Abstract

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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References

  1. 1.

    Ricklefs, R. E. Disintegration of the ecological community. Am. Nat. 172, 741–750 (2008).

  2. 2.

    Pärtel, M., Bennett, J. A. & Zobel, M. Macroecology of biodiversity: disentangling local and regional effects. New. Phytol. 211, 404–410 (2016).

  3. 3.

    Kier, G. et al. Global patterns of plant diversity and floristic knowledge. J. Biogeogr. 32, 1107–1116 (2005).

  4. 4.

    Šímová, I. et al. Global species-energy relationship in forest plots: role of abundance, temperature and species climatic tolerances. Glob. Ecol. Biogeogr. 20, 842–856 (2011).

  5. 5.

    Götzenberger, L. et al. Ecological assembly rules in plant communities: approaches, patterns and prospects. Biol. Rev. Camb. Philos. Soc. 87, 111–127 (2012).

  6. 6.

    Reich, P. B., Frelich, L. E., Voldseth, R. A., Bakken, P. & Adair, E. C. Understorey diversity in southern boreal forests is regulated by productivity and its indirect impacts on resource availability and heterogeneity. J. Ecol. 100, 539–545 (2012).

  7. 7.

    Lessard, J.-P., Belmaker, J., Myers, J., Chase, J. M. & Rahbek, C. Inferring local ecological processes amid species pool influences. Trends Ecol. Evol. 27, 600–607 (2012).

  8. 8.

    Diamond, J. M. in Ecology and Evolution of Communities (eds. Cody, M. L. C. & Diamond, J. M.) 342–444 (Harvard Univ. Press, Cambridge, MA, 1975).

  9. 9.

    Eriksson, O. The species-pool hypothesis and plant community diversity. Oikos 68, 371–374 (1993).

  10. 10.

    Zobel, M. The relative role of species pools in determining plant species richness: an alternative explanation of species coexistence? Trends Ecol. Evol. 2, 266–269 (1997).

  11. 11.

    Cornell, H. V. & Harrison, S. P. What are species pools and when are they important? Annu. Rev. Ecol. Evol. Syst. 45, 45–67 (2014).

  12. 12.

    Herben, T. Correlation between richness per unit area and the species pool cannot be used to demonstrate the species pool effect. J. Veg. Sci. 11, 123–126 (2000).

  13. 13.

    Zobel, M. The species pool concept as a framework for studying patterns of plant diversity. J. Veg. Sci. 27, 8–18 (2016).

  14. 14.

    Mittelbach, G. G. & Schemske, D. W. Ecological and evolutionary perspectives on community assembly. Trends Ecol. Evol. 30, 241–247 (2015).

  15. 15.

    Pärtel, M., Szava-Kovats, R. & Zobel, M. Community completeness: linking local and dark diversity within the species pool concept. Folia Geobot. 48, 307–317 (2013).

  16. 16.

    Ricklefs, R. E. Community diversity: relative roles of local and regional processes. Science 235, 167–171 (1987).

  17. 17.

    Pither, J. & Aarssen, L. W. The evolutionary species pool hypothesis and patterns of freshwater diatom diversity along a pH gradient. J. Biogeogr. 32, 503–513 (2005).

  18. 18.

    Pärtel, M. Local plant diversity patterns and ecolutionary history at the reigonal scale. Ecology 83, 2361–2366 (2002).

  19. 19.

    Zobel, M. et al. The formation of species pools: historical habitat abundance affects current local diversity. Glob. Ecol. Biogeogr. 20, 251–259 (2011).

  20. 20.

    Scheiner, S. M. et al. The underpinnings of the relationship of species richness with space and time. Ecol. Monogr. 81, 195–213 (2011).

  21. 21.

    Brown, J. H. & Nicoletto, P. F. Spatial scaling of species composition: body masses of north american land mammals. Am. Nat. 138, 1478–1512 (1991).

  22. 22.

    Graves, G. R. & Rahbek, C. Source pool geometry and the assembly of continental avifaunas. Proc. Natl. Acad. Sci. USA 102, 7871–7876 (2005).

  23. 23.

