobbie S., Seabloom E.W., Ladwig L., Bakker J.D., Chu C., Collins S., Davies K.F., Firn J., Hillebrand H., La Pierre K.J., MacDougall A., Melbourne B.,  McCulley R.L., Morgan J., Orrock J.L., Prober S.M., Risch A.C., Schuetz M., Wragg P.D. 2015. Anthropogenic nitrogen deposition predicts local grassland primary production worldwide. Ecology. 96:1459–1465.

Bracken, M. H. Hillebrand; E. Borer, E. Seabloom, J. Cebrian, E. Cleland, J. Elser, D. Gruner, W.S. Harpole, J. Ngai, J. Smith. 2015. Signatures of nutrient limitation and co-limitation: responses of autotroph internal nutrient concentrations to nitrogen and phosphorus additions. Oikos 124:113-121. Editor’s Choice.

Harpole, W.S., J.T. Ngai, E.E. Cleland, E.T. Borer, M.E.S. Bracken, J.J. Elser, D.S. Gruner, H. Hillebrand, E.W. Seabloom, J.B. Shurin, J.E. Smith. 2011. Nutrient co-limitation of primary producer communities. Ecology Letters. 14:852-862. Faculty 1000 Must-Read.

Cleland, E. E., W. S. Harpole. 2010. Nitrogen enrichment and plant communities. Annals of the New York Academy of Sciences. 1195:46-61.

Hille Ris Lambers, J., W.S. Harpole, S. Schnitzer, D. Tilman, P. Reich. 2009. CO2, nitrogen and diversity differentially affect seed production of prairie plants. Ecology 90:1810-1820.

Everard, K., E. Seabloom, W.S. Harpole, C. de Mazancourt. 2009. Plant water use affects competition for nitrogen: why drought favors invasive species in California. American Naturalist. 175:85-97.

Elser, J.J., M.E.S. Bracken, E.E. Cleland, D.S. Gruner, W.S. Harpole, H. Hillebrand, J.T. Ngai, E.W. Seabloom, J.B. Shurin, J.E. Smith. 2007. Global analysis of nitrogen and phosphorus limitation of primary producers in freshwater, marine, and terrestrial ecosystems. Ecology Letters. 10(12):1135-1142. Faculty 1000 Exceptional, Must-Read.

Harpole, W.S., D.L. Potts, K.N. Suding. 2007. Ecosystem responses to water and nitrogen amendment in a California grassland. Global Change Biology. 13:2341–2348.

Harpole, W.S., D. Tilman. 2007. Grassland species loss due to reduced niche dimension. Nature. 446:791-793. Faculty 1000 Must-Read (2).

Harpole, W.S., L. Goldstein, R. Aicher. Resource limitation. 2007. In C. D’Antonio, J. Corbin, and M. Stromberg, editors. Ecology and Management of California Grassland. University of California Press. 119-127.