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

Niche dimensionality provides a general theoretical explanation for biodiversity—more niches, defined by more limiting factors, allow for more ways that species can coexist1. Because plant species compete for the same set of limiting resources, theory predicts that addition of a limiting resource eliminates potential trade-offs, reducing the number of species that can coexist2. Multiple nutrient limitation of plant production is common and therefore fertilization may reduce diversity by reducing the number or dimensionality of belowground limiting factors. At the same time, nutrient addition, by increasing biomass, should ultimately shift competition from belowground nutrients towards a one-dimensional competitive trade-off for light3. Here we show that plant species diversity decreased when a greater number of limiting nutrients were added across 45 grassland sites from a multi-continent experimental network4. The number of added nutrients predicted diversity loss, even after controlling for effects of plant biomass, and even where biomass production was not nutrient-limited. We found that elevated resource supply reduced niche dimensionality and diversity and increased both productivity5 and compositional turnover. Our results point to the importance of understanding dimensionality in ecological systems that are undergoing diversity loss in response to multiple global change factors.

%B Nature %V 537 %P 93-96 %G eng %U https://www.nature.com/articles/nature19324 %M KNZ001755 %R 10.1038/nature19324 %0 Journal Article %J Nature %D 2016 %T Integrative modelling reveals mechanisms linking productivity and plant species richness %A Grace, J.B. %A Anderson, T.M. %A Seabloom, E.W. %A E.T. Borer %A P. Adler %A Harpole, W.S. %A Hautier, Y. %A Hillebrand, H. %A Lind, E.M. %A Pärtel, M. %A J.D. Bakker %A Buckley, Y.M. %A Crawley, M.J. %A Damschen, E.I. %A Davies, K.F. %A Fay, P.A. %A Firn, J. %A Gruner, D.S. %A Hector, A. %A Knops, J.M.H. %A MacDougall, A.S. %A Melbourne, B.A. %A J.W. Morgan %A Orrock, J.L. %A Prober, S.M. %A M.D. Smith %X

How ecosystem productivity and species richness are interrelated is one of the most debated subjects in the history of ecology1. Decades of intensive study have yet to discern the actual mechanisms behind observed global patterns2, 3. Here, by integrating the predictions from multiple theories into a single model and using data from 1,126 grassland plots spanning five continents, we detect the clear signals of numerous underlying mechanisms linking productivity and richness. We find that an integrative model has substantially higher explanatory power than traditional bivariate analyses. In addition, the specific results unveil several surprising findings that conflict with classical models4, 5, 6, 7. These include the isolation of a strong and consistent enhancement of productivity by richness, an effect in striking contrast with superficial data patterns. Also revealed is a consistent importance of competition across the full range of productivity values, in direct conflict with some (but not all) proposed models. The promotion of local richness by macroecological gradients in climatic favourability, generally seen as a competing hypothesis8, is also found to be important in our analysis. The results demonstrate that an integrative modelling approach leads to a major advance in our ability to discern the underlying processes operating in ecological systems.

%B Nature %V 529 %P 390 - 393 %G eng %U https://www.nature.com/articles/nature16524 %N 7586 %M KNZ001787 %R 10.1038/nature16524 %0 Journal Article %J Nature Plants %D 2015 %T Grassland productivity limited by multiple nutrients %A Fay, P.A. %A Prober, S.M. %A Harpole, W.S. %A Knops, J.M.H. %A J.D. Bakker %A E.T. Borer %A Lind, E.M. %A MacDougall, A.S. %A Seabloom, E.W. %A Wragg, P.D. %A P. Adler %A D.M. Blumenthal %A Buckley, Y.M. %A Chu, C. %A Cleland, E.E. %A Scott. L. Collins %A Davies, K.F. %A G. Du %A Feng, X. %A Firn, J. %A Gruner, D.S. %A Hagenah, N. %A Hautier, Y. %A Heckman, R.W. %A Jin, V.L. %A Kirkman, K.P. %A Klein, J. %A L. Ladwig %A Li, Q. %A McCulley, R.L. %A Melbourne, B.A. %A Mitchell, C.E. %A Joslin L. Moore %A J.W. Morgan %A A. Risch %A schütz, M. %A Stevens, C.J. %A Wedin, D.A. %A Yang, L.H. %X

