Leaf dynamics and shoot phenology of eleven warm-temperate evergreen broad-leaved trees near their northern limit in central Japan

Dynamic features of shoot phenology including leaf emergence and leaf fall, and leaf life span for eleven evergreen broad-leaved tree species were investigated in a warm-temperate rain forest in Mount Kiyosumi, central Japan. All species had periodic leaf emergence or flushing pattern, and were classified into two types; single and multiple flush and only one species, Eurya japonica, represented the latter type and the rest had single flush in spring. The single Bush type can further be subdivided into two groups according to their duration of shoot growth; short and long flush. Seasonal patterns of leaf fall were categorized into four; unimodal, bimodal, broad unimodal, and multimodal type though they were not fixed pattern. The leaf emergence and leaf fall patterns were correlated for the eleven species, and five phenological types were categorized. Four of them were the single flush types, i.e., short flush of leaf emergence with unimodal leaffall (SSU) type of Castanopsis sieboldii and Quercus salicina, short flush with bimodal leaf fall (SSB) type of Quercus acuta, Machilus thunbergii, Neolitsea sericea, and Cinnamomum japonicum, long flush with bimodal leaffall (SLB) type of Myrsine seguinii, and long flush with broad unimodal leaffall (SLR) type of Symplocos prunifolia, Cleyera japonica, and Illicium anisatum. The multiple flush type is only one species, Eurya japonica, and it had multimodal leaffall pattern (MM type). The phenological pattern varied in relation to leaf life span, leaf size, and tree habit. Leaf life span ranged from 1.1 to 5.8 yr. The short flush species or SSU and SSB types were all canopy or subcanopy trees, and the former had short and the latter had long leaf life spans. The long flush species were all microphyllous small trees, and SLB type had a relatively long leaf life span in understory, SLR type had a long leaf life span in understory or in open habitat and/or forest gap as a pioneer tree. MM type had a long leaf life span and colonizing species in open habitat but they can survive in understory as well. The phenological attributes of evergreen trees were well corresponded to the ecological guild of the tree in both forest structure and successional stage, and were also constrained by phylogenetic groups.