Annals of Forest Science

, 75:101 | Cite as

Covariation between tree size and shade tolerance modulates mixed-forest productivity

  • Thomas Cordonnier
  • Thomas Bourdier
  • Georges Kunstler
  • Christian Piedallu
  • Benoît Courbaud
Research Paper
Part of the following topical collections:
  1. Mensuration and modelling for forestry in a changing environment


Key message

In tree communities, tree size inequality reduces productivity and interacts with tree shade tolerance to modulate stand productivity, with a higher productivity in stands where shade-intolerant species dominate shade-tolerant species in size.


Positive diversity–productivity relationships have been reported in different plant communities, including tree communities. These effects may be strongly related to both structural diversity and functional diversity, but also to their interactions if there is a non-random distribution of species functional characteristics among canopy layers.


We explore the relative effects on forest productivity of tree species diversity, tree size inequality, and species shade tolerance diversity, as well as the effect of the distribution of tree shade tolerance in the canopy.


We used 11,054 mixed-species forest plots from the French Forest Inventory (IGN) distributed throughout France (2006–2011). We analyzed the effects of species richness, shade tolerance diversity, and height inequality on forest plot productivity, represented by basal area annual increment over a period of 5 years, while controlling for first-order structure characteristics (basal area and quadratic mean diameter) and environmental factors (soil water budget and sum of growing degree days). Using the covariance between tree height and shade tolerance in mixed species canopies, we also explored the effect of the distribution of species’ shade tolerance among canopy layers.


The results showed a positive effect of species richness (effect size, 0.02) and a negative effect of height inequality (− 0.05) on mixed-forest productivity. We also showed that a negative covariance between shade tolerance and height (e.g., higher proportion of shade-tolerant species in lower height classes) increased productivity (0.01). Shade tolerance diversity did not affect productivity.


In tree communities, as shown previously in monospecific forest stands, tree size inequality reduces productivity. This effect is modulated by the distribution of shade tolerance among canopy layers. Previous studies on species diversity effect have generally overlooked the importance of the size structure and the size hierarchy of functional characteristics. These effects are, however, crucial and deserve to be explored in greater detail.


Species richness Size inequality Stand productivity Functional characteristics Gini index 



This article has been supported by the DISTIMACC project (BGF, no. Ecofor 2014–23) and the GIS-COOP “Cooperative for data on forest tree and stand growth”. LESSEM is part of Labex OSUG@2020 (ANR10 LABX56).

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.


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Copyright information

© INRA and Springer-Verlag France SAS, part of Springer Nature 2018

Authors and Affiliations

  1. 1.Univ. Grenoble Alpes, Irstea, UR LESSEMSaint-Martin-d’Hères cedexFrance
  2. 2.AgroParisTech, Inra, SilvaUniversité de LorraineNancyFrance

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