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


Department for Physiological Diversity
German Centre for Integrative Biodiversity Research (iDiv)
Deutscher Platz 5a
D-04103 Leipzig

phone: +49 341 973 3171

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


2019 (2)
2018 (7)
2017 (4)
2016 (10)
2015 (6)

You could use our publication index for further requests.

2019 (2)

  • Firn, J., McGree, J.M., Harvey, E., Flores-Moreno, H., Schütz, M., Buckley, Y.M., Borer, E.T., Seabloom, E.W., La Pierre, K.J., MacDougall, A.M., Prober, S.M., Stevens, C.J., Sullivan, L.L., Porter, E., Ladouceur, E., Allen, C., Moromizato, K.H., Morgan, J.W., Harpole, W.S., Hautier, Y., Eisenhauer, N., Wright, J.P., Adler, P.B., Arnillas, C.A., Bakker, J.D., Biederman, L., Broadbent, A.A.D., Brown, C.S., Bugalho, M.N., Caldeira, M.C., Cleland, E.E., Ebeling, A., Fay, P.A., Hagenah, N., Kleinhesselink, A.R., Mitchell, R., Moore, J.L., Nogueira, C., Peri, P.L., Roscher, C., Smith, M.D., Wragg, P.D., Risch, A.C., (2019):
    Leaf nutrients, not specific leaf area, are consistent indicators of elevated nutrient inputs
    Nat. Ecol. Evol. 3 , 400 - 406
    full text (url)
  • Weise, H., Auge, H., Baessler, C., Bärlund, I., Bennett, E.M., Berger, U., Bohn, F., Bonn, A., Borchardt, D., Brand, F., Chatzinotas, A., Corstanje, R., De Laender, F., Dietrich, P., Dunker, S., Durka, W., Fazey, I., Groeneveld, J., Guilbaud, C.S.E., Harms, H., Harpole, S., Harris, J.A., Jax, K., Jeltsch, F., Johst, K., Joshi, J., Klotz, S., Kühn, I., Kuhlicke, C., Müller, B., Radchuk, V., Reuter, H., Rinke, K., Schmitt-Jansen, M., Seppelt, R., Singer, A.S., Standish, R.J., Thulke, H.-H., Tietjen, B., Weitere, M., Wirth, C., Wolf, C., Grimm, V., (2019):
    Resilience trinity: safeguarding ecosystem services across three different time horizons and decision contexts [bioRxiv,]
    full text (url)
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2018 (7)

