Effects of canopy cover and seasonal reduction in rainfall on leaf phenology and leaf traits of the fern Oleandra pistillaris in a tropical montane forest, Indonesia

Abstract

Leaf phenology and leaf traits of the fern Oleandra pistillaris were examined in relation to canopy cover (open and understorey) and seasonal reduction in rainfall in a wet tropical montane forest, Indonesia. Although the annual rainfall is high, rainfall is relatively less in June and July. Stomatal density and diameter were greater in the open than in the understorey (229 versus 167 mm⁻² for stomatal density and 33 versus 29 μm for stomatal diameter). The stable carbon isotope ratio (δ¹³C) of leaves, positively correlated with water use efficiency, was higher in the open than in the understorey (mean δ¹³C −30 versus −33‰). Therefore, it is considered that leaves have high gas-exchange capacities per leaf area in the open where water availability would be limited, compared with the understorey. In contrast, leaf mass per area (LMA) was lower and leaf life span was longer in the understorey than in the open (25 versus 34 g m⁻² for LMA and 2.1 and 1.6 y for leaf life span). These thin leaves with a long life span in the understorey would contribute to efficient light capture and photosynthetic production per leaf mass. The number of leaves per stem decreased during the period with less rainfall in both the open and understorey conditions, which should reduce the water loss from plants, but increased again after the period with less rainfall. Stem growth rate was higher in the open than in the understorey, and the seasonal reduction in rainfall hardly affected stem growth rate in either open or understorey conditions. This study concludes that O. pistillaris responds to canopy cover and seasonal reduction in rainfall by adjusting leaf traits and leaf phenology, respectively.