Terrestrial ecosystem productivity is widely accepted to be nutrient limited1. Although nitrogen (N) is deemed a key determinant of aboveground net primary production (ANPP)2,3, the prevalence of co-limitation by N and phosphorus (P) is increasingly recognized4,​5,​6,​7,​8. However, the extent to which terrestrial productivity is co-limited by nutrients other than N and P has remained unclear. Here, we report results from a standardized factorial nutrient addition experiment, in which we added N, P and potassium (K) combined with a selection of micronutrients (K+μ), alone or in concert, to 42 grassland sites spanning five continents, and monitored ANPP. Nutrient availability limited productivity at 31 of the 42 grassland sites. And pairwise combinations of N, P, and K+μ co-limited ANPP at 29 of the sites. Nitrogen limitation peaked in cool, high latitude sites. Our findings highlight the importance of less studied nutrients, such as K and micronutrients, for grassland productivity, and point to significant variations in the type and degree of nutrient limitation. We suggest that multiple-nutrient constraints must be considered when assessing the ecosystem-scale consequences of nutrient enrichment.

%B Nature Plants %V 1 %P 15080 %G eng %U https://www.nature.com/articles/nplants201580 %N 7 %M KNZ001788 %R 10.1038/nplants.2015.80 %0 Journal Article %J Nature Communications %D 2015 %T Plant species’ origin predicts dominance and response to nutrient enrichment and herbivores in global grasslands %A Seabloom, E.W. %A E.T. Borer %A Buckley, Y. %A Cleland, E.E. %A Davies, K.F. %A Firn, J. %A Harpole, W.S. %A Hautier, Y. %A Lind, E. %A MacDougall, A.S. %A Orrock, J.L. %A Prober, S.M. %A P. Adler %A Anderson, T.M. %A J.D. Bakker %A L.A. Biederman %A D.M. Blumenthal %A C.S. Brown %A Brudvig, L.A. %A Cadotte, M. %A Chu, C. %A Cottingham, K.L. %A Crawley, M.J. %A Damschen, E.I. %A D’Antonio, C.M. %A CeCrappeo, N.M. %A G. Du %A Fay, P.A. %A Frater, P. %A Gruner, D.S. %A Hagenah, N. %A Hector, A. %A Hillebrand, H. %A Hofmockel, K.S. %A Humphries, H.C. %A Jin, V.L. %A Kay, A.D. %A Kirkman, K.P. %A Klein, J.A. %A Knops, J.M.H. %A Kimberly J. La Pierre %A L. Ladwig %A Lambrinos, J.G. %A Li, Q. %A Li, W. %A Marushia, R. %K Biological sciences %K ecology %K Plant sciences %X

Exotic species dominate many communities; however the functional significance of species’ biogeographic origin remains highly contentious. This debate is fuelled in part by the lack of globally replicated, systematic data assessing the relationship between species provenance, function and response to perturbations. We examined the abundance of native and exotic plant species at 64 grasslands in 13 countries, and at a subset of the sites we experimentally tested native and exotic species responses to two fundamental drivers of invasion, mineral nutrient supplies and vertebrate herbivory. Exotic species are six times more likely to dominate communities than native species. Furthermore, while experimental nutrient addition increases the cover and richness of exotic species, nutrients decrease native diversity and cover. Native and exotic species also differ in their response to vertebrate consumer exclusion. These results suggest that species origin has functional significance, and that eutrophication will lead to increased exotic dominance in grasslands.

