Partitioning the effects of biodiversity and environmental heterogeneity for productivity and mortality in a tropical tree plantation

1. Over 5000 trees were grown in plots of differing diversity levels (1, 3 and 6 species) in a plantation established in Panama. Four and five years after establishment, we analysed parameters related to the productivity of this tropical plantation (tree survival, height and biomass as well as plot basal area) to test for the presence of biodiversity effects. The relative importance of environmental heterogeneity (such as soil, topography, and drainage) and biodiversity on tree growth and mortality was determined using partial redundancy analysis. 2. Hierarchical clustering revealed nine different soil clusters based on soil quality and drainage. By chance, the six-species plots were apparently established on more variable soils then on the other diversity levels. We found little evidence for spatial autocorrelation between subplots, with the exception of four subplots located on a ridge that extends on the North-South axis of the plantation and corresponds to a zone of higher productivity. 3. The redundancy analysis indicated that environmental heterogeneity and biodiversity together explained around 50% of the variation in subplot productivity and tree mortality. Environment explained 35-57% of the variation in productivity and mortality, respectively, whereas diversity explained an additional 23-30%. 4. Our simulation model revealed a significant positive effect of biodiversity on growth but no effect of biodiversity on mortality. The standardized effect sizes that we used to detect over- or under-yielding or no effect in comparison with monoculture were highly variable and the variability was largely explained by traits related to site topography. 5. Synthesis. In our tropical tree plantation, we detected biodiversity effects at a scale relevant to conservation and quantified the relative importance of environmental heterogeneity and diversity on tree growth and mortality. Our results support the idea that environmental factors could act as hidden sources of variability in biodiversity experiments. Environmental and spatial heterogeneity induced variable responses to biodiversity and amplified the differences between three- and six-species plots. Species identity explained more variation in productivity than did the species diversity. One species, Cedrela odorata, was associated with increased productivity.