Changes in Leaf Functional Traits with Leaf Age for Coexisting Woody Species in Temperature Forest of Northern China

Leaf-trait variation has traditionally been focused on both within and among species along environmental gradients, while leaf age has received less attention. By measuring leaf morphological, stomatal, and stoichiometric traits of 40 coexisting woody species in temperate forest in northern China, we analyzed their variation pattern and the correlations among different plant life forms and leaf age. We found that leaf age has significant effects on leaf functional traits. The young leaves of both shrub and tree species revealed a lower stoma density (SD) and a higher stoma length (SL), stoma width (SW), and leaf N content (LNC) than mature leaves. Shrub species have a higher SLA and SD than tree species for both young and mature leaves. The traits of young leaves generally revealed a higher variation than those of mature leaves. Although correlations between traits are similar between young leaves and mature leaves, the slopes of the SLA-SD and SD-LNC relationships were significantly affected by leaf age. These findings elucidate the adaptive changes of leaf traits during leaf maturation and underscore the trade-off between stomatal safety and efficiency, as well as the trade-off between leaf hydraulic and economic traits in temperate woody species during leaf development. We conclude that variation in leaf traits with age may play a potentially important role in understanding the ecological function of woody species in temperate forests.