Plant Phylogeny and Growth Form as Drivers of the Altitudinal Variation in Woody Leaf Vein Traits

Variation in leaf veins along environmental gradients reflects an important adaptive strategy of plants to the external habitats, because of their crucial roles in maintaining leaf water status and photosynthetic capacity. However, most studies concentrate on a few species and their vein variation across horizontal spatial scale, we know little about how vein traits shift along the vertical scale, e.g., elevational gradient along a mountain, and how such patterns are shaped by plant types and environmental factors. Here, we aimed to investigate the variation in leaf vein traits (i.e., vein density, VD; vein thickness, VT; and vein volume per unit leaf area, VV) of 93 woody species distributed along an elevational gradient (1,374-3,375 m) in a temperate mountain in China. Our results showed that altitude-related trends differed between growth forms. Tree plants from higher altitudes had lower VD but higher VT and VV than those from lower altitude; however, the opposite tend was observed in VD of shrubs, and no significant altitudinal changes in their VT or VV. Plant phylogenetic information at the clade level rather than climate explained most of variation in three leaf vein traits (17.1-86.6% vs. <0.011-6.3% explained variance), supporting the phylogenetic conservatism hypothesis for leaf vein traits. Moreover, the phylogenetic effects on vein traits differed between trees and shrubs, with the vein traits of trees being relatively more conserved. Together, our study provides new picture of leaf vein variation along the altitude, and highlights the importance of taking plant phylogeny into consideration when discussing trait variation from an ecological to a biogeographic scale.