Convergence and correlations among leaf size and function in seed plants: A comparative test using independent contrasts

Prior studies of a broad array of seed plants have reported strong correlations among leaf life span, specific leaf area, nitrogen concentration, and carbon assimilation rates, which have been interpreted as evidence of coordinated leaf physiological strategies. However, it is not known whether these relationships reflect patterns of evolutionary convergence, or whether they are due to contrasting characteristics of major seed plant lineages. We reevaluated a published data set for these seven traits measured in over 100 species, using phylogenetic independent contrasts calculated over a range of alternative seed plant phylogenies derived from recent molecular systematic analyses. In general, pairwise correlations among these seven traits were similar with and without consideration of phylogeny, and results were robust over a range of alternative phylogenies. We also evaluated relationships between these seven traits and lamina area, another important aspect of leaf function, and found moderate correlations with specific leaf area (0.64), mass-based photosynthesis (0.54), area-based nitrogen (-0.56), and leaf life span (-0.42). However, several of these correlations were markedly reduced using independent contrasts; for example, the correlation between leaf life span and lamina area was reduced to close to zero. This change reflects the large differences in both these traits between conifers and angiosperms and the absence of a relationship between the traits within these groups. This analysis illustrates that most interspecific relationships among leaf functional traits, considered across a broad range of seed plant taxa, reflect significant patterns of correlated evolutionary change, lending further support to the adaptive interpretation of these relationships.