Summary1 Spatial patterns of two codominant dioecious tree species, Podocarpus nagi and Neolitsea aciculata, were analysed at Mt. Mikasa, Nara City, Japan. Podocarpus nagi has a higher shade tolerance but a narrower seed dispersal range than N. aciculata. We examined the eects of dioecy on regeneration and coexistence of the two species. 2 Seeds of P. nagi are dispersed by gravity around female trees. Young plants of P. nagi were clumped and showed signi®cant attraction to large female trees and signi®cant repulsion from large male trees. Dioecy therefore aected the spatial heterogeneity of plant density in the P. nagi population. 3 Seeds of N. aciculata are widely dispersed by birds, and young plants of N. aciculata therefore showed no signi®cant attraction to female trees. This wide dispersal of seeds moderated the eects of dioecy on the spatial pattern in N. aciculata. 4 Large N. aciculata trees were clumped and showed signi®cant attraction to large male P. nagi trees, suggesting that growth of N. aciculata is facilitated where young P. nagi plants are uncommon and competition is therefore less intense. 5 One eect of dioecy may be to produce a population structure for P. nagi that promotes its coexistence with N. aciculata.

The effects of El Niño-induced droughts on dipterocarp forests must be quantified to evaluate the implications of future global climatic changes for the tropical forests of Southeast Asia. We studied the mortality of trees ! 1 cm in diameter in a lowland dipterocarp forest in Borneo before, during, and after the 1997/1998 El Niño drought. The annual mortality rates were 1.30, 1.75, and 1.66 percent/yr for the pre-drought, drought, and post-drought periods, respectively. The effect of drought was tree size-dependent being greater for larger trees. Modified logistic regression analysis revealed a significant interaction effect between species' habitat association and edaphic condition on mortality rates in all periods. For species associated with wet habitat, drought effect was greater in dry conditions than in wet conditions, in both the drought and post-drought periods. The mortality rates of dry-habitat species were less affected by the drought both in dry and wet conditions. A similar pattern was also found in common Dipterocarpaceae species; mortality rates increased more in species associated with wet-habitat in the drought and post-drought periods. Species and families with higher mortality in the pre-drought period tended to experience greater mortality increases during the drought and post-drought periods. These results suggest that changes in drought regimes alter the species composition and spatial distribution of dipterocarp forests.

We analyzed sex ratio, growth rates, and spacing among individuals of Podocarpus nagi, a dioecious tree, on Mt. Mikasa, Nara City, Japan. The sex ratio of reproductive trees ‡ 5 cm in stem diameter at breast height (dbh, 130 cm above ground level) was significantly male-biased. The sex ratio was male-biased in the < 20 cm and ‡ 50 cm size classes, while it did not depart from 1:1 in the 20 £ dbh < 50 cm class. Growth rate varied with tree size in males but not in females. The precocity and vigor of males suggests that differences in reproductive costs between sexes induce the biased sex ratio. Random labeling tests on the positions of reproductive trees showed that in the < 30 cm class, males and females were distributed randomly and independently from each other. In the ‡ 30 cm class, males were significantly clumped, whereas females were randomly distributed. Males and females showed significant repulsion, i.e., a spatial segregation of sexes. Both intra-and intersexual effects on the growth rate of crowding by neighbors were significant for females, but not for males. Maximum competitive interference was observed at a distance of 5 m, which corresponded approximately to the radius of clumps of large males and to the significant repulsive distance between large males and females. These results suggest that sexual differences in sensitivity to local crowding are related to the formation of gender-dependent spatial patterns. Formation of female-repulsive male clumps and a male-biased sex ratio may intensify the decreased probability of regeneration near males, as suggested by the limited seed-dispersal range of this species, thereby promoting coexistence with other species.

