Woody plant encroachment amplifies spatial heterogeneity of soil phosphorus to considerable depth

The geographically extensive phenomenon of woody plant encroachment into grass-dominated ecosystems has strong potential to influence biogeochemical cycles at ecosystem to global scales. Previous research has focused almost exclusively on quantifying pool sizes and flux rates of soil carbon and nitrogen (N), while few studies have examined the impact of woody encroachment on soil phosphorus (P) cycling. Moreover, little is known regarding the impact of woody encroachment on the depth distribution of soil total P at the landscape scale. We quantified patterns of spatial heterogeneity in soil total P along a soil profile by taking spatially explicit soil cores to a depth of 120 cm across a subtropical savanna landscape that has undergone encroachment by Prosopis glandulosa (an N2 -fixer) and other tree/shrub species during the past century. Soil total P increased significantly following woody encroachment throughout the entire 120-cm soil profile. Large groves (>100 m2 ) and small discrete clusters (<100 m2 ) accumulated 53 and 10 g P/m2 more soil P, respectively, compared to grasslands. This P accumulation in soils beneath woody patches is most likely attributable to P uplift by roots located deep in the soil profile (>120 cm) and transfer to upper portions of the profile via litterfall and root turnover. Woody encroachment also altered patterns of spatial heterogeneity in soil total P in the horizontal plane, with highest values at the centers of woody patches, decreasing toward the edges, and reaching lowest values in the surrounding grassland matrix. These spatial patterns were evident throughout the upper 1.2 m of the soil profile, albeit at reduced magnitude deeper in the soil profile. Spatial generalized least squares models indicated that fine root biomass explained a significant proportion of the variation in soil total P both across the landscape and throughout the profile. Our findings suggest that transfer of P from deeper soil layers enlarges the P pool in upper soil layers where it is more actively cycled may be a potential strategy for encroaching woody species to satisfy their P demands.