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Addition of multiple limiting resources reduces grassland diversity



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.

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Figure 1: Biodiversity and number of resources.
Figure 2: Biomass and light.
Figure 3: Multiple resource limitation.
Figure 4: Community composition.


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We thank the Minnesota Supercomputer Institute for hosting project data, the University of Minnesota Institute on the Environment for hosting Nutrient Network meetings, and each site investigator for funding their site-level operations. Network coordination and data management were supported by funds from the National Science Foundation Research Coordination Network (NSF-DEB-1042132) to E.T.B. and E.W.S. from the Long Term Ecological Research program (NSF-DEB-1234162) to the Cedar Creek LTER, and from the Institute on the Environment (DG-0001-13). Konza NutNet site was funded by the Konza Prairie LTER; the Saline Experimental Range NutNet site was funded by a Yale Institute for Biospheric Studies Pilot Grant. Nitrogen fertilizer was donated to the Nutrient Network by Crop Production Services, Loveland, Colorado. We thank N. Gotelli for discussions.

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Authors and Affiliations



W.S.H. analysed the data and wrote the paper with contributions and input from all authors. L.L.S., E.M.L. and J.F. contributed to data analysis. W.S.H., E.W.S. and E.T.B. developed and framed the research questions. W.S.H., E.W.S., E.T.B. and E.M.L. are Nutrient Network coordinators. All authors collected data used in this analysis. Author contribution matrix provided as Supplementary Table 2.

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Correspondence to W. Stanley Harpole.

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The authors declare no competing financial interests.

Additional information

Reviewer Information Nature thanks J. Levine, B. Schmid and the other anonymous reviewer(s) for their contribution to the peer review of this work.

Extended data figures and tables

Extended Data Table 1 The effects of nutrient addition on diversity loss and richness loss increase with time
Extended Data Table 2 The number of added resources predicts diversity loss after controlling for other variables
Extended Data Table 3 The number of added resources is an important predictor even after controlling for other variables, for sites that had light and litter data
Extended Data Table 4 Diversity loss due to addition of nutrients associated with soil properities

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Supplementary Information

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Harpole, W., Sullivan, L., Lind, E. et al. Addition of multiple limiting resources reduces grassland diversity. Nature 537, 93–96 (2016).

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