A Second Dimension to the Leaf Economics Spectrum Predicts Edaphic Habitat Association in a Tropical Forest

Background: Strong patterns of habitat association are frequent among tropical forest trees and contribute to the maintenance of biodiversity. The relation of edaphic differentiation to tradeoffs among leaf functional traits is less clear, but may provide insights into mechanisms of habitat partitioning in these species rich assemblages. Methodology/Principal Findings: We quantify the leaf economics spectrum (LES) for 16 tree species within a Bornean forest characterized by highly pronounced habitat specialization. Our findings suggest that the primary axis of trait variation in light-limited, lowland tropical forests was identical to the LES and corresponds with the shade tolerance continuum. There was no separation with respect to edaphic variation along this primary axis of trait variation. However, a second orthogonal axis determined largely by foliar P concentrations resulted in a near-perfect separation of species occupying distinct soil types within the forest. Conclusions/Significance: We suggest that this second axis of leaf trait variation represents a leaf edaphic habitat spectrum'' related to foliar P and potentially other nutrients closely linked to geological substrate, and may generally occur in plant communities characterized by strong edaphic resource gradients.