%B Nature Communications %V 6 %P 7710 - %G eng %U https://www.nature.com/articles/ncomms8710 %M KNZ001697 %R 10.1038/ncomms8710 %0 Journal Article %J Nature %D 2014 %T Eutrophication weakens stabilizing effects of diversity in natural grasslands %A Hautier, Y. %A Seabloom, E.W. %A E.T. Borer %A P. Adler %A Harpole, W.S. %A Hillebrand, H. %A Lind, E.M. %A MacDougall, A.S. %A Stevens, C.J. %A J.D. Bakker %A Buckley, Y.M. %A Chu, C. %A Scott. L. Collins %A Daleo, P. %A Damschen, E.I. %A Davies, K.F. %A Fay, P.A. %A Firn, J. %A Gruner, D.S. %A Jin, V.L. %A Klein, J.A. %A Knops, J.M.H. %A Kimberly J. La Pierre %A Li, W. %A McCulley, R.L. %A Melbourne, B.A. %A Joslin L. Moore %A O’Halloran, L.R. %A Prober, S.M. %A A. Risch %A Sankaran, M. %A Schuetz, M. %A Hector, A. %K Biodiversity %K Community ecology %K Grassland ecology %X

Studies of experimental grassland communities1, 2, 3, 4, 5, 6, 7 have demonstrated that plant diversity can stabilize productivity through species asynchrony, in which decreases in the biomass of some species are compensated for by increases in others1, 2. However, it remains unknown whether these findings are relevant to natural ecosystems, especially those for which species diversity is threatened by anthropogenic global change8, 9, 10, 11. Here we analyse diversity–stability relationships from 41 grasslands on five continents and examine how these relationships are affected by chronic fertilization, one of the strongest drivers of species loss globally8. Unmanipulated communities with more species had greater species asynchrony, resulting in more stable biomass production, generalizing a result from biodiversity experiments to real-world grasslands. However, fertilization weakened the positive effect of diversity on stability. Contrary to expectations, this was not due to species loss after eutrophication but rather to an increase in the temporal variation of productivity in combination with a decrease in species asynchrony in diverse communities. Our results demonstrate separate and synergistic effects of diversity and eutrophication on stability, emphasizing the need to understand how drivers of global change interactively affect the reliable provisioning of ecosystem services in real-world systems.

%B Nature %V 508 %P 521 -525 %G eng %U https://www.nature.com/articles/nature13014 %M KNZ001632 %R 10.1038/nature13014 %0 Journal Article %J Nature %D 2014 %T Herbivores and nutrients control grassland plant diversity via light limitation %A E.T. Borer %A Seabloom, E.W. %A Gruner, D.S. %A Harpole, W.S %A Hillebrand, H. %A Lind, E.M. %A P. Adler %A J. Alberti %A Anderson, T.M. %A J.D. Bakker %A L.A. Biederman %A D.M. Blumenthal %A C.S. Brown %A Brudvig, L.A. %A Buckley, Y.M. %A Cadotte, M. %A Chu, C. %A Cleland, E.E. %A Crawley, M.J. %A Daleo, P. %A Damschen, E.I. %A Davies, K.F. %A DeCrappeo, N.M. %A G. Du %A Firn, J. %A Hautier, Y. %A Heckman, R.W. %A Hector, A. %A HilleRisLambers, J. %A Iribarne, Oscar %A Klein, J.A. %A Knops, J.M.H. %A Kimberly J. La Pierre %A Leakey, A.D.B. %A Li, W. %A MacDougall, A.S. %A McCulley, R.L. %A Melbourne, B.A. %A Mitchell, C.E. %A Joslin L. Moore %A Mortensen, B. %A O'Halloran, L.R. %A Orrock, J.L. %A Pascual, J. %A Prober, S.M. %A Pyke, D.A. %A A. Risch %A Schuetz, M. %A M.D. Smith %A Stevens, C.J. %A L.L. Sullivan %A Williams, R.J. %A Wragg, P.D. %A Wright, J.P. %A Yang, L.H. %B Nature %V 508 %P 517 - 520 %G eng %U https://www.nature.com/articles/nature13144 %N 7497 %M KNZ001793 %R 10.1038/nature13144 %0 Journal Article %J Ecology Letters %D 2013 %T Life-history constraints in grassland plant species: a growth-defence trade-off is the norm %A Lind, E.M. %A E.T. Borer %A Seabloom, E. %A P. Adler %A J.D. Bakker %A D.M. Blumenthal %A Crawley, M. %A Davies, K. %A Firn, J. %A Gruner, D.S. %A Harpole, W.S. %A Hautier, Y. %A Hillebrand, H. %A Knops, J. %A Melbourne, B. %A Mortensen, B. %A A. Risch %A Schuetz, M. %A Stevens, C. %A Wragg, P.D. %X