  • Anderson, T.M., Griffith, D.M., Grace, J.B., Lind, E.M., Adler, P.B., Biederman, L.A., Blumenthal, D.M., Daleo, P., Firn, J., Hagenah, N., Harpole, W.S., MacDougall, A.S., McCulley, R.L., Prober, S.M., Risch, A.C., Sankaran, M., Schuetz, M., Seabloom, E.W., Stevens, C.J., Sullivan, L.L., Wragg, P.D., Borer, E.T., (2018):
    Herbivory and eutrophication mediate grassland plant nutrient responses across a global climatic gradient
    Ecology 99 (4), 822 - 831
    full text (url)
  • Bannar-Martin, K.H., Kremer, C.T., Ernest, S.K.M., Leibold, M.A., Auge, H., Chase, J., Declerck, S.A.J., Eisenhauer, N., Harpole, S., Hillebrand, H., Isbell, F., Koffel, T., Larsen, S., Narwani, A., Petermann, J.S., Roscher, C., Cabral, J.S., Supp, S.R., (2018):
    Integrating community assembly and biodiversity to better understand ecosystem function: the Community Assembly and the Functioning of Ecosystems (CAFE) approach
    Ecol. Lett. 21 (2), 167 - 180
    full text (url)
  • Frater, P.N., Borer, E.T., Fay, P.A., Jin, V., Knaeble, B., Seabloom, E., Sullivan, L., Wedin, D.A., Harpole, W.S., (2018):
    Nutrients and environment influence arbuscular mycorrhizal colonization both independently and interactively in Schizachyrium scoparium
    Plant Soil 425 (1–2), 493 - 506
    full text (url)
  • Hautier, Y., Isbell, F., Borer, E.T., Seabloom, E.W., Harpole, W.S., Lind, E.M., MacDougall, A.S., Stevens, C.J., Adler, P.B., Alberti, J., Bakker, J.D., Brudvig, L.A., Buckley, Y.M., Cadotte, M., Caldeira, M.C., Chaneton, E.J., Chu, C., Daleo, P., Dickman, C.R., Dwyer, J.M., Eskelinen, A., Fay, P.A., Firn, J., Hagenah, N., Hillebrand, H., Iribarne, O., Kirkman, K.P., Knops, J.M.H., Pierre, K.J.L., McCulley, R.L., Morgan, J.W., Pärtel, M., Pascual, J., Price, J.N., Prober, S.M., Risch, A.C., Sankaran, M., Schuetz, M., Standish, R.J., Virtanen, R., Wardle, G.M., Yahdjian, L., Hector, A., (2018):
    Local loss and spatial homogenization of plant diversity reduce ecosystem multifunctionality
    Nat. Ecol. Evol. 2 (1), 50 - 56
    full text (url)
  • Hillebrand, H., Blasius, B., Borer, E.T., Chase, J.M., Downing, J.A., Eriksson, B.K., Filstrup, C.T., Harpole, W.S., Larsen, S., Lewandowska, A.M., Seabloom, E.W., Van de Waal, D.B., Ryabov, A.B., (2018):
    Biodiversity change is uncoupled from species richness trends: Consequences for conservation and monitoring
    J. Appl. Ecol. 55 (1), 169 - 184
    full text (url)
  • Hodapp, , Borer, E.T., Harpole, W.S., Lind, E.M., Seabloom, E.W., Adler, P.B., Alberti, J., Arnillas, C.A., Bakker, J.D., Biederman, L., Cadotte, M., Cleland, E.E., Collins, S., Fay, P.A., Firn, J., Hagenah, N., Hautier, Y., Iribarne, O., Knops, J.M.H., McCulley, R.L., MacDougall, A., Moore, J.L., Morgan, J.W., Mortensen, B., La Pierre, K.J., Risch, A.C., Schütz, M., Peri, P., Stevens, C.J., Wright, J., Hillebrand, H., (2018):
    Spatial heterogeneity in species composition constrains plant community responses to herbivory and fertilisation
    Ecol. Lett. 21 (9), 1364 - 1371
    full text (url)
  • Mortensen, B., Danielson, B., Harpole, W.S., Alberti, J., Arnillas, C.A., Biederman, L., Borer, E.T., Cadotte, M.W., Dwyer, J.M., Hagenah, N., Hautier, Y., Peri, P.L., Seabloom, E.W., (2018):
    Herbivores safeguard plant diversity by reducing variability in dominance
    J. Ecol. 106 (1), 101 - 112
    full text (url)
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2017 (4)

  • Biederman, L.A., Phelps, J., Ross, B.J., Polzin, M., Harpole, W.S., (2017):
    Biochar and manure alter few aspects of praire development: A field test
    Agric. Ecosyst. Environ. 236 , 78 - 87
    full text (url)
  • Borer, E.T., Grace, J.B., Harpole, W.S., MacDougall, A.S., Seabloom, E.W., (2017):
    A decade of insights into grassland ecosystem responses to global environmental change
    Nat. Ecol. Evol. 1 (5), art. 0118
    full text (url)
  • Ceulemans, T., Bodé, S., Bollyn, J., Harpole, S., Coorevits, K., Peeters, G., Van Acker, K., Smolders, E., Boeckx, P., Honnay, O., (2017):
    Phosphorus resource partitioning shapes phosphorus acquisition and plant species abundance in grasslands
    Nat. Plants 3 , art. 16224
    full text (url)
  • Harpole, W.S., Sullivan, L.L., Lind, E.M., Firn, J., Adler, P.B., Borer, E.T., Chase, J., Fay, P.A., Hautier, Y., Hillebrand, H., MacDougall, A.S., Seabloom, E.W., Bakker, J.D., Cadotte, M.W., Chaneton, E.J., Chu, C., Hagenah, N., Kirkman, K., La Pierre, K.J., Moore, J.L., Morgan, J.W., Prober, S.M., Risch, A.C., Schuetz, M., Stevens, C.J., (2017):
    Out of the shadows: multiple nutrient limitations drive relationships among biomass, light and plant diversity
    Funct. Ecol. 31 (9), 1839 - 1846
    full text (url)
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2016 (10)

