Skip to main content

Thank you for visiting You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser (or turn off compatibility mode in Internet Explorer). In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript.

Levels of forest ecosystem services depend on specific mixtures of commercial tree species


Global and local ecosystem change resulting in diversity loss has motivated efforts to understand relationships between species diversity and ecosystem services. However, it is unclear how such a general understanding can inform policies for the management of ecosystem services in production systems, because these systems are primarily used for food or fibre, and are rarely managed for the conservation of species diversity. Here, using data from a nationwide forest inventory covering an area of 230,000 km2, we show that relative abundances of commercial tree species in mixed stands strongly influence the potential to provide ecosystem services. The mixes provided higher levels of ecosystem services compared to respective plant monocultures (overyielding or transgressive overyielding) in 35% of the investigated cases, and lower (underyielding) in 9% of the cases. We further show that relative abundances, not just species richness per se, of specific tree-species mixtures affect the potential of forests to provide multiple ecosystem services, which is crucial information for policy and sustainable forest management.

This is a preview of subscription content, access via your institution

Relevant articles

Open Access articles citing this article.

Access options

Buy article

Get time limited or full article access on ReadCube.


All prices are NET prices.

Fig. 1: Transgressive overyielding was found for several, but not all, forest ecosystem services at the national scale.
Fig. 2: Transgressive overyielding, overyielding or underyielding of forest ecosystem services was found in 44% of the investigated cases.

Data availability

The data that support the findings of this study are available from the corresponding author upon reasonable request.


  1. Daily, G. C. Nature’s Services: Societal Dependence on Natural Ecosystems (Island Press, Washington DC, 1997).

  2. Costanza, R. et al. Twenty years of ecosystem services: how far have we come and how far do we still need to go?. Ecosyst. Serv. 28, 1–16 (2017).

    Article  Google Scholar 

  3. Mace, G. M., Norris, K. & Fitter, A. H. Biodiversity and ecosystem services: a multilayered relationship. Trends Ecol. Evol. 27, 19–26 (2012).

    Article  Google Scholar 

  4. Cardinale, B. J. et al. Biodiversity loss and its impacts on humanity. Nature 486, 59–67 (2012).

    Article  CAS  Google Scholar 

  5. Balvanera, P. et al. Linking biodiversity and ecosystem services: current uncertainties and the necessary next steps. Bioscience 64, 49–57 (2014).

    Article  Google Scholar 

  6. Diaz, S. et al. The IPBES conceptual framework – connecting nature and people. Curr. Opin. Environ. Sustain. 14, 1–16 (2015).

    Article  Google Scholar 

  7. Larigauderie, A. et al. Biodiversity and ecosystem services science for a sustainable planet. The DIVERSITAS vision for 2012-20. Curr. Opin. Environ. Sustain. 4, 101–105 (2012).

    Article  Google Scholar 

  8. Gamfeldt, L. et al. Higher levels of multiple ecosystem services are found in forests with more tree species. Nat. Commun. 4, 1340 (2013).

    Article  Google Scholar 

  9. Grime, J. P. Biodiversity and ecosystem function: the debate deepens. Science 277, 1260–1261 (1997).

    Article  CAS  Google Scholar 

  10. Huston, M. A. Hidden treatments in ecological experiments: re-evaluating the ecosystem function of biodiversity. Oecologia 110, 449–460 (1997).

    Article  Google Scholar 

  11. Cardinale, B. J. et al. The functional role of producer diversity in ecosystems. Am. J. Bot. 98, 572–592 (2011).

    Article  Google Scholar 

  12. Pretzsch, H. et al. Comparison between the productivity of pure and mixed stands of Norway spruce and European beech along an ecological gradient. Ann. For. Sci. 67, 712 (2010).

    Article  Google Scholar 

  13. Toïgo, M. et al. Overyielding in mixed forests decreases with site productivity. J. Ecol. 103, 502–512 (2015).

    Article  Google Scholar 

  14. Felton, A. et al. Replacing monocultures with mixed-species stands: ecosystem service implications of two production forest alternatives in Sweden. Ambio 45, S124–S139 (2016).

