Abstract
Central European grasslands vary widely in productivity and in mowing and grazing regimes. The resulting differences in competition and heterogeneity among grasslands might have direct effects on plants, but might also affect the growth and morphology of their offspring through maternal effects or adaptive evolution. To test for such transgenerational effects, we grew plants of the clonal herb Trifolium repens from seeds collected in 58 grassland sites differing in productivity and mowing and grazing intensities in different treatments: without competition, with homogeneous competition, and with heterogeneous competition. In the competition-free treatment, T. repens from more productive, less frequently mown, and less intensively grazed sites produced more vegetative offspring, but this was not the case in the other treatments. When grown among or in close proximity to competitors, T. repens plants did not show preferential growth towards open spaces (i.e., no horizontal foraging), but did show strong vertical foraging by petiole elongation. In the homogeneous competition treatment, petiole length increased with the productivity of the parental site, but this was not the case in the heterogeneous competition treatment. Moreover, petiole length increased with mowing frequency and grazing intensity of the parental site in all but the homogeneous competition treatment. In summary, although the expression of differences between plants from sites with different productivities and land-use intensities depended on the experimental treatment, our findings imply that there are transgenerational effects of land use on the morphology and performance of T. repens.





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Acknowledgments
We thank Eva Völler and Harald Auge for collecting the seeds, and Yuanye Zhang, Lidewij Keser, Anne Kempel, Andreas Ensslin, Gemma Rutten, Yvonne Zürcher, Joël Baumann, and Michael Moser for help with the harvesting. ZW and MvK acknowledge the support of the Sino-Swiss Science and Technology Cooperation (EG 14-092009). We thank the managers of the three Exploratories—Swen Renner, Sonja Gockel, Kerstin Wiesner, and Martin Gorke—for their work in maintaining the plot and project infrastructure; Simone Pfeiffer and Christiane Fischer for providing support through the central office, Michael Owonibi for managing the central database, and Eduard Linsenmair, Dominik Hessenmöller, Jens Nieschulze, Ingo Schöning, François Buscot, Ernst-Detlef Schulze, Wolfgang W. Weisser, and the late Elisabeth Kalko for their role in setting up the Biodiversity Exploratories project. The work has been partly funded by the DFG Priority Program 1374 “Infrastructure-Biodiversity-Exploratories” and NSFC grants (41371077, 31570530). The experiment complies with the current laws of Switzerland (the country in which the experiment was performed).
Author contribution statement
All authors together conceived and designed the experiment. ZW and MVK performed the experiment. MVK analyzed the data. OB, MF and DP provided seed materials and data on land use intensities. ZW and MVK wrote the manuscript, with comments and help from the other authors.
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Communicated by Jennifer A. Lau.
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Wang, Z., Bossdorf, O., Prati, D. et al. Transgenerational effects of land use on offspring performance and growth in Trifolium repens . Oecologia 180, 409–420 (2016). https://doi.org/10.1007/s00442-015-3480-6
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DOI: https://doi.org/10.1007/s00442-015-3480-6


