Abstract
In experimental plant communities, relationships between biodiversity and ecosystem functioning have been found to strengthen over time1,2, a fact often attributed to increased resource complementarity between species in mixtures3 and negative plant–soil feedbacks in monocultures4. Here we show that selection for niche differentiation between species can drive this increasing biodiversity effect. Growing 12 grassland species in test monocultures and mixtures, we found character displacement between species and increased biodiversity effects when plants had been selected over 8 years in species mixtures rather than in monocultures. When grown in mixtures, relative differences in height and specific leaf area between plant species selected in mixtures (mixture types) were greater than between species selected in monocultures (monoculture types). Furthermore, net biodiversity and complementarity effects1,2 were greater in mixtures of mixture types than in mixtures of monoculture types. Our study demonstrates a novel mechanism for the increase in biodiversity effects: selection for increased niche differentiation through character displacement. Selection in diverse mixtures may therefore increase species coexistence and ecosystem functioning in natural communities and may also allow increased mixture yields in agriculture or forestry. However, loss of biodiversity and prolonged selection of crops in monoculture may compromise this potential for selection in the longer term.
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Acknowledgements
This study was supported by the Swiss National Science Foundation (grant number 130720 to B.S.) and the University Research Priority Program Global Change and Biodiversity of the University of Zurich. Thanks to D. Trujillo Villegas, L. Oesch, T. Zwimpfer, M. Furler, R. Husi, the gardeners of the Jena Experiment and student helpers for technical assistance. G.B.D.D. acknowledges the NWO-ALW VIDI grant scheme for financial support.
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B.S. and J.S.P. conceptualized the study; D.Z.-D. designed the experimental procedure and carried out the experiment with the help of B.S., D.F.B.F. and V.Y.; B.S., D.Z.-D. and D.F.B.F. analysed the data; D.Z.-D., B.S. and D.F.B.F. wrote the paper with input from J.S.P., V.Y. and G.B.D.D. All authors discussed study design, field and glasshouse work, and analysis.
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Data have been deposited at the Dryad Data Repository (http://dx.doi.org/10.5061/dryad.750df).
Extended data figures and tables
Extended Data Figure 1 Designed number of pots planted for each combination of species.
Plants from three different selection histories (monoculture, plot containing one species; monofunctional group, plot containing at least four species of the same functional group of plants; mixed-functional group, plot containing at least four species of four different functional groups) were grown in three different types of test communities (monoculture, monofunctional mixture, mixed-functional mixture). Twelve species in the Jena Experiment were chosen from all four functional groups: grass (g) (Festuca pratensis, Festuca rubra, Poa pratensis), small herb (sh) (Plantago lanceolata, Prunella vulgaris, Veronica chamaedrys), tall herb (th) (Crepis biennis, Galium mollugo, Geranium pratense), legume (l) (Lathyrus pratensis, Onobrychis viciifolia, Trifolium repens); numbers after the letter abbreviations refer to the different species. This design was used once with plants raised from cuttings (Block 1) and once with plants raised from seedlings (Block 2). Overall we aimed to obtain the same 12 monocultures and 48 two-species combinations as test communities for each block. Availability of species precluded some of the two-species combinations in each block, such that they had to be replaced by other combinations. This yielded a total of 50 combinations across the two blocks, with several that were unique within a block. Each monoculture and each two-species combination was assembled three times for each of the three types of selection histories in each block. Some monocultures and some two-species combinations could not be realized with all types of selection histories in both blocks. Overall, there were 855 pots, 168 monocultures and 687 two-species mixtures; for 545 of the latter, the net biodiversity effect could be partitioned into complementarity and sampling effects. Some missing monocultures precluded the calculation of biodiversity effects in certain mixtures.
Extended Data Figure 2 Biodiversity–productivity relationship is stronger for plants with a common selection history.
Aboveground net primary productivity of communities in an experimental manipulation of plant species richness and selection history (common history versus no common history). In this experiment, species represented an expanded set from the present experiment (52 species), and were planted within a large-scale field experiment in Jena, Germany, on mixed soil from 48 plots from which plants had been selected, thus equalizing potential effects of soil legacy among treatments. Plants without selection history were grown from seed from a seed company, while plants with selection history were seed progeny from plots of exactly the same species composition as the one in which they were replanted (same propagation procedure as for the 12 species used in the test communities of the present study). The slope of the biodiversity–productivity relationship was steeper for plants with a common selection history (significance of slope differences tested with interaction term log(species richness) × selection history in mixed model with random-effects factor for 48 specific plant communities; P < 0.001, n = 96).
Extended Data Figure 3 Selection for different biochemical features in monocultures and mixtures.
Ordinations (non-metric multidimensional scaling (NMDS)) of second derivative of spectral wavenumbers of 8 of the 12 species used in the present study, showing effects of 8-year selection history on plant individuals derived from monoculture and mixture communities (Jena Experiment). This can be an indication of selection for different biochemical features over 8 years in monoculture and mixtures. Stress values reflect a measure of goodness of fit for NMDS, with lower values showing better representation of the original data.
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Zuppinger-Dingley, D., Schmid, B., Petermann, J. et al. Selection for niche differentiation in plant communities increases biodiversity effects. Nature 515, 108–111 (2014). https://doi.org/10.1038/nature13869
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DOI: https://doi.org/10.1038/nature13869
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