Frequency-dependent feedback constrains plant community coexistence


Ecological theory suggests that coexistence of many species within communities requires negative frequency-dependent feedbacks to prevent exclusion of the least fit species. For plant communities, empirical evidence of negative frequency dependence driving species coexistence and diversity patterns is rapidly accumulating, but connecting these findings to theory has been difficult as corresponding theoretical frameworks only consider small numbers of species. Here, we show how frequency-dependent feedback constrains community coexistence, regardless of the number of species and inherent fitness inequalities between them. Any interaction network can be characterized by a single community interaction coefficient, IC, which determines whether community-level feedback is positive or negative. Negative feedback is a necessary (but not sufficient) condition for persistence of the entire community. Even in cases where the coexistence equilibrium state cannot recover from perturbations, IC < 0 can enable species persistence via cyclic succession. The number of coexisting species is predicted to increase with the average strength of negative feedback. This prediction is supported by patterns of tree species diversity in more than 200,000 deciduous forest plots in the eastern United States, which can be reproduced in simulations that span the observed range of community feedback. By providing a quantitative metric for the strength of negative feedback needed for coexistence, we can now integrate theory and empirical data to test whether observed feedback–diversity correlations are strong enough to infer causality.

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Fig. 1: Conceptual representation of how the frequency-dependent feedback framework can be extended to the community level for any number of plant species (here, n = 5).
Fig. 2: Two hundred runs of plant community assembly show that more species can stably coexist under stronger negative feedback.
Fig. 3: Effects of frequency-dependent feedback on plant community structure.
Fig. 4: Application of the theoretical framework reveals that observed patterns of negative frequency dependence in over 200,000 forest plots in the eastern United States are consistent with predictions from theoretical communities in which community-level feedback determines species richness.


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This work was conducted as part of the ‘Plant–soil feedback theory’ Working Group at the National Institute for Mathematical and Biological Synthesis, sponsored by the National Science Foundation through NSF Award DBI-1300426, with additional support from The University of Tennessee, Knoxville. The work was further supported by NSF grant DEB-1738041, awarded to J.D.B. We thank K. C. Abbott, J. Bauer, L. S. Comita, K. M. Crawford, S. A. Mangan, J. Umbanhowar and K. N. Suding for discussions of this work during working group meetings, J. Aning for assistance with the FIA data analysis, and T. Markus for assistance with the preparation of the figures. We also thank F. May for comments that improved the manuscript. Finally, we thank all of the FIA employees who collected the data that enabled us to test model predictions.

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M.B.E., D.J.J., K.M.L.M., A.E.S. and J.D.B. designed the study. M.B.E. and M.B. performed the analytical model analyses. M.B.E., M.B., J.J., K.M.L.M. and A.E.S. developed the code, and performed and analysed the model simulations. M.B.E., M.B., D.J.J. and J.D.B. analysed the field observation data. M.B.E. and J.D.B. wrote the first draft, after which all authors contributed to improving the manuscript.

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Correspondence to Maarten B. Eppinga.

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Supplementary information

Supplementary Information

Supplementary Information sections 1–7 and Supplementary Figures 1–13

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Supplementary Table 1

A spreadsheet containing the values of average pair-wise feedback summarized in Fig. 4b of the main text

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Eppinga, M.B., Baudena, M., Johnson, D.J. et al. Frequency-dependent feedback constrains plant community coexistence. Nat Ecol Evol 2, 1403–1407 (2018).

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