Within-stand variability of leaf phenology in deciduous tree species: characterization and ecological implications
Abstract:The vast majority of phenological studies address questions relative to the spatial or temporal variability of phenological timings integrated at the forest stand (i.e. tree population) scale. Within a forest stand, the inter-individual variability of phenological timings is expected to affect a range of tree functions among which the access to light, the use of carbon and nitrogen reserves, the absorption of minerals and the sensitivity to pathogens. Hence the individual’s phenological traits are likely to be strongly selected, resulting in an adaptation of the population to local conditions, as evidenced by latitudinal and altitudinal clines observed in common garden experiments. Studies dedicated to the within-stand variability of the timing of phenophases have to date been mostly designed for contrasting the behaviours of understory versus overstory species or seedlings compared to their adult conspecifics. The few published papers studying the phenological timings among adult conspecifics revealed unclear patterns. We aimed at clarifying the understanding of the within-stand variability of tree phenology of three of the main European deciduous species (Quercus petraea, Quercus robur and Fagus sylvatica) through the analysis of a unique phenological database collected over 44 (28 Oak sites, 16 Beech stands) forest stands at the tree level for 4 years over France.
We show that within a forest stand, individual trees have a distinct “phenological identity” resulting in a year to year conservation of (a) the individuals’ spring and autumn phenological rankings and (b) the individuals’ critical temperature sums required for budburst and senescence. The individual’s spring “phenological identity” affects its functioning and, ultimately, its competitive ability: big trees burst earlier. Acknowledging that Angiosperms show low genetic diversity between populations, we show that the between-site variability of critical temperature sums needed for budburst or senescence is low. We show that this between-site variability is linked to climate and geographical gradients, which suggest local adaptations of the populations’ phenological traits.