%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 Philosophical Transactions of the Royal Society B: Biological Sciences %D 2016 %T Climate modifies response of non-native and native species richness to nutrient enrichment %A Flores-Moreno, H. %A P.B. Reich %A Lind, E.M. %A L.L. Sullivan %A Seabloom, E.W. %A Yahdjian, L. %A MacDougall, A.S. %A Reichmann, L.G. %A J. Alberti %A Báez, S. %A J.D. Bakker %A Cadotte, M.W. %A Caldeira, M.C. %A Chaneton, E.J. %A D'Antonio, C.M. %A Fay, P.A. %A Firn, J. %A Hagenah, N. %A Harpole, W. S. %A Iribarne, O. %A Kirkman, K.P. %A Knops, J.M. H. %A Kimberly J. La Pierre %A Laungani, R. %A Leakey, A.D. B. %A McCulley, R.L. %A Joslin L. Moore %A Pascual, J. %A E.T. Borer %X

Ecosystem eutrophication often increases domination by non-natives and causes displacement of native taxa. However, variation in environmental conditions may affect the outcome of interactions between native and non-native taxa in environments where nutrient supply is elevated. We examined the interactive effects of eutrophication, climate variability and climate average conditions on the success of native and non-native plant species using experimental nutrient manipulations replicated at 32 grassland sites on four continents. We hypothesized that effects of nutrient addition would be greatest where climate was stable and benign, owing to reduced niche partitioning. We found that the abundance of non-native species increased with nutrient addition independent of climate; however, nutrient addition increased non-native species richness and decreased native species richness, with these effects dampened in warmer or wetter sites. Eutrophication also altered the time scale in which grassland invasion responded to climate, decreasing the importance of long-term climate and increasing that of annual climate. Thus, climatic conditions mediate the responses of native and non-native flora to nutrient enrichment. Our results suggest that the negative effect of nutrient addition on native abundance is decoupled from its effect on richness, and reduces the time scale of the links between climate and compositional change.

%B Philosophical Transactions of the Royal Society B: Biological Sciences %V 3719371 %P 20150273 %G eng %U https://royalsocietypublishing.org/doi/10.1098/rstb.2015.0273 %N 1694 %M KNZ001747 %R 10.1098/rstb.2015.0273 %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 Ecology %D 2015 %T Anthropogenic nitrogen deposition predicts local grassland primary production worldwide %A Stevens, C.J. %A Lind, E.M. %A Hautier, Y. %A Harpole, W.S. %A E.T. Borer %A Hobbie, S. %A Seabloom, E.Q. %A L. Ladwig %A J.D. Bakker %A Chu, C. %A Scott. L. Collins %A Davies, K.F. %A Firn, J. %A Hillebrand, H. %A Kimberly J. La Pierre %A MacDougall, A.S. %A Melbourne, B.A. %A McCulley, R.L. %A J.W. Morgan %A Orrock, J.L. %A Prober, S.M. %A A. Risch %A Schultz, M. %A Wragg, P.D. %X

Humans dominate many important Earth system processes including the nitrogen (N) cycle. Atmospheric N deposition affects fundamental processes such as carbon cycling, climate regulation, and biodiversity, and could result in changes to fundamental Earth system processes such as primary production. Both modelling and experimentation have suggested a role for anthropogenically altered N deposition in increasing productivity, nevertheless, current understanding of the relative strength of N deposition with respect to other controls on production such as edaphic conditions and climate is limited. Here we use an international multiscale data set to show that atmospheric N deposition is positively correlated to aboveground net primary production (ANPP) observed at the 1-m2 level across a wide range of herbaceous ecosystems. N deposition was a better predictor than climatic drivers and local soil conditions, explaining 16% of observed variation in ANPP globally with an increase of 1 kg N·ha−1·yr−1 increasing ANPP by 3%. Soil pH explained 8% of observed variation in ANPP while climatic drivers showed no significant relationship. Our results illustrate that the incorporation of global N deposition patterns in Earth system models are likely to substantially improve estimates of primary production in herbaceous systems. In herbaceous systems across the world, humans appear to be partially driving local ANPP through impacts on the N cycle.

