Survival time and mortality rate of regeneration in the deep shade of a primeval beech forest

Low mortality rates and slow growth differentiate shade-tolerant from shade-intolerant species and define the survival strategy of juvenile trees growing in deep shade. While radial stem growth has been widely used to explain mortality in juvenile trees, the leaf area ratio (LAR), known to be a key component of shade tolerance, has been neglected so far. We assessed the effects of LAR, radial stem growth and tree height on survival time and the age-specific mortality rate of juvenile Fagus sylvatica L. (European beech), Acer pseudoplatanus L. (sycamore maple) and Acer platanoides L. (Norway maple) in a primeval beech forest (Ukraine). Aboveground and belowground biomass and radial stem growth were analysed for 289 living and 179 dead seedlings and saplings. Compared with the other species, F. sylvatica featured higher LAR, slower growth and a lower mortality rate. The average survival time of F. sylvatica juveniles (72 years) allows it to reach the canopy more often than its competitors in forests with low canopy turnover rate. In contrast, a combination of lower LAR, higher growth rate and higher age-specific mortality rate of the two Acer species resulted in their shorter survival times and thus render their presence in the canopy a rare event. Overall, this study suggests that shade tolerance, commonly defined as a relationship between sapling mortality and growth, can alternatively be formulated as a relationship between survival time and the interplay of growth and LAR.