Disentangling the effects of biomass and productivity in plant competition.

The relationship between competition and productivity in plant communities is unclear, and this is likely to be due to (1) a confusion in the literature between productivity and biomass, (2) the lack of studies assessing variation in competition in all combinations of biomass and productivity. We assessed the outcome of plant-plant interactions by removing the neighbors around five focal species in 14 herbaceous communities with contrasting biomasses and productivities: meadows with high biomass and productivity, heathlands with high biomass and low productivity, understory communities of deciduous forests with low biomass and high productivity and calcareous grasslands with low biomass and low productivity. Competition intensity was quantified with the relative interaction index (RII) calculated for both survival and growth of the transplanted targets assessed with the increase in leaf number. To examine which traits better explain variation in competition and what drives variation in diversity, we also quantified litter decomposition rate, species composition and diversity and six morphological traits related to plant size and growth rate for eight dominant species of each community. Our main questions were: (1) Is competition mostly related to biomass or productivity? (2) Which traits of the community dominants better explain variation in competition? (3) Is variation in competition and related traits correlated with variation in diversity? Competition for survival significantly increased with increasing community biomass (but not productivity). In addition, competition for survival increased with the size traits and competitive effects of the dominant species of the communities, whereas diversity decreased. Competition for growth also increased with increasing productivity, but only for high-biomass communities. Additionally, the increase in competition for growth with increasing soil fertility, as measured with litter decomposition rate, was only due to an increase in target growth in plots without neighbors and was unrelated to community competitive effects and species diversity. The results of our study illustrate how the confusion between productivity and biomass could have contributed to the long-standing debate on variation in competition along productivity gradients and its consequence for diversity.