Abstract

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.

Highlights

  • Leaf veins are composed of xylem and phloem cells that are embedded in parenchyma, or sometimes sclerenchyma, and are surrounded by bundle sheath cells (Sack and Scoffoni, 2013)

  • As an important criterion of plant classification, leaf vein traits should be relatively invariant for a given species, and we propose the leaf vein trait phylogenetic conservatism hypothesis, which means that differences in leaf vein networks can be mainly attributed to intrinsic evolutionary adaptations among major phylogenetic clades, with plants evolving denser and thinner minor leaf veins to counter the reduction in atmospheric CO2 concentrations that occurred during the Cretaceous (Brodribb and Feild, 2010; Feild and Brodribb, 2013; Sack and Scoffoni, 2013)

  • When all species were considered, no clear trends were observed for the vein traits along the altitudinal gradient, and phylogenetic effect explained most of the variation at the clade level, whereas climate exerted very weak influence

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Summary

Introduction

Leaf veins are composed of xylem and phloem cells that are embedded in parenchyma, or sometimes sclerenchyma, and are surrounded by bundle sheath cells (Sack and Scoffoni, 2013). Based on a survey of 485 global species, Sack et al (2012) emphasized that both larger leaves from moist regions and small leaves from dry regions may possess high VD. These studies suggest that trait–environment relationships can be obscured by different underlying processes, such as adaptive strategies and phylogenetic constraints, and, require further study (Blonder et al, 2017; Schneider et al, 2018)

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