%B Ecology %V 96 %P 1459 -1465 %G eng %U https://esajournals.onlinelibrary.wiley.com/doi/full/10.1890/14-1902.1 %M KNZ001698 %R http://dx.doi.org/10.1890/14-1902.1 %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 Ecology Letters %D 2015 %T Plant diversity predicts beta but not alpha diversity of soil microbes across grasslands worldwide %A Prober, S.M. %A Leff, J.W. %A Bates, S.T. %A E.T. Borer %A Firn, J. %A Harpole, W.S. %A Lind, E.M. %A Seabloom, E.W. %A P. Adler %A J.D. Bakker %A E.E Cleland %A DeCrappeo, N.M. %A DeLorenze, E. %A Hagenah, N. %A Hautier, Y. %A Hofmockel, K.S. %A Kirkman, K.P. %A Knops, J.M.H. %A Kimberly J. La Pierre %A MacDougall, A.S. %A McCulley, R.L. %A Mitchell, C.E. %A A. Risch %A Schuetz, M. %A Stevens, C.J. %A Williams, R.J. %A Fierer, N. %X

Aboveground–belowground interactions exert critical controls on the composition and function of terrestrial ecosystems, yet the fundamental relationships between plant diversity and soil microbial diversity remain elusive. Theory predicts predominantly positive associations but tests within single sites have shown variable relationships, and associations between plant and microbial diversity across broad spatial scales remain largely unexplored. We compared the diversity of plant, bacterial, archaeal and fungal communities in one hundred and forty-five 1 m2 plots across 25 temperate grassland sites from four continents. Across sites, the plant alpha diversity patterns were poorly related to those observed for any soil microbial group. However, plant beta diversity (compositional dissimilarity between sites) was significantly correlated with the beta diversity of bacterial and fungal communities, even after controlling for environmental factors. Thus, across a global range of temperate grasslands, plant diversity can predict patterns in the composition of soil microbial communities, but not patterns in alpha diversity.

%B Ecology Letters %V 18 %P 85 -95 %G eng %U https://onlinelibrary.wiley.com/doi/abs/10.1111/ele.12381 %M KNZ001678 %R 10.1111/ele.12381 %0 Journal Article %J Global Ecology and Biogeography %D 2014 %T Anthropogenic-based regional-scale factors most consistently explain plot-level exotic diversity in grasslands %A MacDougall, A.S. %A Bennett, J.R. %A Firn, J. %A Seabloom, E.W. %A E.T. Borer %A Lind, E.M. %A Orrock, J.L. %A Harpole, W.S. %A Hautier, Y. %A P. Adler %A Cleland, E. %A Davies, K. %A Melbourne, B. %A Prober, S.M. %A J.D. Bakker %A Fay, P.A. %A Jin, V.L. %A Kendig, A. %A Kimberly J. La Pierre %A Joslin L. Moore %A J.W. Morgan %A Stevens, C.J. %X

Aim Evidence linking the accumulation of exotic species to the suppression of native diversity is equivocal, often relying on data from studies that have used different methods. Plot-level studies often attribute inverse relationships between native and exotic diversity to competition, but regional abiotic filters, including anthropogenic influences, can produce similar patterns. We seek to test these alternatives using identical scale-dependent sampling protocols in multiple grasslands on two continents. Location Thirty-two grassland sites in North America and Australia. Methods We use multiscale observational data, collected identically in grain and extent at each site, to test the association of local and regional factors with the plot-level richness and abundance of native and exotic plants. Sites captured environmental and anthropogenic gradients including land-use intensity, human population density, light and soil resources, climate and elevation. Site selection occurred independently of exotic diversity, meaning that the numbers of exotic species varied randomly thereby reducing potential biases if only highly invaded sites were chosen. Results Regional factors associated directly or indirectly with human activity had the strongest associations with plot-level diversity. These regional drivers had divergent effects: urban-based economic activity was associated with high exotic : native diversity ratios; climate- and landscape-based indicators of lower human population density were associated with low exotic : native ratios. Negative correlations between plot-level native and exotic diversity, a potential signature of competitive interactions, were not prevalent; this result did not change along gradients of productivity or heterogeneity. Main conclusion We show that plot-level diversity of native and exotic plants are more consistently associated with regional-scale factors relating to urbanization and climate suitability than measures indicative of competition. These findings clarify the long-standing difficulty in resolving drivers of exotic diversity using single-factor mechanisms, suggesting that multiple interacting anthropogenic-based processes best explain the accumulation of exotic diversity in modern landscapes.

%B Global Ecology and Biogeography %V 23 %P 802 - 810 %G eng %U https://onlinelibrary.wiley.com/doi/abs/10.1111/geb.12157 %N 7 %M KNZ001792 %R 10.1111/geb.12157 %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