Interspecific integration of trait dimensions at local scales: the plant phenotype as an integrated network

Plant phenotypic diversity is shaped by the interplay of trade-offs and constraints in evolution. Closely integrated groups of traits (i.e. trait dimensions) are used to classify plant phenotypic diversity into plant strategies, but we do not know the degree of interdependence among trait dimensions. To assess how selection has shaped the phenotypic space, we examine whether trait dimensions are independent. We gathered data on saplings of 24 locally coexisting tree species in a temperate forest, and examined the correlation structure of 20 leaf, branch, stem and root traits. These traits fall into three well-established trait dimensions (the leaf economic spectrum, the wood spectrum and Corner's Rules) that characterize vital plant functions: resource acquisition, sap transport, mechanical support and canopy architecture. Using ordinations, network analyses and Mantel tests, we tested whether the sapling phenotype of these tree species is organized along independent trait dimensions. Across species, the sapling phenotype is not structured into clear trait dimensions. The trait relationships defining trait dimensions are either weak or absent and do not dominate the correlation structure of the sapling phenotype as a whole. Instead traits from the three commonly recognized trait dimensions are organized into an integrated trait network. The effect of phylogeny on trait correlations is minimal. Our results indicate that trait dimensions apparent in broad-based interspecific surveys do not hold up among locally coexisting species. Furthermore, architectural traits appear central to the phenotypic network, suggesting a pivotal role for branching architecture in linking resource acquisition, mechanical support and hydraulic functions. Synthesis. Our study indicates that local and global patterns of phenotypic integration differ and calls into question the use of trait dimensions at local scales. We propose that a network approach to assessing plant function more effectively reflects the multiple trade-offs and constraints shaping the phenotype in locally co-occurring species.