Good things take time—Diversity effects on tree growth shift from negative to positive during stand development in boreal forests

Abstract

Long-term grassland biodiversity experiments have shown that diversity effects on productivity tend to strengthen through time, as complementarity among coexisting species increases. But it remains less clear whether this pattern also holds for other ecosystems such as forests, and if so why. Here we explore whether diversity effects on tree growth change predictably during stand development in Finland's boreal forests. Using tree ring records from mature forests, we tested whether diameter growth trajectories of dominant tree species growing in mixture differed from those in monoculture. We then compared these results with data from the world's longest running tree diversity experiment, where the same combinations of species sampled in mature forests were planted in 1999. We found that diversity effects on tree growth strengthened progressively through time, only becoming significantly positive around 20 years after seedling establishment. This shift coincided with the period in which canopy closure occurs in these forests, at which time trees begin to interact and compete above-ground. These temporal trends were remarkably consistent across different tree species sampled in mature forests, and broadly matched growth responses observed in the much younger experimental plots. Synthesis. Our results mirror those from grassland ecosystems and suggest that canopy closure is a key phase for promoting niche complementarity in diverse tree communities. They also provide a series of testable hypotheses for the growing number of tree diversity experiments that have been established in recent years.