Shade adaptation and shade tolerance in saplings of three Acer species from eastern North America.

Saplings of three, co-occurring maple species in a mature maple-beech forest differed in a suite of structural and physiological characters that separated the canopy species, Acer, saccharum, from the two subcanopy species, A. pensylvanicum and A. spicatum. Acer saccharum had both more dense wood and tougher and heavier but thinner leaves than the subcanopy species. Acer pensylvanicum had the largest, lightest leaves with high stomatal density and its canopy architecture was the most effective in terms of leaf display for light interception. Acer spicatum had weaker wood similar to that of A. pensylvanicum but also small, soft and relatively poorly displayed leaves. Both subcanopy species maintained marginally higher average rates of photosynthesis over the growing season in the understory environment. We consider juvenile A. saccharum only shade-tolerant, capable of persisting through long periods in the closed canopy until a gap occurs but not specifically adapted to the understory environment. Juvenile A. sacchrum appears to be constrained functionally by the requirements set by the canopy environment that adults will occupy. Characters such as high wood density are already expressed in the understory sapling; this investment in denser wood slows the growth of saplings, but is necessary for structural reasons in the adult. Juvenile A. saccaharum have morphological and photosynthetic characters better suited to gas exchange and extension growth under the increased photon flux densities in large forest gaps, characteristics that will also be advantageous in the sunlit canopy environment of adults.Both subcanopy maples appear to be more truly shade-adapted, although in somewhat different ways. Acer pensylvanicum has characteristics that enhance the potential for capture and utilization of sunflecks and is able to sustain higher growth rates than A. saccharum in the shaded subcanopy environment. Acer spicatum shares some shade-adapted features with A. pensylvanicum, and its habit of lateral spread through stem layering may confer an additional advantage in foraging for small light gaps.