Plant growth can be limited by resource acquisition and defence against consumers, leading to contrasting trade-off possibilities. The competition-defence hypothesis posits a trade-off between competitive ability and defence against enemies (e.g. herbivores and pathogens). The growth-defence hypothesis suggests that strong competitors for nutrients are also defended against enemies, at a cost to growth rate. We tested these hypotheses using observations of 706 plant populations of over 500 species before and following identical fertilisation and fencing treatments at 39 grassland sites worldwide. Strong positive covariance in species responses to both treatments provided support for a growth-defence trade-off: populations that increased with the removal of nutrient limitation (poor competitors) also increased following removal of consumers. This result held globally across 4 years within plant life-history groups and within the majority of individual sites. Thus, a growth-defence trade-off appears to be the norm, and mechanisms maintaining grassland biodiversity may operate within this constraint.

%B Ecology Letters %V 16 %P 513 - 521 %G eng %U https://onlinelibrary.wiley.com/doi/abs/10.1111/ele.12078 %N 4 %M KNZ001791 %R 10.1111/ele.12078 %0 Journal Article %J Global Change Biology %D 2013 %T Predicting invasion in grassland ecosystems: is exotic dominance the real embarrassment of richness? %A Seabloom, E.W. %A E.T. Borer %A Buckley, Y. %A Cleland, E.E. %A Davies, K. %A Firn, J. %A Harpole, W.S. %A Hautier, Y. %A Lind, E. %A MacDougall, A. %A Orrock, J.L. %A Prober, S.M. %A P. Adler %A J. Alberti %A Anderson, M.T. %A J.D. Bakker %A L.A. Biederman %A D.M. Blumenthal %A C.S. Brown %A Brudvig, L.A. %A Caldeira, M. %A Chu, C. %A Crawley, M.J. %A Daleo, P. %A Damschen, E.I. %A D'Antonio, C.M. %A DeCrappeo, N.M. %A Dickman, C.R. %A G. Du %A Fay, P.A. %A Frater, P. %A Gruner, D.S. %A Hagenah, N. %A Hector, A. %A Helm, A. %A Hillebrand, H. %A Hofmockel, K.S. %A Humphries, H.C. %A Iribarne, O. %A Jin, V.L. %A Kay, A. %A Kirkman, K.P. %A Klein, J.A. %A Knops, J.M.H. %A Kimberly J. La Pierre %A L. Ladwig %A Lambrinos, J.G. %A Leakey, A.D.B. %A Li, Q. %A Li, W. %A McCulley, R. %A Melbourne, B. %A Mitchell, C.E. %A Joslin L. Moore %A J.W. Morgan %A Mortensen, B. %A O'Halloran, L.R. %A Pärtel, M. %A Pascual, J %A Pyke, D.A. %A A. Risch %A Salguero-Gomez, R. %A Sankaran, M. %A Schuetz, M. %A Simonsen, A. %A M.D. Smith %A Stevens, C. %A Sullivan, L. %A Wardle, G.M. %A Wolkovich, E.M. %A Wragg, P.D. %A Wright, J. %A Yang, L. %X

