Vatica cauliflora P.S. Ashton (Dipterocarpaceae) is a threatened tree species endemic to Kapuas Hulu District, West Kalimantan Province, Indonesia. The species is only known from the type specimen collected in 1953. After this first collection, there was no record confirming the presence of this tree in its natural habitat. Our recent surveys in 2019 and 2020 located 179 individuals of the species in six unprotected locations. The population's size structure is dominated (62.6%) by young individuals within the 0–5 cm diameter class. Our surveys also showed that the habitat of V. cauliflora is degraded as a result of the negative effects of agriculture and logging. Assessment with the IUCN Red List criteria indicates that V. cauliflora should be categorized as Critically Endangered.

Dipterocarpaceae, a dominant family of trees in South-East Asian tropical forests, are remarkable in that they exhibit supra-annual mass-flowering events. The flowering patterns are related to the El Niño Southern Oscillation, but the mechanism that precipitates mass-flowering is still debated. Here, we test if a cumulative-trigger model that tracks resource availability, specifically light, may better explain dipterocarp phenology than a direct-environmental-trigger mechanism. Using 11 flowering time series with an average length of 29 y and variety of candidate predictor variables (precipitation, cloud cover, minimum temperature and El Niño indices) we could not find a plausible direct-environmental-trigger (median AUCs across regions from 0.53 to 0.57 indicating near random predictions). The cumulative-trigger model based on El Niño indices showed better predictive results (AUC 0.67), which could further be improved by resetting the resource at known flowering events (AUC 0.76). Additional support for a cumulative-trigger model comes from the observation that regional differences in the time of year of peak flowering correspond to where El Niño effects are strongest. We conclude that cumulative resource tracking is an evolutionary plausible trigger mechanism that has other primary evolutionary advantages, such as predator satiation.

South-East Asian tropical rain forests experience sporadic, but profuse, seed production after general flowering, leading to the synchronous emergence of various seedlings and subsequent seedling dynamics, which play a crucial role in determining species distribution and coexistence. We examined the relative importance of both biotic (initial height, conspecific seedling density) and abiotic (canopy openness, per cent sand, soil water content) drivers using survival data for 1842 seedlings of 12 dipterocarp species for 1.5 y following mast fruiting in an old-growth Bornean tropical rain forest. More than 30% of all dipterocarp seedlings survived 1.5 y after mast fruiting. When all species were analysed together, we found that initial seedling height, canopy openness and conspecific seedling density affected dipterocarp seedling survival. Negative density dependence indicated that predators were not satiated, but dipterocarp seedlings rather suffered from host-specific natural enemies or intraspecific competition. Species-level analyses of seven dipterocarp species showed large variation in response to biotic and abiotic factors. These results suggest that interspecific differences in the relative importance of biotic and abiotic effects on seedling survival might contribute to species coexistence.

The responses of plants to shade and foliar herbivory jointly affect growth rates and community assembly. We grew 600 seedlings of ten species of the economically important Dipterocarpaceae in experimental gradients of shading (0.3–47.0% of full sunlight) and defoliation (0, 25%, 50% or 75% of leaf area removed). We assessed stem diameters initially, after 2 and 4 mo, and calculated relative growth rates (RGR) with a linear model. Shading interacted with defoliation, reducing RGR by 21.6% in shaded conditions and 8.9% in well-lit conditions. We tested three hypotheses for interspecific trade-offs in growth responses to shading and defoliation. They could be positively related, because both reduce a plant's access to carbon, or inversely related because of trade-offs between herbivore resistance and tolerance. We observed, however, that species varied in their response to shading, but not defoliation, precluding an interspecific trade-off and suggesting that plants tolerate shade and herbivory with differing strategies. Shading most strongly reduced the growth of species with less-dense wood and larger seeds. The strong and variable growth responses to shade, contrasted with the weak and uniform responses to defoliation, suggest that variation in light availability more strongly affects the growth of tropical tree seedlings, and thus community assembly, than does variation in herbivory.