    Fraser, L. H. et al. Response to comment on ‘worldwide evidence of a unimodal relationship between productivity and plant species richness’. Science 350, 1177–1178 (2016).

  24. 24.

    Jiménez-Alfaro, B., Marcenò, C., Bueno, Á., Gavilán, R. & Obeso, J. R. Biogeographic deconstruction of alpine plant communities along altitudinal and topographic gradients. J. Veg. Sci. 25, 160–171 (2014).

  25. 25.

    Horvat, V., Biurrun, I. & García-Mijangos, I. Herb layer in silver fir-beech forests in the western Pyrenees: does management affect species diversity? For. Ecol. Manag. 385, 87–96 (2017).

  26. 26.

    Paillet, Y. et al. Biodiversity differences between managed and unmanaged forests: meta-analysis of species richness in Europe. Conserv. Biol. 24, 101–112 (2010).

  27. 27.

    Magri, D. Patterns of post-glacial spread and the extent of glacial refugia of European beech (Fagus sylvatica). J. Biogeogr. 35, 450–463 (2008).

  28. 28.

    Fukami, T. Historical contingency in community assembly: integrating niches, species pools, and priority effects. Annu. Rev. Ecol. Evol. Syst. 46, 1–23 (2015).

  29. 29.

    Willner, W., Di Pietro, R. & Bergmeier, E. Phytogeographical evidence for post-glacial dispersal limitation of European beech forest species. Ecography 32, 1011–1018 (2009).

  30. 30.

    Sabatini, F. M., Jiménez-Alfaro, B., Burrascano, S., Lora, A. & Chytrý, M. Beta-diversity of Central European forests decreases along an elevational gradient due to the variation in local community assembly processes. Ecography https://doi.org/10.1111/ecog.02809 (2017).

  31. 31.

    Stein, A., Gerstner, K. & Kreft, H. Environmental heterogeneity as a universal driver of species richness across taxa, biomes and spatial scales. Ecol. Lett. 17, 866–880 (2014).

  32. 32.

    Gavin, D. G. et al. Climate refugia: joint inference from fossil records, species distribution models and phylogeography. New. Phytol. 204, 37–54 (2014).

  33. 33.

    Wohlgemuth, T., Moser, B., Brändli, U.-B., Kull, P. & Schütz, M. Diversity of forest plant species at the community and landscape scales in Switzerland. Plant Biosyst. 142, 604–613 (2008).

  34. 34.

    Ujházyová, M. et al. Diversity of beech forest vegetation in the Eastern Alps, Bohemian Massif and the Western Carpathians. Preslia 88, 435–457 (2016).

  35. 35.

    Ewald, J. The calcareous riddle: why are there so many calciphilous species in the Central European flora? Folia Geobot. 38, 357–366 (2003).

  36. 36.

    Biurrun, I., Campos, J. A., García-Mijangos, I., Herrera, M. & Loidi, J. Floodplain forests of the Iberian Peninsula: vegetation classification and climatic features. Appl. Veg. Sci. 19, 336–354 (2016).

  37. 37.

    Franklin, J., Serra-Diaz, J. M., Syphard, A. D. & Regan, H. M. Big data for forecasting the impacts of global change on plant communities. Glob. Ecol. Biogeogr. 26, 6–17 (2017).

  38. 38.

    Härdtle, W., von Oheimb, G. & Westphal, C. The effects of light and soil conditions on the species richness of the ground vegetation of deciduous forests in northern Germany (Schleswig-Holstein). For. Ecol. Manag. 182, 327–338 (2003).

  39. 39.

    Tzedakis, P. C., Lawson, I. T., Frogley, M. R., Hewitt, G. M. & Preece, R. C. Buffered tree population changes in a Quaternary refugium: evolutionary implications. Science 297, 2044–2047 (2002).

  40. 40.

    Karger, D. N. et al. Delineating probabilistic species pools in ecology and biogeography. Glob. Ecol. Biogeogr. 25, 489–501 (2016).

  41. 41.

