Variation in the aboveground stand structure and fine-root biomass of Bornean heath (kerangas) forests in relation to altitude and soil nitrogen availability

To elucidate the biotic and abiotic factors affecting the variation in fine-root biomass (FRB, < 2 mm diameter) of trees growing under nutrient-poor environments in Sabah, North Borneo, we investigated FRB in different forests with varying soil nitrogen (N) availability. We selected two study sites at different altitudes: the Maliau Basin (ca. 1000 m asl) and Nabawan (ca. 500 m asl). Both sites included tropical heath (kerangas) forest, on infertile soils (podzols) with a surface organic horizon overlying a bleached (eluviated) mineral horizon, and taller forests on more fertile non-podzolic soils. FRB was obtained from each plot by soil coring (to a depth of 15 cm). FRB increased with decreasing soil inorganic N content (NH4-N and NO3-N), tree height, and aboveground biomass. Thus, higher proportions of carbon resources were allocated to fine-roots in stands with lower N availability. FRB was significantly greater at the Maliau Basin than at Nabawan, reflecting lower soil N availability at higher altitude. Our results demonstrate high variation in FRB among the heath forests, and suggest that fine-root development is more prominent under a cooler climate where N availability limits tree growth owing to slower decomposition. The variation in N availability under the same climate (i.e., at the same altitude) appears to be related to the extent of soil podzolization.