Mixing has limited impacts on the foliar nutrition of European beech and Scots pine trees across Europe

Géraud de Streel*, Christian Ammer, Peter Annighöfer, Ignacio Barbeito, Kamil Bielak, Andrés Bravo-Oviedo, Gediminas Brazaitis, Włodzimierz Buraczyk, Catherine Collet, Vaclav Hurt, Viktor Kurylyak, Jan den Ouden, Maciej Pach, Hans Pretzsch, Jerzy Skrzyszewski, Vit Sramek, Jolanta Stankevičiūtė, Katarina Strelcova, Miroslav Svoboda, Kris VerheyenTzvetan Zlatanov, Quentin Ponette

*Corresponding author for this work

Research output: Contribution to journalArticleAcademicpeer-review


Tree species-mixing has been suggested as one option to counteract the adverse effects of global change on tree mineral nutrition, yet the effect of mixing on nutrient availability remains poorly documented. We therefore analyzed the current foliar nutrient (N, P, K, Ca, Mg) quantities and ilr balances (isometric log transformed ratios between elements or groups of elements) for 261 European beech and 248 Scots pine trees from 15 sites, each consisting of one beech-pine mixed stand and the respective monocultures, across a gradient of environmental conditions in Europe. We hypothesized an overall positive effect of mixing on tree foliar nutrient content, and that this mixing effect would be stronger on nutrient-poor sites. Using linear mixed models and multivariate linear regression models, we first tested for the effects of species (beech/pine) and composition (pure/mixed) across all sites; we then investigated whether the species-mixing effect was related to site fertility. The nutrient composition of beech leaves and pine needles differed significantly for all ilr balances. For both species, significant mixing effects were detected for some nutrients and ilr balances; those effects, however, could not be consistently related to contrasted nutrient composition between species. For most nutrients and ilr balances, the mixing effect was influenced by the site nutritional status, but the pattern differed from expectation: absence or minor differences between monocultures and mixtures at the lower end of the chemical fertility gradient, and maximum differences in rich soils. The contrasting foliar nutrient composition of pine and beech trees and the site nutrient status only partly explained the mixing effects on tree mineral nutrition. Our results claim for a better understanding of nutrient-related mechanisms associated with complementarity and points towards the need to further expand the existing frameworks to account for the multivariate nature of tree nutrition.

Original languageEnglish
Article number118551
JournalForest Ecology and Management
Early online date10 Sep 2020
Publication statusE-pub ahead of print - 10 Sep 2020


  • Complementarity
  • Fagus sylvatica L.
  • Foliar nutrition
  • Pinus sylvestris L.
  • Species mixture

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