The space occupation and use by tree crowns explain variations of individual growth rates in an old-growth temperate forest in Japan

Relative growth rates (RGR), i.e., growth rates per unit biomass (ΔM/M, where M is plant mass and ΔM is the change in M over a period of time), reflect growth strategies of plant species. Partitioning of RGR to net assimilation rate (ΔM/leaf area) and leaf area ratio (leaf area/M) provides further insights into allocation strategies. To apply this analytical approach to large canopy tree species, we used crown area (A_c) as a proxy for leaf area to understand variations of RGR partitioned to space use efficiency (SUE, ΔM/A_c) and space occupation efficiency (SOE, A_c/M). With UAV imagery, we measured A_c of 226 co-occurring individuals of 14 canopy tree species in a 1-ha stand in a temperate old-growth mixed-forest in Japan, and analyzed how RGR was related to SUE and SOE. The results show that deciduous species exhibited higher SOE and lower SUE compared to evergreen species, even though their RGR values largely overlapped. Late successional species tended to have higher RGR through higher SUE than early-to-middle successional species. We also analyzed the relationship of absolute growth rates (AGR) with several functional traits including DBH (diameter at breast height), A_c, leaf- and stem traits. Both A_c and DBH were strong determinants of AGR across species. Low specific leaf area (SLA, leaf area per unit leaf mass) and high wood density positively contributed to AGR across species, offering long-term growth advantages for old-growth natural forest. The analytical framework introduced in this study may be useful to elucidate the variation of tree growth strategies of canopy trees in natural forest stands.