Contrasting patterns of tree species mixture effects on wood δ13C along an environmental gradient

  • G. de StreelEmail author
  • C. Collet
  • I. Barbeito
  • K. Bielak
  • A. Bravo-Oviedo
  • G. Brazaitis
  • L. Coll
  • L. Drössler
  • D. Forrester
  • M. Heym
  • M. Löf
  • M. Pach
  • H. Pretzsch
  • R. Ruiz-Peinado
  • J. Skrzyszewski
  • J. Stankevičiūtė
  • M. Svoboda
  • K. Verheyen
  • T. Zlatanov
  • D. Bonal
  • Q. Ponette
Original Paper


Establishing mixed-species stands is frequently proposed as a strategy to adapt forests to the increasing risk of water scarcity, yet contrasted results have been reported regarding mixing effects on tree drought exposure. To investigate the drivers behind the spatial and temporal variation in water-related mixing effects, we analysed the δ13C variation in 22-year tree ring chronologies for beech and pine trees sampled from 17 pure and mixed pine–beech stands across a large gradient of environmental conditions throughout Europe. In the pure stands, average δ13C values were lower for beech (−27.9‰ to −22.2‰) than for pine (−26.0‰ to −21.1‰), irrespective of site conditions. Decreasing SPEI values (calculated over June to September) were associated with an increase in δ13C for both species, but their effect was influenced by stand basal area for pine and site water availability for beech. Mixing did not change the temporal constancy of δ13C nor the tree reaction to a drought event, for any of the species. While the mixing effect (Δ δ13C = δ13C pure stands − δ13C mixed stands) was on average positive for beech and non-significant for pine across the whole gradient, this effect strongly differed between sites. For both species, mixing was not significant at extremely dry sites and positive at dry sites; on moderately wet sites, mixing was positive for beech and negative for pine; at sites with permanent water supply, no general patterns emerge for any of the species. The pattern of mixing effect along the gradient of water availability was not linear but showed threshold points, highlighting the need to investigate such relation for other combinations of tree species.


Stable carbon isotope composition Species mixture Fagus sylvatica L. Pinus sylvestris L. Drought 



The networking in this study has been supported by COST Action FP1206 EUMIXFOR. All contributors thank their national funding institutions to establish, measure and analyse data from the triplets. The main author obtained a PhD grant from the “Fonds National de la Recherche Scientifique” (FNRS-FRIA) and additional funding support from the Walloon forest service (Service Public de Wallonie—Département de la Nature et des Forêts) through the 5-year research programme “Accord-cadre de recherches et de vulgarisation forestières”. All authors thank the numerous persons who contributed to the fieldwork in the different sites across Europe. We also thank Maud Antoine who helped carry out the chemical analyses. Finally, we thank the two anonymous reviewers and the main editor who contributed to improving a previous version of this manuscript. We acknowledge the E-OBS dataset from the EU-FP6 project ENSEMBLES ( and the data providers in the ECA&D project (”.

Author’s contributions

GdS, DB and QP conceived the ideas and designed methodology; all authors contributed to data collection; GdS, DB and QP analysed the data and led the writing of the manuscript; all authors contributed to the drafts and gave the final approval for publication.

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.

Supplementary material

10342_2019_1224_MOESM1_ESM.docx (943 kb)
Supplementary material 1 (DOCX 942 kb)


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Copyright information

© Springer-Verlag GmbH Germany, part of Springer Nature 2019

Authors and Affiliations

  • G. de Streel
    • 1
    Email author
  • C. Collet
    • 2
  • I. Barbeito
    • 3
  • K. Bielak
    • 4
  • A. Bravo-Oviedo
    • 5
  • G. Brazaitis
    • 6
  • L. Coll
    • 7
  • L. Drössler
    • 8
  • D. Forrester
    • 9
  • M. Heym
    • 10
  • M. Löf
    • 3
  • M. Pach
    • 11
  • H. Pretzsch
    • 12
  • R. Ruiz-Peinado
    • 13
    • 14
  • J. Skrzyszewski
    • 11
  • J. Stankevičiūtė
    • 6
  • M. Svoboda
    • 15
  • K. Verheyen
    • 16
  • T. Zlatanov
    • 17
  • D. Bonal
    • 2
  • Q. Ponette
    • 1
  1. 1.Earth and Life InstituteUniversité catholique de LouvainLouvain-La-NeuveBelgium
  2. 2.Université de Lorraine, AgroParisTech, INRA, UMR SilvaNancyFrance
  3. 3.Southern Swedish Forest Research CentreSwedish University of Agricultural SciencesAlnarpSweden
  4. 4.Department of SilvicultureWarsaw University of Life SciencesWarsawPoland
  5. 5.Department of Biogeography and Global ChangeNational Museum of Natural Sciences, MNCN-CSICMadridSpain
  6. 6.Vytautas Magnus UniversityKaunasLithuania
  7. 7.Department of Agriculture and Forest Engineering – Forest Sciences Centre of Catalonia (CTFC)University of LleidaLleidaSpain
  8. 8.Institute of EcologyIlia State UniversityTbilisiGeorgia
  9. 9.Swiss Federal Institute of Forest, Snow and Landscape Research WSLBirmensdorfSwitzerland
  10. 10.Chair of Forest Growth and Yield Science, TUM School of Life Sciences WeihenstephanTechnical University of MunichFreisingGermany
  11. 11.Department of Silviculture, Institute of Forest Ecology and SilviculltureUniversity of Agriculture in KrakowKrakówPoland
  12. 12.Chair for Forest Growth and Yield ScienceTechnische Universität MünchenMunichGermany
  13. 13.Forest Research CentreINIAMadridSpain
  14. 14.iuFOR, Sustainable Forest Management Research InstituteUniversity of Valladolid - INIAPalenciaSpain
  15. 15.Faculty of Forestry and Wood SciencesCzech University of Life SciencesPraha 6 SuchdolCzech Republic
  16. 16.Forest and Nature Lab, Department of EnvironmentGhent UniversityMelle-GontrodeBelgium
  17. 17.Institute of Biodiversity and Ecosystem ResearchBulgarian Academy of SciencesSofiaBulgaria

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