TREE DIVERSITY EFFECTS ON FOREST PRODUCTIVITY: DISENTANGLING THE EFFECTS
OF TREE SPECIES ADDITION VS. SUBSTITUTION
Abstract
1. Mixture effect on stand productivity is usually apprehended through a
substitutive approach, whereby productivity in mixed stands is compared
to productivity in monocultures, at equivalent stand density. This
approach has proved that in many cases mixed stands perform better than
monospecific forests, however, we do not yet have a solid theory about
species behaviour in the mixture or even guidelines for combining
species. The addition of a second tree species to an existing
mono-specific stand has received much less consideration. Yet, this
approach has the potential to separate the facilitation effect from the
complementarity effect. 2. We compared the effect of tree species
substitution vs. addition on the productivity of maritime pine and
silver birch in a young tree diversity experiment implemented in 2008 in
SW France. 3. Substituting pines with birches to create two-species
mixtures resulted in an increase of tree productivity at stand level
beyond what was expected from monocultures (i.e., overyielding). In
contrast, creating mixture through the addition of birches to pine
stands had no effect on the maritime pine stand productivity
(transgressive mixture effect not significant). This absence of effect
is produced by two distinct density-dependence responses at an
individual level. 4. Our results allow clarifying the cases in which a
mixed stand can be considered as an alternative to a monoculture of a
productive species. In particular, the addition of a pioneer and soil
low-demanding species during young developmental stages is a possibility
to diversify the stand and potentially to increase ecosystem services
without altering the productivity of the target species.