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Global patterns and drivers of tree diversity integrated across a continuum of spatial grains


Controversy remains over what drives patterns in the variation of biodiversity across the planet. The resolution is obscured by lack of data and mismatches in their spatial grain (scale), and by grain-dependent effects of the drivers. Here we introduce cross-scale models integrating global data on tree-species richness from 1,336 local forest surveys and 282 regional checklists, enabling the estimation of drivers and patterns of biodiversity across spatial grains. We uncover grain-dependent effects of both environment and biogeographic regions on species richness, with a striking positive effect of Southeast Asia at coarse grain that disappears at fine grains. We show that, globally, biodiversity cannot be attributed purely to environmental or regional drivers, as the regions are environmentally distinct even within a single latitudinal band. Finally, we predict global maps of biodiversity at local (plot-based) and regional grains, identifying areas of exceptional beta-diversity in China, East Africa and North America. By allowing the importance of drivers of diversity to vary with grain in a single model, our approach unifies disparate results from previous studies regarding environmental versus biogeographic predictors of biodiversity, and enables efficient integration of heterogeneous data.

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Data availability

All data and R codes used for the analyses are available under CC-BY 4.0 license in a GitHub repository at, which is also mirrored at figshare at Please note that if the data on species richness are reused, the original data sources should be credited.


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We thank D. Craven and I. Šímová for valuable advice, and H. Kreft, J. Coyle, R. Ricklefs, and S. Blowes for critical comments that greatly improved the manuscript. We acknowledge the support of the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig funded by the German Research Foundation (FZT 118).

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P.K. formalized the ideas, collated the data, performed the analyses, and led the writing. J.M.C. proposed the initial idea, contributed to its development, discussed the results, and contributed to the writing.

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Correspondence to Petr Keil.

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Supplementary Figures 1–13, Supplementary Table 1, Supplementary Methods, Supplementary Discussion and Supplementary References

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Keil, P., Chase, J.M. Global patterns and drivers of tree diversity integrated across a continuum of spatial grains. Nat Ecol Evol 3, 390–399 (2019).

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