Complex range shifts among forest functional types under the contemporary warming.

The direction and magnitude of species distribution shifts tend to differ among species and functional types (FTs). Quantifying functional trait variation and species interactions will improve our understanding of the complex mechanisms that govern ecosystem dynamics and their responses to climate change. Here, we analyzed differences in the juvenile and adult temperature ranges of Japanese tree species at the mean, colder edge, and warmer edge of their distributions to reveal how functional traits affect interactions between different FT groups (e.g., deciduous and evergreen broad-leaved trees), using linear models and permutation tests. Overall, juveniles preferred cooler sites, but with high variation. The variation among species was partly explained by the difference in seed mass where species with lighter seeds tend to colonize colder sites. On the other hand, the distribution range of FTs showed complex behavior at the ecotones of different FTs. Specifically, in three of eight ecotones, nonparallel range shifts between FTs were detected, which includes cold shifting in deciduous broad-leaved FT where a warm shift by subalpine FT happened, and cold shifting in subtropical FT where warm shifts by either the deciduous broad-leaved or the evergreen broad-leaved FTs happened. Our results suggest that past warming has caused a general cold shift at species level, whereas different mechanisms, such as light seeds disperse farther in distribution's colder edge and heavy seeds (e.g., evergreen broad-leaved) compete better in warmer edge, create nonparallel responses of FT distribution ranges leading to the observed homogenization at several ecotones among FTs. These complex range shifts at FT level have crucial implications for climate change mitigation and adaptation.