The family Dipterocarpaceae includes 470 tree species from 13 genera in South and South-East Asian tropical forests (Ashton 1982). Many dipterocarp species in aseasonal lowland rain forests of western Malesia flower synchronously during masting (or general flowering) events, which usually occur at irregular intervals of 2–10 y (Ashton et al. 1988). Very few individuals flower at other times, and successful recruitment of seedlings is limited to those masting events (Ashton et al. 1988, Curran et al. 1999).

Stemonoporus Thw. (Dipterocarpaceae), a genus endemic to Sri Lanka, contains as many as 26 species, all but one confined to the perhumid forests of the island. Several species reach 1800 m, the highest altitude recorded for any dipterocarp species in Sri Lanka. We report the first quantitative data on Stemonoporus-domirated stands in the montane zone (1500–1700 m). Stemonoporus cordifolius and 5. rigidus show clear single-species dominance of their respective stands. Stemonoporus gardneri is a co-dominant with Palaquium rubiginosum in its two stands. All Stemonoporus species appear to have a unique geographical and ecological range and no two Stemonoporus species co-exist in our stands.

Numbers of lianas ≥2 cm gbh were recorded on trees ≥30 cm gbh in two 4 ha plots of dipterocarp forest. Associations between lianas and trees of the 12 commonest families and 16 commonest species were analysed. Liana density was 882 ha-1. Fifty-seven per cent of trees supported lianas and the mean number lianas per tree was 2.1. Lianas were highly aggregated on individual trees in most families; numbers of lianas per tree followed the negative binomial distribution for the Dipterocarpaceae but not Euphorbiaceae. Dipterocarpaceae and Euphorbiaceae differed in susceptibility, with 41 and 64% of their trees respectively supporting lianas. The proportions of trees with lianas (i.e. % trees laden) were correlated between plots at the species, but not the family, level. They were not related to tree girth; differences in proportions between families and species were maintained for trees in the 30–40 cm gbh class. At the species level, proportions of trees with lianas and the mean number of lianas per tree were significantly inversely correlated with mean branch-free bole height of trees in the 30–40 cm gbh class. Increased branch-free bole height and associated branch shedding may therefore decrease the susceptibility of trees to lianas and explain the lower susceptibility in the Dipterocarpaceae than the Euphorbiaceae.

Seeds of Dryobalanops aromatica and D. lanceolata (Dipterocarpaceae) were sown in gaps and understorey on a ridge and a valley in a Bornean rainforest. Germination rates were 92–100% regardless of site and species. On the ridge, the proportion of seedlings that opened primary leaves was larger in D. aromatica (19–30%) than in D. lanceolata (7–19%) due to higher root predation on D. lanceolata. In the valley, on the other hand, the proportion was larger in D. lanceolata (71–74%) than in D. aromatica (56%). After primary leaf production, seedlings of both species had higher survivorship in the gaps than in the understorey. Interactions between predators, soil conditions and light environment and their effects on seedling establishment are discussed.

Tropical rain forest is described as a mosaic of ecologically uniform patches. A method is proposed to recognize and inventory the patches and to draw mosaic maps, based on the size and architecture of canopy trees. Mosaic maps from three different forest localities in Indonesia are compared in terms of the relative area occupied by each kind of patch, and the surface range and number of constituent patches. The comparison shows that the mosaic patterns are specific to the sites and can be used for the characterization of patch dynamics in different forest types. Possible implications for forest management are discussed in the light of the mosaic maps.

Site data from 291 forest inventory plots in Mixed Dipterocarp Forest in central Sarawak were simplified by factor analysis. The distribution of the 33 commonest canopy tree species was related to site factor scores by ‘t’ tests. There appears to be a degree of edaphic influence on the distribution of many species. Fairly constant soil characteristics related to the lithology of the parent material appear to be more important than ephemerally variable pro perties such as organic matter or exchangeable cations. Magnesium appears to be particularly important, possibly because of an effect on the capacity of mycorrhizal root systems to absorb phosphorus.

