Abstract
The ongoing climate change is triggering plant community thermophilization. This selection process ought to shift community composition towards species adapted to warmer climates but may also lead to biotic homogenization. The link between thermophilization and homogenization and the community dynamics that drive them (colonization and extinction) remain unknown but is critical for understanding community responses under rapid environmental change. We used 14,167 pairs of plots to study shifts in plant community during 10 years of rising temperature in 80 forest ecoregions of France. We computed community mean thermal optimum (thermophilization) and Δβ-diversity (homogenization) for each ecoregion and partitioned these changes into extinction and colonization dynamics of cold- and warm-adapted species. Forest understorey communities thermophilized on average by 0.12 °C per decade and up to 0.20 °C per decade in warm ecoregions. This rate was entirely driven by extinction dynamics. Extinction of cold-adapted species was a driver of homogenization but it was compensated for by the colonization of rare species and the extinction of common species, resulting in the absence of an apparent homogenization trend. Here we show a dieback of present cold-adapted species rather than an adaptation of communities via the arrival of warm-adapted species, with a mutually cancelling effect on β-diversity. These results suggest that a future loss of biodiversity and delayed biotic homogenization should be considered.
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Data availability
French National Forest Inventory data are freely distributed by IGN at https://inventaire-forestier.ign.fr. The dataset and the code used to reproduce our analysis can be downloaded from GitHub https://github.com/Jeremy-borderieux/Article_thermo_beta_part.git.
Code availability
The code and workflow to run and reproduce our analysis can be downloaded from GitHub https://github.com/Jeremy-borderieux/Article_thermo_beta_part.git.
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Acknowledgements
We are grateful to the French Institute for Geographic and Forest Information (IGN) and their field technicians for providing the NFI data and a precise description of the forest ecoregions. We acknowledge the funding from the Labex Arbre. J.B. was funded by a joint AgroParisTech and Région Grand-Est grant (grant no. 19_GE8_01020p05035) and J.M.S.-D. was funded by the ANR-JCJC (Agence Nationale de la Recherche, jeunes chercheuses et jeunes chercheurs) SEEDFOR (ANR-21-CE32-0003). J.M.S.-D. acknowledges the support from NASA for UConn’s Ecological Modeling Institute (no. 80NSSC 22K0883).
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J.B., J.-C.G. and J.M.S.-D. conceived this article. J.B. was responsible for analysis, visualization, methodology and software. J.-C.G. and J.M.S.-D. undertook supervision and funding acquisition. J.B. wrote the manuscript with contributions from all co-authors.
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Extended data
Extended Data Fig. 1 Histogram of the randomized thermophilization values of the 80 ecoregions.
Results of the 200 iterations of the random thermal optimum model (thermal optima randomly assigned to the species). In this figure, the runs are not averaged: the 80 ecoregions randomized 200 times are displayed. The average values of thermophilization, Δβ-diversity and their contribution of the original dataset are displayed.
Extended Data Fig. 2 Thermophilization and Δβ-diversity change in lowland, mountain and Mediterranean ecoregions.
Thermophilization (a) and Δβ-diversity (b) in lowland, mountain and Mediterranean ecoregion clusters (Extended Data Fig. 1). Lowland (8,271 pairs, 45 ecoregions), mountain (4,116 pairs, 29 ecoregions), Mediterranean (377 pairs, 6 ecoregions). Each dot represents the values of one of the 80 ecoregions (ntot = 80).
Extended Data Fig. 3 Subsequent partitioning, into ‘rare’ and ‘common’ components, of the Thermophilization and Δβ-diversity change.
Partitioning of the data presented in Fig. 3. The contributions to a) thermophilization (°C decade−1) and b) Δβ-diversity (unitless) were partitioned on the basis of species declining or increasing in occurrences, of their thermal optimum relatively to their ecoregion and whether these species were rare (baseline occurrences < 10% of the plots) or common (baseline occurrences > 10% of the plots). Each dot represents the values of one of the 80 ecoregions (ntot = 80). The dashed grey line delineates the colonization and extinction components. The mean of each component is displayed. White dot; mean value of the thermophilization null model. The statistical difference between the null model value and the original dataset, obtained with a two-sided Wilcoxon test, is also displayed: p < 0.05 (*), p < 0.01 (**), p < 0.001 (***). Exact P values are available in Supplementary Table 1. Boxes; 25th centile, median and 75th centile; whiskers, 1.5 times the interquartile range.
Extended Data Fig. 4 Map of the sampled ecoregions and display of the plots of one ecoregion.
(a) Map of the 86 forest ecoregions of France, with a coloured gradient representing the number of plot pairs. Three main biomes (lowland, Mediterranean, mountain) cluster different ecoregions delineated with bold black lines. The clusters without a label are mountain ecoregions. The zoomed ecoregion in (b) is outlined in red in (a). (b) Example of the plot pair sampling design, with NFI plot localization. Some plots may overlap. Green, forested areas. Basemap credits82.
Supplementary information
Supplementary Information
Supplementary Tables 1–4, Figs. 1 and 2 and equations.
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Borderieux, J., Gégout, JC. & Serra-Diaz, J.M. Extinction drives recent thermophilization but does not trigger homogenization in forest understorey. Nat Ecol Evol 8, 695–704 (2024). https://doi.org/10.1038/s41559-024-02362-3
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DOI: https://doi.org/10.1038/s41559-024-02362-3