The ecology of very small-seeded shade-tolerant trees and shrubs in lowland rain forest in Singapore

A study was made in Bukit Timah Nature Reserve of niche differentiation among 11 woody species that have very small seeds (22–460 µg) and establish in deep shade, i.e., where the indirect site factor (isf) under cloudy conditions is 2%, and one taxonomically related light-demander (seed mass 33 µg). Comparative observations were made on changing light requirement with age. The species varied from shrubs and treelets (Urophyllum) via small to medium-height trees (Ficus, Pternandra) to tall trees (Gynotroches and Pellacalyx). In general, the shorter species fruited in deep shade while the taller needed direct irradiance. All produced ripe fruit at least once a year.In 1–3% isf seedling mortality over 11 months was 24% for Urophyllum hirsutum and 57% for Pternandra echinata (two strongly shade-tolerant species); survivors produced appreciable new stem and leaf tissue. For two more light-demanding species (Ficus grossularioides and F. fistulosa) mean lamina area was much larger in small gaps (10% isf) than in the understorey (<3% isf), while for Urophyllum hirsutum it was much smaller. These three species showed the greatest difference in SLA. There were few significant intraspecific differences between gaps and understorey in concentrations of N, P or K. In a valley where there was no significant difference in isf between a path and adjacent understorey, seedlings of species with seeds <1 mg mean dry mass were five times as dense on transects along steep pathside banks as in the understorey 5 m upslope. The mean mid-bank slope was 66° (cf. 20° upslope). On mid-bank transects litter had a cover of 36% (cf. 95% upslope), and was thinner (0.8 vs 3.2 cm). The soil at 0–7 cm depth into the mid-bank had much lower concentrations of organic matter, total N and P (but not total K) and fine roots than that of upslope transects. All species were found to have vesicular-arbuscular mycorrhiza when 1–2 m tall in deep shade, several having very dense infections. Seedling form and responses are discussed in relation to potential adaptive value and possible evolutionary origin.