Grow slowly, persist, dominate-Explaining beech dominance in a primeval forest

Being able to persist in deep shade is an important characteristic of juvenile trees, often leading to a strong dominance of shade-tolerant species in forests with low canopy turnover and a low disturbance rate. While leaf, growth, and storage traits are known to be key components of shade tolerance, their interplay during regeneration development and their influence on juveniles' survival time remains unclear. We assessed the ontogenetic effects of these three traits on the survival time of beech (Fagus sylvatica), and Norway and sycamore maples (Acer pseudoplatanus, Acer platanoides) in a primeval beech forest. Biomass allocation, age, and content of nonstructural carbohydrates (NSC) were measured in the stems and roots of 289 seedlings and saplings in high- and low-vitality classes. Saplings experienced a trade-off between absolute growth rate (AGR) and storage (NSC) as the leaf area ratio (LAR) decreases with biomass development. High LAR but low AGR and low NSC corresponded to beech with a marked ability to persist in deep shade while awaiting canopy release. In turn, a comparably small LAR in combination with a high AGR and higher storage (NSC), as observed in Norway maple and sycamore maple, reduced sapling survival time, thus offering an explanation for beech dominance and maple disappearance in the undergrowth of old-growth beech forests.