    Gerhold, P., Cahill, J. F., Winter, M., Bartish, I. V. & Prinzing, A. Phylogenetic patterns are not proxies of community assembly mechanisms (they are far better). Funct. Ecol. 29, 600–614 (2015).

  42. 42.

    Wiser, S. K. Achievements and challenges in the integration, reuse and synthesis of vegetation plot data. J. Veg. Sci. 27, 868–879 (2016).

  43. 43.

    Lehsten, D. et al. Modelling the Holocene migrational dynamics of Fagus sylvatica L. and Picea abies (L.) H. Karst. Glob. Ecol. Biogeogr. 23, 658–668 (2014).

  44. 44.

    Willner, W. et al. Classification of European beech forests: a Gordian Knot? Appl. Veg. Sci. 20, 494–512 (2017).

  45. 45.

    Chytrý, M. et al. European Vegetation Archive (EVA): an integrated database of European vegetation plots. Appl. Veg. Sci. 19, 173–180 (2016).

  46. 46.

    Hennekens, S. M. & Schaminée, J. H. J. TURBOVEG, a comprehensive data base management system for vegetation data. J. Veg. Sci. 12, 589–591 (2001).

  47. 47.

    Tichý, L. JUICE, software for vegetation classification. J. Veg. Sci. 13, 451–453 (2002).

  48. 48.

    Houston Durrant, T., de Rigo, D. & Caudullo, G. in European Atlas of Forest Tree Species (eds. San-Miguel-Ayanz, J., de Rigo, D., Caudullo, G., Houston Durrant, T. & Mauri, A.) 94–95 (Publications Office of the EU, Luxembourg, 2016).

  49. 49.

    Elith, J. et al. A statistical explanation of MaxEnt for ecologists. Divers. Distrib. 17, 43–57 (2011).

  50. 50.

    Magyari, E. K. et al. Late Pleniglacial vegetation in eastern-central Europe: are there modern analogues in Siberia? Quat. Sci. Rev. 95, 60–79 (2014).

  51. 51.

    Colwell, R., Mao, C. & Chang, J. Interpolating, extrapolating, and comparing incidence-based species accumulation curves. Ecology 85, 2717–2727 (2004).

  52. 52.

    Ellenberg, H. et al. Zeigerwerte von Pflanzen in Mitteleuropa. Scr. Geobot. 18, 1–248 (1991).

  53. 53.

    Zelený, D. & Schaffers, A. P. Too good to be true: pitfalls of using mean Ellenberg indicator values in vegetation analyses. J. Veg. Sci. 23, 419–431 (2012).

  54. 54.

    Wildi, O. Why mean indicator values are not biased. J. Veg. Sci. 27, 40–49 (2016).

  55. 55.

    Mayor, M. Ecología de la Fflora y Vegetación del Principado de Asturias (Real Instituto de Estudios Asturianos, Oviedo, 1999).

  56. 56.

    Pignatti, S. Valori di bioindicazione delle piante vascolari della flora d’Italia. Braun-Blanquetia 39, 1–97 (2005).

  57. 57.

    Elith, J., Leathwick, J. R. & Hastie, T. A working guide to boosted regression trees. J. Anim. Ecol. 77, 802–813 (2008).

  58. 58.

    Grace, J. B. Structural Equation Modeling and Natural Systems (Cambridge Univ. Press, Cambridge, UK, 2006).

  59. 59.

    Lefcheck, J. S. piecewiseSEM: piecewise structural equation modelling in R for ecology, evolution, and systematics. Methods Ecol. Evol. 7, 573–579 (2016).

  60. 60.

    Kissling, W. D. & Carl, G. Spatial autocorrelation and the selection of simultaneous autoregressive models. Glob. Ecol. Biogeogr. 17, 59–71 (2008).

  61. 61.

    Bivand, R., Hauke, J. & Kossowski, T. Computing the Jacobian in Gaussian spatial autoregressive models: an illustrated comparison of available methods. Geogr. Anal. 45, 150–179 (2013).

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Acknowledgements

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

Affiliations

  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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Contributions

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.

Corresponding author

Correspondence to Borja Jiménez-Alfaro.

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https://doi.org/10.1038/s41559-017-0462-6