Population maintenance of the short-lived shrub Sambucus in a deciduous forest

This study quantitatively clarifies the life history of a shrub, Sambucus racemosa ssp. sieboldiana, in an old-growth forest, the Ogawa Forest Reserve, Japan, by a demographic approach using a projection matrix model that incorporates interactions between demographic parameters and canopy height dynamics. S. racemosa is a common deciduous shrub in central Japan and is known to grow predominantly at forest edges or roadsides. This indicates that it is a highly light-demanding species, and occurrence in gaps in old-growth stands suggests its "fugitive," gap-dependent life history in old-growth forests. We found that one distinctive feature of this species was that its seedlings can survive well in shaded conditions by alternating stems every year like perennial herb species. Matrix model analyses demonstrated that S. racemosa can continuously regenerate under the present disturbance regime of this forest and is highly adaptable to the structural dynamics of the old-growth forest. The maturity of S. racemosa shrubs depends on their size, and nearly all (>90%) of the mature (reproducing) individuals were found in gaps or near gaps. But wide seed dispersal by birds and the ability to form both seed banks and seedling banks, the latter of which has been regarded as a common characteristic of shade-tolerant climax species, probably increase the species' chances to encounter canopy gaps. Dynamic-canopied matrix models showed that the greatest elasticity is with shaded seedling survival. The frequent stem alternation of shaded seedlings often makes the growth rate negative, but the survival rate of seedlings in low light awaiting new gap creation is remarkably high (0.93 yr(-1)). The lower survival rate of the larger individuals and smaller minimum size to start reproduction than other canopy or subcanopy shade-tolerant species indicate that S. racemosa has the potential to reproduce before the closure of the encountered gaps and to complete its life history rapidly.