ONTOGENETIC CHANGES IN THE NUMBERS OF SHORT- VS. LONG-SHOOTS ACCOUNT FOR DECREASING SPECIFIC LEAF AREA IN ACER RUBRUM (ACERACEAE) AS TREES INCREASE IN SIZE

Specific leaf area (SLA) is reported to decrease with increasing plant size among dicot tree species despite a strong positive correlation between SLA and relative growth rate. This diminishing returns in SLA may result from changes in the relative numbers of different shoot types bearing leaves with different SLAs as trees increase in overall size. This ontogenetic shift hypothesis was examined for 15 Acer rubrum trees differing in basal stem diameter (0.01 m <= D <= 0.62 m). Detailed analyses of the largest tree showed that short-shoots produced leaves with significantly smaller SLA than the leaves produced by long-shoots regardless of the location of shoots within the canopy. A combination of random effect and split-plot (main-effect) ANOVA models showed that >94% of the variance observed for SLA was attributable to shoot type rather than to the location of leaves in the canopy. Further, with increasing trunk diameter, the number of short-shoots increased rapidly relative to the number of long-shoots. Although the leaves of short-shoots gain disproportionately more surface area per unit mass investment compared to the leaves produced by long-shoots, our data show that ontogenetic shifts occurring at the shoot and whole plant level account for size-dependent decreases in total canopy SLA.