Scale-dependent effects of pines on the herbaceous layer diversity in a semi-arid mediterranean ecosystem

In arid and semi-arid ecosystems, the presence of woody neighbours affects the existence of several herbaceous species by modifying critical aspects of the environment (e.g., soil humidity, nutrient content or light availability) beneath their canopies. Herbaceous species growing in the understory of Pinus pinaster may be distinct from those in open areas due to litter fall, light interception and changes in nutrient availability. We suggest that the overall effect of woody neighbours on herbaceous layer diversity may vary with the scale focus of analysis. To examine this hypothesis, we collected data on the abundance of herbaceous species in open pineland forests of the central Iberian Peninsula (Spain) using sample quadrats of 0.5 m x 0.5 m distributed beneath, at the edge, and outside the canopy of pines in a landscape composed of dunes and plains. The results of CCA ordination revealed significant spatial segregation of herbaceous species reflecting the occurrence of pines and dunes in the landscape. Nested ANOVA disclosed markedly lower species richness beneath the pines, particularly in the dune sites. Species richness partitioning showed higher pine-induced heterogeneity than expected from the sample-based randomized model, leading to significantly increased species richness at the patch level. Hence, the outcome of pine-induced effects on the herbaceous plant diversity is scale-dependent, negative if we focus on separate communities, but positive if the scale focus is extended to whole patches comprising the sum of communities beneath, at the edge, and outside pine tree canopies. These results emphasize the necessity of using various scale perspectives to clarify the different ways in which pines and other woody nurse species affect structure of herbaceous communities in semi-arid Mediterranean ecosystems.