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Complementarity among four highly productive grassland species depends on resource availability

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Abstract

Positive species richness–productivity relationships are common in biodiversity experiments, but how resource availability modifies biodiversity effects in grass–legume mixtures composed of highly productive species is yet to be explicitly tested. We addressed this question by choosing two grasses (Arrhenatherum elatius and Dactylis glomerata) and two legumes (Medicago × varia and Onobrychis viciifolia) which are highly productive in monocultures and dominant in mixtures (the Jena Experiment). We established monocultures, all possible two- and three-species mixtures, and the four-species mixture under three different resource supply conditions (control, fertilization, and shading). Compared to the control, community biomass production decreased under shading (−56 %) and increased under fertilization (+12 %). Net diversity effects (i.e., mixture minus mean monoculture biomass) were positive in the control and under shading (on average +15 and +72 %, respectively) and negative under fertilization (−10 %). Positive complementarity effects in the control suggested resource partitioning and facilitation of growth through symbiotic N2 fixation by legumes. Positive complementarity effects under shading indicated that resource partitioning is also possible when growth is carbon-limited. Negative complementarity effects under fertilization suggested that external nutrient supply depressed facilitative grass–legume interactions due to increased competition for light. Selection effects, which quantify the dominance of species with particularly high monoculture biomasses in the mixture, were generally small compared to complementarity effects, and indicated that these species had comparable competitive strengths in the mixture. Our study shows that resource availability has a strong impact on the occurrence of positive diversity effects among tall and highly productive grass and legume species.

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Acknowledgments

This study was part of the Jena Experiment and supported by the German Science Foundation (RO2397/1 within FOR456). We acknowledge A. Weigelt for field coordination. We thank U. Gerighausen, S. Ferber, C. Kups, the gardeners, and several student helpers for their support with the establishment and management of the four-species experiment and during sampling; I. Hilke, S. Matthei, and H. Geilmann for conducting chemical and stable isotope analyses; F. Voigt and B. Schlöffel for the construction of the shade roof; K. Hippler, M. Hertel, and A. Fastnacht for setting up and maintaining the weather station; and three anonymous reviewers for their helpful comments on a previous version of the manuscript.

Author contribution statement

CR, E-DS, BS, and OK conceived and designed the experiment. CR and OK performed the experiment. CR analyzed the data and wrote the manuscript; E-DS and BS discussed the results and contributed to writing the manuscript; OK provided editorial advice.

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Correspondence to Christiane Roscher.

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Communicated by Scott Collins.

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Roscher, C., Schmid, B., Kolle, O. et al. Complementarity among four highly productive grassland species depends on resource availability. Oecologia 181, 571–582 (2016). https://doi.org/10.1007/s00442-016-3587-4

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  • DOI: https://doi.org/10.1007/s00442-016-3587-4

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