  • Dunker, S., Nadrowski, K., Jakob, T., Kasprzak, P., Becker, A., Langner, U., Kunath, C., Harpole, S., Wilhelm, C., (2016):
    Assessing in situ dominance pattern of phytoplankton classes by dominance analysis as a proxy for realized niches
    Harmful Algae 58 , 74 - 84
    full text (url)
  • Flores-Moreno, H., Reich, P.B., Lind, E.M., Sullivan, L.L., Seabloom, E.W., Yahdjian, L., MacDougall, A.S., Reichmann, L.G., Alberti, J., Báez, S., Bakker, J.D., Cadotte, M.W., Caldeira, M.C., Chaneton, E.J., D'Antonio, C.M., Fay, P.A., Firn, J., Hagenah, N., Harpole, W.S., Iribarne, O., Kirkman, K.P., Knops, J.M.H., La Pierre, K.J., Laungani, R., Leakey, A.D.B., McCulley, R.L., Moore, J.L., Pascual, J., Borer, E.T., (2016):
    Climate modifies response of non-native and native species richness to nutrient enrichment
    Philos. Trans. R. Soc. B-Biol. Sci. 371 (1694), art. 20150273
    full text (url)
  • Grace, J.B., Anderson, T.M., Seabloom, E.W., Borer, E.T., Adler, P.B., Harpole, W.S., Hautier, Y., Hillebrand, H., Lind, E.M., Pärtel, M., Bakker, J.D., Buckley, Y.M., Crawley, M.J., Damschen, E.I., Davies, K.F., Fay, P.A., Firn, J., Gruner, D.S., Hector, A., Knops, J.M.H., MacDougall, A.S., Melbourne, B.A., Morgan, J.W., Orrock, J.L., Prober, S.M., Smith, M.D., (2016):
    Integrative modelling reveals mechanisms linking productivity and plant species richness
    Nature 529 (7586), 390 - 393
    full text (url)
  • Harpole, W.S., (2016):
    The hydrological niche and spatially structured species co-existence
    J. Veg. Sci. 27 (2), 215 - 216
    full text (url)
  • Harpole, W.S., Sullivan, L.L., Lindahl, B., Firn, J., Adler, P.B., Borer, E.T., Chase, J., Fay, P.A., Hautier, Y., Hillebrand, H., MacDougall, A.S., Seabloom, E.W., Williams, R., Bakker, J.D., Cadotte, M.W., Chaneton, E.J., Chu, C., Cleland, E.E., D’Antonio, C., Davies, K.F., Gruner, D.S., Hagenah, N., Kirkman, K., Knops, J.M.H., La Pierre, K.J., McCulley, R.L., Moore, J.L., Morgan, J.W., Prober, S.M., Risch, A.C., Schuetz, M., Stevens, C.J., Wragg, P.D., (2016):
    Addition of multiple limiting resources reduces grassland diversity
    Nature 537 (7618), 93 - 96
    full text (url)
  • Lewandowska, A.M., Biermann, A., Borer, E.T., Cebrián-Piqueras, M.A., Declerck, S.A.J., De Meester, L., Van Donk, E., Gamfeldt, L., Gruner, D.S., Hagenah, N., Harpole, W.S., Kirkman, K.P., Klausmeier, C.A., Kleyer, M., Knops, J.M.H., Lemmens, P., Lind, E.M., Litchman, E., Mantilla-Contreras, J., Martens, K., Meier, S., Minden, V., Moore, J.L., Venterink, H.O., Seabloom, E.W., Sommer, U., Striebel, M., Trenkamp, A., Trinogga, J., Urabe, J., Vyverman, W., Van de Waal, D.B., Widdicombe, C.E., Hillebrand, H., (2016):
    The influence of balanced and imbalanced resource supply on biodiversity–functioning relationship across ecosystems
    Philos. Trans. R. Soc. B-Biol. Sci. 371 (1694), art. 20150283
    full text (url)
  • Schuman, M.C., van Dam, N.M., Beran, F., Harpole, W.S., (2016):
    How does plant chemical diversity contribute to biodiversity at higher trophic levels?
    Curr. Opin. Insect Sci. 14 , 46 - 55
    full text (url)
  • Stein, C., Harpole, W.S., Suding, K.N., (2016):
    Transitions and invasion along a grazing gradient in experimental California grasslands
    Ecology 97 (9), 2319 - 2330
    full text (url)
  • Sullivan, L.L., Danielson, B.J., Harpole, W.S., (2016):
    Mammalian herbivores alter the population growth and spatial establishment of an early-establishing grassland species
    PLOS One 11 (2), e0147715
    full text (url)
  • Tredennick, A.T., Adler, P.B., Grace, J.B., Harpole, W.S., Borer, E.T., Seabloom, E.W., Anderson, T.M., Bakker, J.D., Biederman, L.A., Brown, C.S., Buckley, Y.M., Chu, C., Collins, S.L., Crawley, M.J., Fay, P.A., Firn, J., Gruner, D.S., Hagenah, N., Hautier, Y., Hector, A., Hillebrand, H., Kirkman, K., Knops, J.M.H., Laungani, R., Lind, E.M., MacDougall, A.S., McCulley, R.L., Mitchell, C.E., Moore, J.L., Morgan, J.W., Orrock, J.L., Peri, P.L., Prober, S.M., Risch, A.C., Schütz, M., Speziale, K.L., Standish, R.J., Sullivan, L.L., Wardle, G.M., Williams, R.J., Yang, L.H., (2016):
    Comment on “Worldwide evidence of a unimodal relationship between productivity and plant species richness”
    Science 351 (6272), 457
    full text (url)
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2015 (6)

