Sapling biomass allocation and growth in the understory of a deciduous hardwood forest.

Above- and belowground tissues of co-occurring saplings (0.1-1 m height) of Acer saccharum Marsh. (very shade tolerant), Acer rubrum L. (shade tolerant), Fraxinus americana L. (intermediate shade tolerant), and Prunus serotina Ehrh. (shade intolerant) were harvested from a forest understory to test the hypothesis that the pattern of biomass allocation varied predictably with shade-tolerance rank. The placement and length of branches along the main axis were consistent with the formation of a monolayer of foliage for the tolerant and intermediate species. Other morphological characteristics did not vary predictably with shade-tolerance rank. The maintenance of high specific leaf area (SLA; leaf area/leaf mass) and leaf area ratio (LAR; leaf area/sapling mass) is considered important for growth under extreme shade, yet these traits were not clearly related to the shade-tolerance rank of these species. Fraxinus americana, an intermediate species, had the highest LAR and growth rate in the understory, and with the exception of P. serotina, the very shade-tolerant A. saccharum had the lowest LAR. Prunus serotina maintained a large starch-rich tap root and shoot dieback was common, yielding the largest root/shoot ratio for these species. The observed allocation patterns were not similar to the long-standing expectation for the phenotypic response of juvenile trees to shade, but were consistent with three hypothetical "growth strategies" in the understory: (1) the low SLA and LAR of A. saccharum may provide a measure of defense against herbivores and pathogens and thus promote persistence in the understory, (2) the high SLA for F. americana and high LAR for F. americana and A. rubrum may enable these species to achieve high growth rates in shade, and (3) the large carbohydrate stores of P. serotina may poise this species for opportunistic growth following disturbance. The relative importance of resistance to herbivores and pathogens vs. the maintenance of high growth rates may be important in evaluating the patterns of biomass allocation in the understory.