Scaling relationships among twig size, leaf size and leafing intensity in a successional series of subtropical forests

Scaling relationships among twig size, leaf size and leafing intensity fundamentally influence the twig-leaf deployment pattern, a property that affects the architecture and functioning of plants. However, our understanding of how these relationships change within a species or between species as a function of forest succession is unclear. We determined log-log scaling relationships between twig cross-sectional area (twig size) and each of total and individual leaf area, and leafing intensity (the number of leaves per twig volume) for 78 woody species along a successional series in subtropical evergreen forests in eastern China. The series included four stages: secondary shrub (S1), young (S2), sub-climax (S3) and climax evergreen broadleaved forests (S4). The scaling slopes in each of the three relationships did not differ among the four stages. The y-intercept did not shift among the successional stages in the relationship between twig cross-sectional area and total leaf area; however, the y-intercept was greatest in S4, intermediate in S3 and lowest in S2 and S1 for the relationship between twig size and individual leaf area, while the opposite pattern was found for the twig size-leafing intensity relationship. This indicates that late successional trees have few but large leaves while early successional trees have more small leaves per unit twig size. For the relationship between twig cross-sectional area and total leaf area, there was no difference in the regression slope between recurrent (appear in more than one stages) and non-recurrent species (appear in only one stage) for each of the S1-S2, S2-S3 and S3-S4 pairs. A significant difference in the y-intercept was found in the S2-S3 pair only. In the relationship between twig cross-sectional area and individual leaf area, the regression slope between recurrent and non-recurrent species was homogeneous in the S1-S2 and S3-S4 pairs, but heterogeneous in the S2-S3 pair. We conclude that forest succession caused the shift in the intercept, but did not affect scaling slopes for relationships among twig size, leaf size and leaf intensity. For recurrent species, the invariant scaling slope in the twig-leaf size relationship between adjacent pairs of successional stages may be related to their phenotypic plasticity by adjusting their twig and leaf deployment strategy to similar to what the non-recurrent species display.