Leaf area accumulation helps juvenile evergreen trees tolerate shade in a temperate rainforest
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Most knowledge of the physiological correlates of interspecific variation in shade tolerance derives from studies of first-year seedlings in artificial environments. The present study relates growth, allocation, foliage turnover, biomass distribution and gas exchange traits to low-light survival of large seedlings (20–100 cm tall) of eight temperate rainforest evergreens under field conditions. Taxa for which natural mortality was not observed in low light during the 14-month study are referred to here as "shade-tolerant" species, and those which did die in the shade are referred to as "light-demanding" species. In low light (2–5% canopy openness), shade-tolerant species had slightly lower light compensation points than light-demanders. Light-demanding species had more plastic aboveground allocation patterns, generally allocating proportionally less aboveground biomass to foliage production than shade-tolerant associates in high light (>10% canopy openness), but more in low light. Foliage turnover was generally much slower in shade-tolerant species (10–40% year–1) than in light-demanding species (30–190%). As these differences in leaf retention outweighed variation in allocation, shade-tolerant species displayed higher leaf areas at all light levels. Furthermore, all shade-tolerant species gained leaf area in low light during the study period, whereas light-demanding taxa showed leaf area declines. Higher leaf area ratios, plus differences in light compensation points, indicate that large seedlings of shade-tolerant evergreens enjoy net carbon gain advantages over light-demanding associates in low light. However, minimal growth rate differences in low light imply higher storage allocation in shade-tolerant species. This study provides a rather different picture from that which has emerged from recent reviews of first-year seedling data, illustrating the long-term consequences of foliage turnover differences for biomass distribution, and suggesting that shade tolerance in juvenile evergreen trees is associated with a suite of traits which enhance net carbon gain, but not growth, in low light. Accumulation of a large foliage area through long leaf retention times is probably a key mechanism enhancing low-light carbon gain in evergreens.
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