Do we Underestimate the Importance of Leaf Size in Plant Economics? Disproportional Scaling of Support Costs Within the Spectrum of Leaf Physiognomy

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Abstract
Broad scaling relationships between leaf size and function do not take into account that leaves of different size may contain different fractions of support in petiole and mid-rib. The fractions of leaf biomass in petiole, mid-rib and lamina, and the differences in chemistry and structure among mid-ribs, petioles and laminas were investigated in 122 species of contrasting leaf size, life form and climatic distribution to determine the extent to which differences in support modify whole-lamina and whole-leaf structural and chemical characteristics, and the extent to which size-dependent support investments are affected by plant life form and site climate. For the entire data set, leaf fresh mass varied over five orders of magnitude. The percentage of dry mass in mid-rib increased strongly with lamina size, reaching more than 40 % in the largest laminas. The whole-leaf percentage of mid-rib and petiole increased with leaf size, and the overall support investment was more than 60 % in the largest leaves. Fractional support investments were generally larger in herbaceous than in woody species and tended to be lower in Mediterranean than in cool temperate and tropical plants. Mid-ribs and petioles had lower N and C percentages, and lower dry to fresh mass ratio, but greater density (mass per unit volume) than laminas. N percentage of lamina without mid-rib was up to 40 % higher in the largest leaves than the total-lamina (lamina and mid-rib) N percentage, and up to 60 % higher than whole-leaf N percentage, while lamina density calculated without mid-rib was up to 80 % less than that with the mid-rib. For all leaf compartments, N percentage was negatively associated with density and dry to fresh mass ratio, while C percentage was positively linked to these characteristics, reflecting the overall inverse scaling between structural and physiological characteristics. However, the correlations between N and C percentages and structural characteristics differed among mid-ribs, petioles and laminas, implying that the mass-weighted average leaf N and C percentage, density, and dry to fresh mass ratio can have different functional values depending on the importance of within-leaf support investments. These data demonstrate that variation in leaf size is associated with major changes in within-leaf support investments and in large modifications in integrated leaf chemical and structural characteristics. These size-dependent alterations can importantly affect general leaf structure vs. function scaling relationships. These data further demonstrate important life-form effects on and climatic differentiation in foliage support costs.