Prof. Dr. W. Stanley Harpole Prof. Stan Harpole     Prof. Stan Harpole


Department for Physiological Diversity
German Centre for Integrative Biodiversity Research (iDiv)
Puschstraße 4
04103 Leipzig

phone: +49 341 973 3171 / +49 341 6025 3387 (Zoom Voice)

Highly Cited Researcher

Google Scholar


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.

  • Helmholtz-Center for Environmental Research (UFZ)
  • German Centre for Integrative Biodiversity Research (iDiv)
  • Martin-Luther-Universität-Halle-Wittenberg (MLU)

2014-2014 Professor, Department Head Physiological Diversity


Weiterführende Recherchen können Sie in unserem Publikationsverzeichnis durchführen.

2024 (5)

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2023 (10)

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2022 (13)

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2021 (10)

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2020 (13)

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2019 (8)

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2018 (8)

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2017 (4)

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2016 (10)

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2015 (6)

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  • 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.
  • Borer, E., E. Seabloom, D. Gruner, W.S. Harpole, H. Hillebrand, E. Lind, P. Adler, J. Alberti, T. Anderson, J. Bakker, L. Biederman, D. Blumenthal, C. Brown, L. Brudvig, Y. Buckley, M. Cadotte, C. Chu, E. Cleland, M. Crawley, P. Daleo, E. Damschen, K. Davies, N. DeCrappeo, G. Du, J. Firn, Y. Hautier, R. Heckman, A. Hector, J. HilleRisLambers, O. Iribarne, J. Klein, J. Knops, K. La Pierre, A. Leakey, W. Li, A. MacDougall, R. McCulley, B. Melbourne, C. Mitchell, J. Moore, B. Mortensen, L. O'Halloran, J. Orrock, L. Pascual, S. Prober, D. Pyke, A. Risch, M. Schuetz, M. Smith, C. Stevens, L. Sullivan, R. Williams, P. Wragg, J. Wright, L. Yang. 2014. Herbivores and nutrients control grassland plant diversity via light limitation. Nature 508:517-520.
  • Grace J.B., P.B. Adler, W.S. Harpole, E.T. Borer, E.W. Seabloom. 2014. Causal networks clarify productivity-richness interrelations, bivariate plots do not. Functional Ecology. 28 (4), 787-798.
  • MacDougall A. S., J. R. Bennett, J. Firn, E. Seabloom, E. Borer, E. Lind, P. Adler, M. Anderson, E. Cleland, S. L. Collins, K. Davies, J. B. Grace, W. S. Harpole, Y. Hautier, A. Kendig, B. Melbourne, J. Orrock. 2014. Anthropogenic-based regional-scale factors most consistently explain plot-level exotic diversity in grasslands on two continents. Global Ecology and Biogeography.
  • Hautier, Y., E. Seabloom, E. Borer, P. Adler, W.S. Harpole, H. Hillebrand, E. Lind, A. MacDougall, C. Stevens, J. Bakker, Y. Buckley, C. Chu, S. Collins, P. Daleo, E. Damschen, K. Davies, P. Fay, J. Firn, D. Gruner, V. Jin, J. Klein, J. Knops, K. La Pierre, W. Li, R. McCulley, B. Melbourne, J. Moore, L. O'Halloran, S. Prober, A. Risch, M. Sankaran, M. Schuetz, A. Hector. 2014. Eutrophication weakens stabilizing effects of diversity in natural grasslands. Nature 508:521-525.
  • Stein, C., L.M. Hallett, W.S. Harpole, K.N. Suding. 2014. Evaluating ecosystem services provided by non-native species: An experimental test in California grasslands. PLoS One 9:9, e75396.
  • Filstrup, C.T., H. Hillebrand, A. Heathcote, W.S. Harpole, J.A. Downing. 2014. Cyanobacteria dominance influences resource use efficiency and community turnover in phytoplankton and zooplankton communities. Ecology Letters 17:464-474.
