Background and aims: The "diminishing returns" hypothesis postulates that the scaling exponent governing the lamina area versus lamina mass scaling relationships has, on average, a numerical value less than one. Theoretically, a similar scaling relationship may exist at the twig level. However, this possibility has not been explored empirically. Methods: We tested both hypotheses by measuring the lamina area and mass, petiole mass of individual leaves, and the total foliage area and stem mass of individual current-year shoots (twigs) of 64 woody species growing in three characteristic forest community types: (1) Evergreen broad-leaved, (2) mixed coniferous and broad-leaved, and (3) deciduous. Key results: The results demonstrate that lamina area vs. mass and lamina area vs. petiole mass differ significantly among the three forest types at both the individual leaf and twig levels. Nevertheless, the scaling exponents of lamina area vs. mass were <1.0 in each of the three community types, as were the corresponding exponents for lamina area vs. petiole mass, both within and across the three community types. Similar trends were observed at the individual twig level. The numerical values of the scaling exponent for lamina area vs. petiole mass and total foliage area vs. stem mass per twig decreased with increased elevation. Conclusions: These data support the "diminishing returns" hypothesis at both the individual leaf level and at the individual twig level, phenomena that can inform future inquiries into the mechanistic basis of biomass allocation patterns to physiological (leaf) and mechanical (stem) plant organs.
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