Living long by staying small: stem layering as an adaptive life-history trait in shade-tolerant tree seedlings

Suppressed seedling banks are important in replacement dynamics in late-successional forests. However, demographic properties of seedling populations are poorly known, and there has been little attention to traits that might affect fitness in suppressed seedlings. Acer saccharum Marsh., a shade-tolerant dominant in eastern North American forests, frequently develops adventitious roots along prostrate portions of stems (“layering”). Measurements of Acer seedlings in old-growth forests in Michigan indicate that layered seedlings proportionally reduce structural allocations to older layered stem tissues, retain leaf area / height ratios of younger unlayered seedlings, and tend to survive longer. In tree seedlings, allometric consequences of normal stem growth lead to declining ratios of photosynthetic to nonphotosynthetic biomass, which potentially reduces shade tolerance and limiting age. The layering habit may defer this penalty by changing the allometry of growth. Resulting increases in life expectancy should increase chance of access to increased light and of reaching the canopy. Thus, because flowering is generally restricted to canopy trees, the tendency to layer may increase fitness. Properties of individuals in suppressed seedling banks may be selectively and ecologically important, shaping life histories and population dynamics.