Forest diversity-productivity relationships have been intensively investigated in recent decades. However, few studies have considered the interplay between species and structural diversity in driving productivity. We analyzed these factors using data from 52 permanent plots in southwestern Germany with more than 53,000 repeated tree measurements. We used basal area increment as a proxy for productivity and hypothesized that: (1) structural diversity would increase tree and stand productivity, (2) diversity-productivity relationships would be weaker for species diversity than for structural diversity, and (3) species diversity would also indirectly impact stand productivity via changes in size structure. We measured diversity using distance-independent indices. We fitted separate linear mixed-effects models for fir, spruce and beech at the tree level, whereas at the stand level we pooled all available data. We tested our third hypothesis using structural equation modeling. Structural and species diversity acted as direct and independent drivers of stand productivity, with structural diversity being a slightly better predictor. Structural diversity, but not species diversity, had a significant, albeit asymmetric, effect on tree productivity. The functioning of structurally diverse, mixed forests is influenced by both structural and species diversity. These sources of trait diversity contribute to increased vertical stratification and crown plasticity, which in turn diminish competitive interferences and lead to more densely packed canopies per unit area. Our research highlights the positive effects of species diversity and structural diversity on forest productivity and ecosystem dynamics.

There is increasing evidence that mixed-species forests can provide multiple ecosystem services at a higher level than their monospecific counterparts. However, most studies concerning tree diversity and ecosystem functioning relationships use data from forest inventories (under noncontrolled conditions) or from very young plantation experiments. Here, we investigated temporal dynamics of diversityproductivity relationships and diversity-stability relationships in the oldest tropical tree diversity experiment. Sardinilla was established in Panama in 2001, with 22 plots that form a gradient in native tree species richness of one-, two-, three-and five-species communities. Using annual data describing tree diameters and heights, we calculated basal area increment as the proxy of tree productivity. We combined tree neighbourhood-and community-level analyses and tested the effects of both species diversity and structural diversity on productivity and its temporal stability.General patterns were consistent across both scales indicating that tree-tree interactions in neighbourhoods drive observed diversity effects. From 2006 to 2016, mean overyielding (higher productivity in mixtures than in monocultures) was 25%-30% in two-and three-species mixtures and 50% in five-species stands. Tree neighbourhood diversity enhanced community productivity but the effect of species diversity was stronger and increased over time, whereas the effect of structural diversity declined. Temporal stability of community productivity increased with species diversity via two principle mechanisms: asynchronous responses of species to environmental variability and overyielding. Overyielding in mixtures was highest during a strong El Niño-related drought. Overall, positive diversity-productivity and diversity-stability relationships predominated, with the highest productivity and stability at the highest levels of diversity. These results provide new insights into mixing effects in diverse, tropical plantations and highlight the importance of analyses of temporal dynamics for our understanding of the complex relationships between diversity, productivity and stability. Under climate change, mixed-species forests may provide both high levels and high stability of production.

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