  • Farrell, K.A., Harpole, W.S., Stein, C., Suding, K.N., Borer, E.T., (2015):
    Grassland arthropods are controlled by direct and indirect interactions with cattle but are largely unaffected by plant provenance
    PLOS One 10 (7), e0129823
    full text (url)
  • Fay, P.A., Prober, S.M., Harpole, W.S., Knops, J.M.H., Bakker, J.D., Borer, E.T., Lind, E.M., MacDougall, A.S., Seabloom, E.W., Wragg, P.D., Adler, P.B., Blumenthal, D.M., Buckley, Y.M., Chu, C., Cleland, E.E., Collins, S.L., Davies, K.F., Du, G., Feng, X., Firn, J., Gruner, D.S., Hagenah, N., Hautier, Y., Heckman, R.W., Jin, V.L., Kirkman, K.P., Klein, J., Ladwig, L.M., Li, Q., McCulley, R.L., Melbourne, B.A., Mitchell, C.E., Moore, J.L., Morgan, J.W., Risch, A.C., Schütz, M., Stevens, C.J., Wedin, D.A., Yang, L.H., (2015):
    Grassland productivity limited by multiple nutrients
    Nat. Plants 1 (7), art. 15080
    full text (url)
  • Leff, J.W., Jones, S.E., Prober, S.M., Barberán, A., Borer, E.T., Firn, J.L., Harpole, W.S., Hobbie, S.E., Hofmockel, K.S., Knops, J.M.H., McCulley, R.L., La Pierre, K., Risch, A.C., Seabloom, E.W., Schütz, M., Steenbock, C., Stevens, C.J., Fierer, N., (2015):
    Consistent responses of soil microbial communities to elevated nutrient inputs in grasslands across the globe
    Proc. Natl. Acad. Sci. U.S.A. 112 (35), 10967 - 10972
    full text (url)
  • Prober, S.M., Leff, J.W., Bates, S.T., Borer, E.T., Firn, J., Harpole, W.S., Lind, E.M., Seabloom, E.W., Adler, P.B., Bakker, J.D., Cleland, E.E., DeCrappeo, N.M., DeLorenze, E., Hagenah, N., Hautier, Y., Hofmockel, K.S., Kirkman, K.P., Knops, J.M.H., LaPierre, K.J., MacDougall, A.S., McCulley, R.L., Mitchell, C.E., Risch, A.C., Schuetz, M., Stevens, C.J., Williams, R.J., Fierer, N., (2015):
    Plant diversity predicts beta but not alpha diversity of soil microbes across grasslands worldwide
    Ecol. Lett. 18 (1), 85 - 95
    full text (url)
  • Seabloom, E.W., Borer, E.T., Buckley, Y.M., Cleland, E.E., Davies, K.F., Firn, J., Harpole, W.S., Hautier, Y., Lind, E.M., MacDougall, A.S., Orrock, J.L., Prober, S.M., Adler, P.B., Anderson, T.M., Bakker, J.D., Biederman, L.A., Blumenthal, D.M., Brown, C.S., Brudvig, L.A., Cadotte, M., Chu, C., Cottingham, K.L., Crawley, M.J., Damschen, E.I., Dantonio, C.M., DeCrappeo, N.M., Du, G., Fay, P.A., Frater, P., Gruner, D.S., Hagenah, N., Hector, A., Hillebrand, H., Hofmockel, K.S., Humphries, H.C., Jin, V.L., Kay, A., Kirkman, K.P., Klein, J.A., Knops, J.M.H., La Pierre, K.J., Ladwig, L., Lambrinos, J.G., Li, Q., Li, W., Marushia, R., McCulley, R.L., Melbourne, B.A., Mitchell, C.E., Moore, J.L., Morgan, J., Mortensen, B., O'Halloran, L.R., Pyke, D.A., Risch, A.C., Sankaran, M., Schuetz, M., Simonsen, A., Smith, M.D., Stevens, C.J., Sullivan, L., Wolkovich, E., Wragg, P.D., Wright, J., Yang, L., (2015):
    Plant species’ origin predicts dominance and response to nutrient enrichment and herbivores in global grasslands
    Nat. Commun. 6 , art. 7710
    full text (url)
  • Stevens, C.J., Lind, E.M., 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., Pierre, K.J.L., 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 (6), 1459 - 1465
    full text (url)
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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.