“…Our analysis of species occurrence as a function of spatial trait values also suggests that a combination of these traits contributes differently to the delimitation of the species range (Table 3), in particular: (i) mortality delimits certain parts of the southern and eastern range of beech, reflecting the climatic marginality of the species in these areas, and meaning that these populations are most threatened and making eastwards expansion of beech difficult (although more studies on regeneration are needed to confirm this result); this is the case for many species whose highest mortality is in the driest part of their range (Anderegg et al, 2015; Benito-Garzón et al, 2013; Camarero, Gazol, Sancho-Benages, & Sangüesa-Barreda, 2015); (ii) the smallest girths are predicted in the southern part of the distribution and the eastern part of the range, suggesting that radial growth is mostly restricted by drought (interaction graph and map, Figure 1b), as has already been pointed out (Farahat & Linderholm, 2018); (iii) with very little variation across climatic gradients, vertical growth alone will not delimit beech range. This is not the case for other tree species, for which tree height is clearly delimiting species range (Chakraborty, Schueler, Lexer, & Wang, 2018), highlighting the fact that no single best trait delimits tree species ranges; (iv) projections of trees growing in southern and south-eastern regions that flush early also have higher mortality and lower growth predictions than elsewhere within the species range. However, when tree height and leaf flushing are pooled together in the two-trait model, this leads to an decrease in vertical growth in the North; (v) it seems that in beech, and likely in other species with local adaptation to photoperiod, phenology could restrict the northern expansion of ranges (Duputié et al, 2015; Saltré, Duputié, Gaucherel, & Chuine, 2015), although the link between phenology, survival and fitness is still unclear, and more experiments are needed to better understand the interaction between photoperiod and phenology.…”