Within-twig leaf distribution patterns differ among plant life-forms in a subtropical Chinese forest

Abstract

In theory, plants can alter the distribution of leaves along the lengths of their twigs (i.e., within-twig leaf distribution patterns) to optimize light interception in the context of the architectures of their leaves, branches and canopies. We hypothesized that (i) among canopy tree species sharing similar light environments, deciduous trees will have more evenly spaced within-twig leaf distribution patterns compared with evergreen trees (because deciduous species tend to higher metabolic demands than evergreen species and hence require more light), and that (ii) shade-adapted evergreen species will have more evenly spaced patterns compared with sun-adapted evergreen ones (because shade-adapted species are generally light-limited). We tested these hypotheses by measuring morphological traits (i.e., internode length, leaf area, lamina mass per area, LMA; and leaf and twig inclination angles to the horizontal) and physiological traits (i.e., light-saturated net photosynthetic rates, Amax; light saturation points, LSP; and light compensation points, LCP), and calculated the 'evenness' of within-twig leaf distribution patterns as the coefficient of variation (CV; the higher the CV, the less evenly spaced leaves) of within-twig internode length for 9 deciduous canopy tree species, 15 evergreen canopy tree species, 8 shade-adapted evergreen shrub species and 12 sun-adapted evergreen shrub species in a subtropical broad-leaved rainforest in eastern China. Coefficient of variation was positively correlated with large LMA and large leaf and twig inclination angles, which collectively specify a typical trait combination adaptive to low light interception, as indicated by both ordinary regression and phylogenetic generalized least squares analyses. These relationships were also valid within the evergreen tree species group (which had the largest sample size). Consistent with our hypothesis, in the canopy layer, deciduous species (which were characterized by high LCP, LSP and Amax) had more even leaf distribution patterns than evergreen species (which had low LCP, LSP and Amax); shade-adapted evergreen species had more even leaf distribution patterns than sun-adapted evergreen species. We propose that the leaf distribution pattern (i.e., 'evenness' CV, which is an easily measured functional trait) can be used to distinguish among life-forms in communities similar to the one examined in this study.