Abstract
The effect of species diversity on forest productivity and its temporal stability is known to be species-, climateand site-dependent and is mostly apprehended through stem diameter. Therefore, it remains largely unknown whether the mixture effect on the growth of tree crowns is similar to its effect on the growth of tree diameter. However, it is commonly accepted that changes in crown architecture are an important component of tree response to tree species diversity. Moreover, the mixture effect on species is often asymmetric, i.e. the effect of a species A on a species B is not equal to the effect of species B on A. It then appears that considering the effects of both species mixture and climate on shoot growth could contrast the results coming mainly from stem growth. We studied the effects of tree species mixture and temperature on the annual growth of shoots and basal area of stems in Fagus sylvatica-Quercus pubescens and Fagus sylvatica-Abies alba stands along a Mediterranean-Alpine gradient, for four years in five sites. The sample design was organized in 10 triplets: four triplets of mono-and bispecific plots of Quercus pubescens and Fagus sylvatica and six triplets of mono-and bi-specific plots of Abies alba and Fagus sylvatica along an altitudinal gradient ranging from 725 m to 1431 m. We found that the mixture effect on annual shoot volume increment (SVI) and on basal area increment (BAI) was asymmetrical in seven out of 10 cases and not significant in the three remaining cases. Mixture effect on SVI ranked from − 56% to 157% and on BAI it ranked from − 40% to 252%. Eventually we found that mixture effect was dependent on the type of limiting factor for growth, with at the driest sites a predominance of competition effects and at the coldest site a positive mixture effect on the two species studied. Branch growth appears as a variable that can be at least as informative as radial growth regarding the tree response to species interactions. This implies that considering only stem diameter in the diversity-productivity relationship can lead to biased conclusions on the global mixture effect on tree growth, which calls for a comprehensive approach of the tree response to tree species diversity. Our results are discussed in the light of the species stress tolerances and strategies to cope with competition.