  • Borer, E., W.S. Harpole, P. Adler, E. Lind, J. Orrock, E. Seabloom, M. Smith. 2014. Finding generality in ecology: A model for globally distributed experiments. Methods in Ecology and Evolution 5:65-73.
  • Harpole, W.S. & L.A. Biederman. 2014. On the importance of accurate reporting: a response to comments on “Biochar and its effects on plant productivity and nutrient cycling: a meta-analysis”. Global Change Biology-Bioenergy 6:172-175.
  • Borer, E.T., M.E.S. Bracken, E.W. Seabloom, J.E. Smith., J Cebrian, E.E. Cleland, J.J. Elser, W. Fagan, D.S. Gruner, W.S. Harpole, H. Hillebrand, A. Kerkoff, J.T. Ngai, J.B. Shurin. 2013. Global biogeography of autotroph chemistry: is insolation a driving force? Oikos. 122:1121–1130.
  • Seabloom, E.W., E.T. Borer, Y. Buckley, E. Cleland, K. Davies, J. Firn, S. Harpole, Y. Hautier, E. Lind, A. MacDougall, J.L. Orrock, S.M. Prober, P. Adler, J. Alberti, T.M. Anderson, J.D. Bakker, L.A. Biederman, D. Blumenthal, C.S. Brown, L.A. Brudvig, M. Caldeira, C. Chu, M.J. Crawley, P. Daleo, E.I. Damschen, C.M. Dantonio, N.M. DeCrappeo, C.R. Dickman, G. Du, P.A. Fay, P. Frater, D.S. Gruner, N. Hagenah, A. Hector, A. Helm, H. Hillebrand, K.S. Hofmockel, H.C. Humphries, O. Iribarne, V.L. Jin, A. Kay, K.P. Kirkman, J.A. Klein, J.M.H. Knops, K.J. La Pierre, L.M. Ladwig, J.G. Lambrinos, A.D.B. Leakey, Q. Li, W. Li, R. McCulley, B. Melbourne, C. Mitchell, J.L. Moore, J. Morgan, B. Mortensen, L.R. O'Halloran, M. Pärtel, J. Pascual, D. Pyke, A.C. Risch, R. Salguero-Gómez, M. Sankaran, M. Schuetz, A. Simonsen, M. Smith, C. Stevens, L. Sullivan, G.M. Wardle, E.M. Wolkovich, P.D. Wragg, J. Wright, L. Yang. 2013. Predicting invasion in grassland ecosystems: is exotic dominance the real embarrassment of richness? Global Change Biology 19:3677-3687.
  • Lind E.M., E. Borer, E. Seabloom, P. Adler, J.D. Bakker, D. Blumenthal, M. Crawley, K. Davies, J. Firn, D.S. Gruner, W.S. Harpole, Y. Hautier, H. Hillebrand, J. Knops, B. Melbourne, B. Mortensen, A.C. Risch, M. Schuetz, C. Stevens, P.D. Wragg. 2013. Life history constraints in grassland plant species: a growth-defense trade-off is the norm. Ecology Letters. 16:513–521.
  • O’Halloran L.R., E.T. Borer, E.W. Seabloom, A.S. MacDougall, E. E. Cleland, R.L. McCulley, S. Hobbie, W.S. Harpole, N.M. DeCrappeo, C. Chu, J.D. Bakker, K.F. Davies, G. Du, J. Firn, N. Hagenah, K. S. Hofmockel, J.M.H. Knops, W. Li, B.A. Melbourne, J.W. Morgan, J.L. Orrock, S.M. Prober, and C.J. Stevens. 2013. Regional contingencies in the relationship between aboveground biomass and litter in the world’s grasslands. PLoS ONE. 8(2): e54988
  • Suding K.N., W.S. Harpole, T. Fukami, A. Kulmatiski, A.S. MacDougall A.S., C. Stein, W.H. van der Putten, M. Hutchings M. 2013. Consequences of plant-soil feedbacks in invasion. Journal of Ecology. 101:298-308.
  • Biederman L., W.S. Harpole. 2013. Biochar and its effects on plant productivity and nutrient cycling: a meta-analysis. Global Change Biology–Bioenergy. 5:202–214. Most Downloaded Article in 2014 & 2015and most cited article published in GCB Bioenergy in 2013, 2014.
  • Hille Ris Lambers, J., P.B. Adler, W.S. Harpole, J.M. Levine, M.M. Mayfield. 2012. Rethinking Community Assembly through the Lens of Coexistence Theory. Annual Review of Ecology, Evolution, and Systematics. 43:227-248
  • Grace J.B., Adler P.B., Seabloom E.W., Borer E.T., Hillebrand H., Hautier Y., Hector A., Harpole W.S., O'Halloran L.R., Anderson T.M., Bakker J.D., Brown C.S., Buckley Y.M., Collins S.L., Cottingham K.L., Crawley M.J., Damschen E.I., Davies K.F., DeCrappeo N.M., Fay P.A., Firn J., Gruner D.S., Hagenah N., Jin V.L., Kirkman K.P., Knops J.M.H., La Pierre K.J., Lambrinos J.G., Melbourne B.A., Mitchell C.E., Moore J.L., Morgan J.W., Orrock J.L., Prober S.M., Stevens C.J., Wragg P.D. & Yang L.H. 2012. Response to Comments on “Productivity Is a Poor Predictor of Plant Species Richness.” Science. 335:1441.
