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


  1. German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, Leipzig, Germany

    • Borja Jiménez-Alfaro
    •  & Ute Jandt
  2. Institute of Biology/Geobotany and Botanical Garden, Martin Luther University Halle-Wittenberg, Halle, Germany

    • Borja Jiménez-Alfaro
    •  & Ute Jandt
  3. Department of Botany and Zoology, Masaryk University, Brno, Czech Republic

    • Borja Jiménez-Alfaro
    • , Milan Chytrý
    •  & Lubomír Tichý
  4. Section for Ecoinformatics and Biodiversity, Department of Bioscience, Aarhus University, Aarhus, Denmark

    • Marco Girardello
    •  & Jens-Christian Svenning
  5. cE3c–Centre for Ecology, Evolution and Environmental Changes, Azorean Biodiversity Group, Angra do Heroísmo, Portugal

    • Marco Girardello
  6. Center for Biodiversity Dynamics in a Changing World (BIOCHANGE), Aarhus University, Aarhus, Denmark

    • Jens-Christian Svenning
  7. VINCA - Vienna Institute for Nature Conservation and Analyses, Wien, Austria

    • Wolfgang Willner
  8. Department of Botany and Biodiversity Research, University of Vienna, Vienna, Austria

    • Wolfgang Willner
  9. LERFoB, INRA, AgroParisTech, Nancy, France

    • Jean-Claude Gégout
  10. Department of Environmental Biology, Sapienza University of Rome, Roma, Italy

    • Emiliano Agrillo
  11. Department of Plant Biology and Ecology, University of Basque Country UPV/EHU, Bilbao, Spain

    • Juan Antonio Campos
  12. Department of Vegetation Ecology, Botanical Garden, University of Wroclaw, Wrocław, Poland

    • Zygmunt Kącki
  13. ZRC SAZU, Institute of Biology, Ljubljana, Slovenia

    • Urban Šilc
  14. Institute of Forest Ecology, Slovak Academy of Sciences, Zvolen, Slovakia

    • Michal Slezák
  15. Plant Science and Biodiversity Center, Institute of Botany, Slovak Academy of Sciences, Bratislava, Slovakia

    • Michal Slezák
  16. Department of Botany, School of Biology, Aristotle University of Thessaloniki, Thessaloniki, Greece

    • Ioannis Tsiripidis
  17. Alexandru Borza Botanical Garden, Babeș-Bolyai University, Cluj-Napoca, Romania

    • Pavel Dan Turtureanu
  18. Department of Applied Ecology, Faculty of Ecology and Environmental Sciences, Technical University in Zvolen, Zvolen, Slovakia

    • Mariana Ujházyová
  19. WSL Swiss Federal Research Institute, Birmensdorf, Switzerland

    • Thomas Wohlgemuth


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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.

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The authors declare no competing financial interests.

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Correspondence to Borja Jiménez-Alfaro.

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