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History and environment shape species pools and community diversity in European beech forests


A central hypothesis of ecology states that regional diversity influences local diversity through species-pool effects. Species pools are supposedly shaped by large-scale factors and then filtered into ecological communities, but understanding these processes requires the analysis of large datasets across several regions. Here, we use a framework of community assembly at a continental scale to test the relative influence of historical and environmental drivers, in combination with regional or local species pools, on community species richness and community completeness. Using 42,173 vegetation plots sampled across European beech forests, we found that large-scale factors largely accounted for species pool sizes. At the regional scale, main predictors reflected historical contingencies related to post-glacial dispersal routes, whereas at the local scale, the influence of environmental filters was predominant. Proximity to Quaternary refugia and high precipitation were the main factors supporting community species richness, especially among beech forest specialist plants. Models for community completeness indicate the influence of large-scale factors, further suggesting community saturation as a result of dispersal limitation or biotic interactions. Our results empirically demonstrate how historical factors complement environmental gradients to provide a better understanding of biodiversity patterns across multiple regions.

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We are grateful to data contributors of the European Vegetation Archive, and also S. Hennekens and I. Knollová for their support with data management. B.J.-A. was supported by the project 'Employment of Best Young Scientists for International Cooperation Empowerment' (CZ.1.07/2.3.00/30.0037) co-financed by the European Social Fund and the state budget of the Czech Republic, and by the German Centre for Integrative Biodiversity Research funded by the German Research Foundation. M.C. and L.T. were supported by the Czech Science Foundation (Centre of Excellence PLADIAS, 14-36079 G). J.-C.S. considers this work a contribution to his VILLUM Investigator project funded by VILLUM FONDEN (grant 16549).

Author information

B.J.-A. designed the study with support of M.C. and J.-C.S. B.J.-A. prepared the data and wrote the manuscript. M.G. performed the statistical analyses. The other co-authors contributed data, interpreted the results and commented on the final version. The first six authors are ordered by relative contribution, the rest are ordered alphabetically.

Competing interests

The authors declare no competing financial interests.

Correspondence to Borja Jiménez-Alfaro.

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Fig. 1: A multi-regional framework for community assembly.
Fig. 2: The number of plant species in European beech forests.
Fig. 3: Path coefficients of structural equation models indicating relationships between large-scale environmental and historical factors, local habitat conditions, species pools and community species richness across European beech forests.