  • Adler, P., Seabloom E.W., Borer E.T., Hillebrand H., Hautier Y., Hector A., Harpole W.S., O'Halloran L.R., Grace J.B., Anderson T.M., Bakker J.D., Biederman L.A., Brown C.S., Buckley Y.M., Calabrese L.B., Chu C.J., Cleland E.E., Collins S.L., Cottingham K.L., Crawley M.J., Damschen E.I., Davies K.F., DeCrappeo N.M., Fay P.A., Firn J., Frater P., Gasarch E.I., Gruner D.S., Hagenah N., Hille Ris Lambers J., Humphries H., Jin V.L., Kay A.D., Kirkman K.P., Klein J.A., Knops J.M., La Pierre K.J., Lambrinos J.G., Li W., Macdougall A.S., McCulley R.L., Melbourne B.A., Mitchell C.E., Moore J.L., Morgan J.W., Mortensen B., Orrock J.L., Prober S.M., Pyke D.A., Risch A.C., Schuetz M., Smith M.D., Stevens C.J., Sullivan L.L., Wang G., Wragg P.D., Wright J.P. & Yang L.H. 2012. Productivity is a poor predictor of plant species richness. Science. 333:1750-1753. Faculty 1000 Must-Read.
  • 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.
  • Isbell, F, V. Calcagno, A. Hector, J. Connolly, W. S. Harpole, P. B. Reich, M. Scherer-Lorenzen, B. Schmid, D. Tilman, J. van Ruijven, A. Weigelt, B. J. Wilsey, E. S. Zavaleta, M. Loreau. 2011. How many species are needed to maintain ecosystem functioning? Nature. 477:199-202.
  • Firn, J., J. Moore, A.S. MacDougall, E. Borer, E. Seabloom, J. Hille Ris Lambers, W.S. Harpole, E. Cleland, C. Brown, J. Knops, S. Prober, D. Pyke, K. Farell, J. Bakker, L. Ries-O’Halloran, Y.M. Buckley. 2011. Species abundance at home predicts abundance away in grasslands. Ecology Letters. 14:274-281. Faculty 1000 Must-Read.
  • Harpole, W.S. and K.N. Suding. 2011. A test of the niche dimension hypothesis in an arid annual grassland. Oecologia. 166:197-205.
  • Cadotte, M.W., E.T. Borer, E.W. Seabloom, J. Cavender-Bares, W.S. Harpole, E. Cleland, K. Davies. 2010. Phylogenetic patterns differ for native and exotic plant communities across a richness gradient in Northern California. Diversity and Distributions. 16:892-901.
  • Harpole, W.S. 2010. Neutral theory of species diversity. Nature Education Knowledge 1(8):31. (Invited submission).
  • Cleland, E. E., W. S. Harpole. 2010. Nitrogen enrichment and plant communities. Annals of the New York Academy of Sciences. 1195:46-61.
  • La Pierre, K.J., W.S. Harpole, K.N. Suding. 2010. Strong feeding preference of an exotic generalist herbivore for an exotic forb: a case against invasional antagonism. Biological Invasions. 12:3025-3031
  • 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.
  • Cardinale, B.J., H. Hillebrand, W.S. Harpole, K. Gross, R. Ptacnik. 2009. Separating the influence of resource ‘availability’ from resource ‘imbalance’ on productivity–diversity relationships. Ecology Letters. 12: 475-487.
  • Hillebrand, H., E.T. Borer, M.E. S. Bracken, B.J. Cardinale, J. Cebrian, E.E. Cleland, J.J. Elser, D.S. Gruner, W.S. Harpole, J.T. Ngai, S. Sandin, E.W. Seabloom, J.B. Shurin, J.E. Smith, M.D. Smith. 2009. Herbivore metabolism and stoichiometry constrain herbivory at different organizational scales across ecosystems. Ecology Letters. 12: 516-527.
  •  Gruner, D.S., J.E. Smith, E.W. Seabloom, S.A. Sandin, J.T. Ngai, H. Hillebrand, W.S. Harpole, J.J. Elser, E.E. Cleland, M.E.S. Bracken, E.T. Borer, B.M. Bolker. 2008. A cross-system synthesis of consumer and nutrient resource control on producer biomass. Ecology Letters. 11:740-755.