Fine-scale spatial genetic structure is increasingly recognized as an important factor in the studies of tropical forest trees as it influences genetic diversity of local populations. The biologic mechanisms that generate fine-scale spatial genetic structure are not fully understood. We studied fine-scale spatial genetic structure in ten coexisting dipterocarp tree species in a Bornean rain forest using microsatellite markers. Six of the ten species showed statistically significant fine-scale spatial genetic structure. Fine-scale spatial genetic structure was stronger at smaller spatial scales ( 100 m) than at larger spatial scales (> 100 m) for each species. Multiple regression analysis suggested that seed dispersal distance was important at the smaller spatial scale. At the larger scale (> 100 m) and over the entire sample range (0-1000 m), pollinators and spatial distribution of adult trees were more important determinants of fine-scale spatial genetic structure. Fine-scale spatial genetic structure was stronger in species pollinated by less mobile small beetles than in species pollinated by the more mobile giant honeybee (Apis dorsata). It was also stronger in species where adult tree distributions were more clumped. The hypothesized mechanisms underlying the negative correlation between clump size and fine-scale spatial genetic structure were a large overlap among seed shadows and genetic drift within clumped species.

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The lowland tropical forests of Southeast Asia are dominated by a single family of canopy and emergent trees, the Dipterocarpaceae. The seeds of dipterocarps are gravity or gyration dispersed. Short distance and limited seed dispersal via these mechanisms result in the aggregation of related individuals and strong fine-scale spatial genetic structure (FSGS). In logged and fragmented forests, where gene flow may be disrupted, tree species with strong FSGS are predicted to exhibit increased inbreeding,

The resource allocation for vegetative growth and female reproduction in three tree species of subgenus Cyclobalanopsis (Quercus, Fagaceae), i.e., Q. salicina, Q. sessilifolia, and Q. acuta, were examined on a per-individual basis in two consecutive reproductive seasons, in order to test whether these trees fit the predictions of the masting hypotheses about resource matching versus resource switching. Since the three Quercus species have a biennial fruiting habit, it takes 3 years for the observation of two reproductive events. Female flower and acorn production per tree were investigated by using a seed-trap method and a numerical analysis of seed dispersal. The net production of each individual was estimated as the sum of the annual increase in the dry mass of vegetative organs and reproductive investment per tree. In the data analyses, the three species were pooled, since all 12 sample trees of the subgenus apparently showed masting in the same year, with no exceptions. Female flower and acorn production per individual tree changed considerably between years. The net production per tree increased with tree size, but did not differ between years. Therefore, the reproductive allocation (proportion of a plant's annual assimilated resources which are used for reproduction) differed dramatically between years. On the other hand, within a year, the reproductive allocation increased with increasing net production per tree. These results suggest that the switching of resource allocation between years within an individual are occurring in subgenus Cyclobalanopsis species, and the intensity of the switching increases with increasing tree size.

Abstract:Tropical tree wood density is often related to other species-specific functional traits, e.g. size, growth rate and mortality. We would therefore expect significant associations within tropical forests between the spatial distributions of stand-level wood density and micro-environments when interspecific variation in wood density is larger than intraspecific variation and when habitat-based species assembly is important in the forest. In this study, we used wood cores collected from 515 trees of 72 species in a 15-ha plot in northern Thailand to analyse intra- and interspecific variation in wood density and the spatial association of stand-level wood density. Intraspecific variation was lower than interspecific variation (20% vs. 80% of the total variation), indicating that species-specific differences in wood density, rather than phenotypic plasticity, are the major source of variation in wood density at the study site. Wood density of individual species was significantly negatively related to maximum diameter, growth rate of sapling diameter and mortality of saplings. Stand-level mean wood density was significantly negatively related to elevation, slope convexity, sapling growth rate and sapling mortality, and positively related to slope inclination. East-facing slopes had significantly lower stand-level mean wood densities than west-facing slopes. We hypothesized that ridges and east-facing slopes in the study forest experience strong and frequent wind disturbance, and that this severe impact may lead to faster stand turnover, creating conditions that favour fast-growing species with low wood density.