    Article  Google Scholar 

  15. Lu, H., Mohren, G. M. J., den Ouden, J., Goudiaby, V. & Sterck, F. J. Overyielding of temperate mixed forest occurs in evergreen-deciduous but not in deciduous-deciduous species mixtures over time in the Netherlands. For. Ecol. Manage. 376, 321–332 (2016).

    Article  Google Scholar 

  16. Pretzsch, H. in Forest Diversity and Function 41–64 (Springer Berlin, Berlin, 2005).

  17. de Wit, C. T. On competition. Versl. Landbouwk. Onderzoek 66, 1–82 (1960).

    Google Scholar 

  18. Trenbath, B. R. Diversity or be damned? Ecologist 5, 76–83 (1975).

    Google Scholar 

  19. Vandermeer, J. H. The Ecology of Intercropping 1st edn (Cambridge Univ. Press, Cambridge, 1989).

  20. Schmid, B., Hector, A., Saha, P. & Loreau, M. Biodiversity effects and transgressive overyielding. J. Plant Ecol. 1, 95–102 (2008).

    Article  Google Scholar 

  21. Nyfeler, D. et al. Strong mixture effects among four species in fertilized agricultural grassland led to persistent and consistent transgressive overyielding. J. Appl. Ecol. 46, 683–691 (2009).

    Article  Google Scholar 

  22. Guenay, Y., Ebeling, A., Steinauer, K., Weisser, W. W. & Eisenhauer, N. Transgressive overyielding of soil microbial biomass in a grassland plant diversity gradient. Soil Biol. Biochem. 60, 122–124 (2013).

    Article  CAS  Google Scholar 

  23. Finney, D. M., White, C. M. & Kaye, J. P. Biomass production and carbon/nitrogen ratio influence ecosystem services from cover crop mixtures. Agron. J. 108, 39–52 (2016).

    Article  CAS  Google Scholar 

  24. Jucker, T., Bouriaud, O. & Coomes, D. A. Crown plasticity enables trees to optimized canopy packing in mixed-species forests. Funct. Ecol. 29, 1078–1086 (2015).

    Article  Google Scholar 

  25. Kahmen, A., Renker, C., Unsicker, S. B. & Buchmann, N. Niche complementarity for nitrogen: an explanation for the biodiversity and ecosystem functioning relationship? Ecology 87, 1244–1255 (2006).

    Article  Google Scholar 

  26. Williams, L. J., Paquette, A., Cavender-Bares, J., Messier, C. & Reich, P. B. Spatial complementarity in tree crowns explains overyielding in species mixtures. Nat. Ecol. Evol. 1, 0063 (2017).

    Article  Google Scholar 

  27. Paquette, A. & Messier, C. The effect of biodiversity on tree productivity: from temperate to boreal forests. Glob. Ecol. Biogeogr. 20, 170–180 (2011).

    Article  Google Scholar 

  28. Dosi, G. Technological paradigms and technological trajectories: a suggested interpretation of the determinants and directions of technical change. Res. Policy 11, 147–162 (1982).

    Article  Google Scholar 

  29. Moen, J. et al. Eye on the Taiga: removing global policy impediments to safeguard the boreal forest. Conserv. Lett. 7, 408–418 (2014).

    Article  Google Scholar 

  30. Blennow, K. Risk management in Swedish forestry – policy formation and fulfilment of goals. J. Risk Res. 11, 237–254 (2008).

    Article  Google Scholar 

  31. Axelsson, A.-L. et al. in National Forest Inventories—Pathways for Common Reporting (eds Tomppo, E. et al.) 541–553 (Springer Netherlands, Dordrecht, 2010).

  32. Haines-Young, R. & Potschin, M. B. Common International Classification of Ecosystem Services (CICES) V5.1 and Guidance on the Application of the Revised Structure (Fabis Consulting Ltd., Nottingham, 2018);

  33. Díaz, S. et al. Assessing nature’s contributions to people. Science 359, 270–272 (2018).

    Article  Google Scholar 

  34. Marklund, L. G. Biomass Functions for Pine, Spruce and Birch in Sweden (Department of Forest Survey, Swedish University of Agricultural Sciences, Umeå, 1988).