  • Potts, D.L., W.S. Harpole, M.L. Goulden, K.N. Suding. 2008. The impact of invasion and subsequent removal of an exotic thistle, Cynara cardunculus, on CO2 and H2O vapor exchange in a coastal California grassland. Biological Invasions. 10:1073-1084.
  • Fox, J.W., and W.S. Harpole. 2008. Revealing how species loss affects ecosystem function: the trait-based Price equation partition. Ecology. 89(1): 269-279.
  • Harpole, W.S. and K.N. Suding. 2007. Frequency-dependence stabilizes competitive interactions among four annual plants. Ecology Letters. 10(12):1164-1169.
  • 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.
  • Hillebrand, H., D.S. Gruner, Borer, E.T., M.E.S. Bracken, E.E. Cleland, J.J. Elser, W.S. Harpole, J.T. Ngai, E.W. Seabloom, J.B. Shurin, J.E. Smith. 2007. Consumer versus resource control of producer diversity depends on ecosystem type and producer community structure. Proceedings of the National Academy of Sciences. 104(26):10904-10909. Faculty 1000 Recommended.
  • Fargione, J., D. Tilman, R. Dybzinski, J. Hille Ris Lambers, C. Clark, W.S. Harpole, J. M. H. Knops, P. B. Reich, M. Loreau. 2007. From selection to complementarity: Shifts in the causes of biodiversity-productivity relationships in a long-term biodiversity experiment. Proceedings of the Royal Society B. 274:871–876.
  • Harpole, W.S., D. Tilman. 2006. Non-neutral patterns of species abundance and species traits. Ecology Letters. 9:15-23. Faculty 1000 Must-Read, Recommended.
  • Harpole, W.S. 2006. Resource-ratio theory and the control of invasive plants. Plant and Soil. 280:23-27.
  • Moe, S.J., R.S. Stelzer, M.R. Forman, W.S. Harpole, T. Daufresne, T. Yoshida. 2005. Recent advances in ecological stoichiometry: insights for population and community ecology. Oikos. 109:29-39.
  • Hille Ris Lambers, J., W.S. Harpole, D. Tilman, J. Knops, P. Reich. 2004. Species and mechanisms contributing to the positive diversity-productivity relationship. Ecology Letters. 7:661-668.
  • Tilman, G.D., J. Hille Ris Lambers, S. Harpole, R. Dybzinski, J. Fargione, C. Clark, C. Lehman. 2004. Does metabolic theory apply to community ecology? It's a matter of scale. Ecology. 85:1797-1799.
  • Seabloom, E., W.S. Harpole, O.J. Reichman, D. Tilman. 2003. Resource competition, seed-limitation, and invasion in California grasslands. Proceedings of the National Academy of Sciences. 100:13384-13389.
  • Parker, I.M., W.S. Harpole, D. Dionne. 1997. Plant community diversity and invasion of the exotic shrub Cytisus scoparius: Testing hypotheses of invasibility and impact. pp. 149-162, in P.V. Dunn and K. Ewing, eds., Ecology and Conservation of the South Puget Sound Prairie Landscape. The Nature Conservancy Press, Seattle.