Invasions have increased the size of regional species pools, but are typically assumed to reduce native diversity. However, global-scale tests of this assumption have been elusive because of the focus on exotic species richness, rather than relative abundance. This is problematic because low invader richness can indicate invasion resistance by the native community or, alternatively, dominance by a single exotic species. Here, we used a globally replicated study to quantify relationships between exotic richness and abundance in grass-dominated ecosystems in 13 countries on six continents, ranging from salt marshes to alpine tundra. We tested effects of human land use, native community diversity, herbivore pressure, and nutrient limitation on exotic plant dominance. Despite its widespread use, exotic richness was a poor proxy for exotic dominance at low exotic richness, because sites that contained few exotic species ranged from relatively pristine (low exotic richness and cover) to almost completely exotic-dominated ones (low exotic richness but high exotic cover). Both exotic cover and richness were predicted by native plant diversity (native grass richness) and land use (distance to cultivation). Although climate was important for predicting both exotic cover and richness, climatic factors predicting cover (precipitation variability) differed from those predicting richness (maximum temperature and mean temperature in the wettest quarter). Herbivory and nutrient limitation did not predict exotic richness or cover. Exotic dominance was greatest in areas with low native grass richness at the site- or regional-scale. Although this could reflect native grass displacement, a lack of biotic resistance is a more likely explanation, given that grasses comprise the most aggressive invaders. These findings underscore the need to move beyond richness as a surrogate for the extent of invasion, because this metric confounds monodominance with invasion resistance. Monitoring species' relative abundance will more rapidly advance our understanding of invasions.

%B Global Change Biology %V 19 %P 3677 - 3687 %G eng %U https://onlinelibrary.wiley.com/doi/abs/10.1111/gcb.12370 %N 12 %M KNZ001790 %R 10.1111/gcb.12370 %0 Journal Article %J Science %D 2011 %T Productivity is a poor predictor of plant species richness %A P. Adler %A Seabloom, E.W. %A E.T. Borer %A Hillebrand, H. %A Hautier, Y. %A Hector, A. %A Harpole, W.S. %A O'Halloran, L.R. %A Grace, J.B. %A Anderson, T.M. %A J.D. Bakker %A L.A. Biederman %A C.S. Brown %A Buckley, Y.M. %A Calabrese, L.B. %A Chu, C.J. %A Cleland, E.E. %A Scott. L. Collins %A Cottingham, K.L. %A Crawley, M.J. %A Damschen, E.I. %A Davies, K.F. %A DeCrappeo, N.M. %A Fay, P.A. %A Firn, J. %A Frater, P. %A Gasarch, E.I. %A Gruner, D.S. %A Hagenah, N. %A HilleRisLambers, J. %A Humphries, H.C. %A Jin, V.L. %A Kay, A. %A Kirkman, K.P. %A Klein, J.A. %A Knops, J.M.H. %A Kimberly J. La Pierre %A Lambrinos, J.G. %A Li, W. %A MacDougall, A.S. %A McCulley, R.L. %A Melbourne, B.A. %A Mitchell, C.E. %A Joslin L. Moore %X

For more than 30 years, the relationship between net primary productivity and species richness has generated intense debate in ecology about the processes regulating local diversity. The original view, which is still widely accepted, holds that the relationship is hump-shaped, with richness first rising and then declining with increasing productivity. Although recent meta-analyses questioned the generality of hump-shaped patterns, these syntheses have been criticized for failing to account for methodological differences among studies. We addressed such concerns by conducting standardized sampling in 48 herbaceous-dominated plant communities on five continents. We found no clear relationship between productivity and fine-scale (meters−2) richness within sites, within regions, or across the globe. Ecologists should focus on fresh, mechanistic approaches to understanding the multivariate links between productivity and richness.

%B Science %V 333 %P 1750 -1753 %G eng %U http://science.sciencemag.org/content/333/6050/1750 %M KNZ001448 %R 10.1126/science.1204498