Combined effects of soil 3D spatial heterogeneity and biotic spatial heterogeneity (plant clumping) on ecosystem processes in grasslands

Soil heterogeneity has been shown to enhance plant diversity, but its effect on grassland productivity is less clear. Even less is known about the effect of plant clumping (intraspecific aggregation) and its potential interaction with soil heterogeneity. The combined effects of soil 3D spatial heterogeneity and species clumping were experimentally studied in grassland mesocosms consisting of four grassland species. These species were planted in three patterns (i.e. completely mixed, clumped by 9 or 36 individuals of the same species) on soils with heterogeneous cells of alternating nutrient-poor and rich soil differing in size from 0 (mixed soil) to 12, 24, and 48 cm (complete poor or rich mesocosm). Moderate soil cell sizes (12-24 cm) consistently increased whole-mesocosm aboveground productivity by more than 20%, which mainly originated from the increased growth of the plants growing on the poor soil cells. In contrast, total mesocosm productivity was not affected by species clumping although there were some species-specific effects, both of clumping and of the interaction of clumping with soil heterogeneity. Our results show that intermediate soil heterogeneity promotes productivity. Clumping can improve the growth of inferior species, thus promoting coexistence, without affecting overall productivity. We found no interaction effect of clumping and soil heterogeneity on productivity at the community level and some minor species-specific effects.