In primeval monodominant Fagus sylvatica L. forests, seedlings (0-130 cm tall) and saplings (131-500 cm tall) often grow in deep shade for extended periods, due to low canopy turnover (Hobi et al., 2015; Runkle, 1985;Valverde & Silvertown, 1997). If canopy turnover is low (i.e., the mean time between recurring gap formation at any point in the forest), saplings that are able to persist for decades in shade are more likely to experience a canopy opening (i.e., a release event) enabling the subsequent promotion to canopy (Canham, 1985(Canham, , 1990. Hence, high juvenile shade tolerance is pivotal in determining the survival time of F. sylvatica and may be compromised in co-occurring species, explaining the low tree species diversity and F. sylvatica dominance frequently observed during succession (Korpel, 1995;Rey et al., 2019). Shade tolerance can be assessed via functional traits, that is, morphological, physiological, and phenological features that reflect a species' ecological strategy (Pérez-Harguindeguy