  35. Petersson, H. & Ståhl, G. Functions for below-ground biomass of Pinus sylvestris, Picea abies, Betula pendula and Betula pubescens in Sweden. Scand. J. For. Res. 21, 84–93 (2006).

  36. Miina, J., Hotanen, J. P. & Salo, K. Modelling the abundance and temporal variation in the production of bilberry (Vaccinium myrtillus L.) in Finnish mineral soil forests. Silva. Fenn. 43, 577–593 (2009).

    Article  Google Scholar 

  37. Cederlund, G., Ljungqvist, H., Markgren, G. & Stålfelt, F. Foods of moose and roe deer at Grimsö in central Sweden—results of rumen content analysis. Swed. Wildlife Res. 11, 167–247 (1980).

    Google Scholar 

  38. Hanley, T. A. A nutritional view of understanding and complexity in the problem of diet selection by deer (Cervidae). Oikos 79, 209–218 (1997).

    Article  CAS  Google Scholar 

  39. Boman, M., Mattsson, L., Ericsson, G. R. & Kriström, B. Moose hunting values in Sweden now and two decades ago: the Swedish hunters revisited. Environ. Resour. Econ. 50, 515–530 (2011).

    Article  Google Scholar 

  40. Agriculture and Forestry in Sweden Since 1900 – A Cartographic Description (ed Jansson, U.) 232 (The Royal Swedish Academy of Agriculture and Forestry, Stockholm, 2011).

  41. Warfvinge, P. & Sverdrup, H. Critical Loads of Acidity to Swedish Forest Soils: Methods, Data and Results (Department of Chemical Engineering II, Lund University, Lund, 1995).

  42. von Arx, G., Dobbertin, M. & Rebetez, M. Spatio-temporal effects of forest canopy on understory microclimate in a long-term experiment in Switzerland. Agric. For. Meteorol. 166-167, 144–155 (2012).

    Article  Google Scholar 

  43. Lieffers, V. J., Messier, C., Stadt, K. J., Gendron, F. & Comeau, P. G. Predicting and managing light in the understory of boreal forests. Can. J. For. Res. 29, 796–811 (1999).

    Article  Google Scholar 

  44. Strengbom, J., Axelsson, E. P., Lundmark, T. & Nordin, A. Trade-offs in the multi-use of managed boreal forests. J. Appl. Ecol. 55, 958–966 (2018).

    Article  Google Scholar 

  45. Joliffe, P. A. The replacement series. J. Ecol. 88, 371–385 (2000).

    Article  Google Scholar 

  46. Bates, D. et al. Fitting linear mixed-effect models using lme4. J. Stat. Softw. 67, 1–48 (2015).

    Article  Google Scholar 

  47. R Core Team R: A Language and Environment for Statistical Computing (R Foundation for Statistical Computing, 2017);

Download references


This work was supported by grant no. 942–2015–1090 from the Swedish research council Formas.

Author information

Authors and Affiliations



All authors planned the study. L.G. and M.J. managed the data. M.J. performed the statistical analyses with support from T.S., and M.J. led the writing of the text with input from all authors.

Corresponding author

Correspondence to Micael Jonsson.

Ethics declarations

Competing interests

The authors declare no competing interests.

Additional information

Publisher’s note: Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Supplementary information

Supplementary Information

Supplementary Figures 1–7 and Supplementary Tables 1–9.

Reporting Summary

Rights and permissions

Reprints and Permissions

About this article

Verify currency and authenticity via CrossMark

Cite this article

Jonsson, M., Bengtsson, J., Gamfeldt, L. et al. Levels of forest ecosystem services depend on specific mixtures of commercial tree species. Nature Plants 5, 141–147 (2019).

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI:

This article is cited by


Quick links

Nature Briefing

Sign up for the Nature Briefing newsletter — what matters in science, free to your inbox daily.

Get the most important science stories of the day, free in your inbox. Sign up for Nature Briefing