Nanophyes shoreae is a pre-dispersal fruit-predator of several species of Dipterocarpaceae. The timing of oviposition of this weevil was monitored in six trees in Pasoh Forest Reserve, West Malaysia from September 1986 to February 1987. A flowering event occurred during this period. Oviposition started within 12 days after peak anthesis, and continued within any one tree for 11–39 days. Host trees of N. shoreae flowered over a 75 day period. Intra-specific flowering synchrony was high but different species flowered sequentially. The behaviour of N. shoreae and the period of the oviposition ‘window’ strongly suggest that this pre-dispersal fruit-predator ‘tracks’ the sequence of flowering shown by its host species. The implications for predator satiation through mass-flowering of the Dipterocarpaceae are discussed.

The relationship between sepal and nut size of fruits was studied in fruit of 394 species of Dipterocarpaceae in Malesia and Sri Lanka. The fruits of many dipterocarps have long twisted sepals which have a role as blades for dispersal by gyration. But long sepals decrease resource allocation to other parts, and increase the chance of fruit being trapped in the canopy. The frequency distribution of the ratio of sepal length to nut length was bimodal. The gyrationdispersed group with longer sepals comprised 74% of Dipterocarpaceae studied. The group with short sepal mainly comprised understorey trees, but there were a few emergents. In both groups, sepal areas were expressed as a power function of nut volumes'. Because fruit weight per sepal area increases with fruit size, larger fruits fall more quickly than smaller fruits, and sepals of the former can not serve so effectively as propeller blades. This is consistent with the fact that a few emergent trees with large nuts have short sepals and are not dispersed by gyration. Therefore, the re-evolution of short fruit sepals appears to have been mainly due to two reasons: reduction of tree height and enlargement of nut size.

Bukit Belalong, Brunei, is a small mountain (913 m) of uniform shale lithology with continuous primary rain forest from its base to its summit. Three 0.25 ha plots were established at each of three altitudes (200, 500 and 850 m) to investigate the altitudinal zonation of the soils and the vegetation. One soil profile from each altitude is described and chemical analyses of the soils indicate that they are similar at all altitudes. In each plot all trees ≥10 cm dbh were measured and identified as far as possible. Maximum tree height was greatest at 200 m (60 m) and least at 850 m (33 m). Tree density (number of individuals ha−1) increased with altitude. The forest at 850 m was the most species- and family-rich, but since the ratio of species to individuals did not vary significantly among altitudes, the higher species richness is attributed partly to the larger number of trees sampled. The Dipterocarpaceae was the most important family in terms of basal area at all altitudes, but its proportion of basal area was much smaller at 850 m than at the other altitudes. The Dipterocarpaceae was the most diverse family at 200 m and 500 m, but at 850 m its diversity was exceeded by the Lauraceae, Myrtaceae and Euphorbiaceae and equalled by the Clusiaceae. The forests at 200 m and 500 m are classified as lowland rain forest and that at 850 m is classified as a lower montane forest.

The general flowering of lowland dipterocarp forests of South-east Asia is a unique phenomenon that occurs intermittently, sometimes many years apart, and may be widespread throughout the region or sporadic. During a general flowering a very large number of tree families, including the well-known Dipterocarpaceae, flower and fruit exceptionally heavily. The huge pulse of flowering is echoed by an apparent explosion in the number of pollinators, and this is further augmented by migration of pollinators from the forest fringes. The huge demand for pollinators seems to be solved among several dipterocarps through utilization of tiny, fast breeding, floral-feeding insects such as thrips and hoppers. Competition for pollinators is further reduced through development of sequential flowering among several groups of trees that share similar pollinators. The resulting massive general fruiting seems to contribute to a heightened level of faunal activity and numbers. This long-intervalled flowering and fruiting phenology which dominates these lowland dipterocarp forests may have partly contributed to the lowered animal biomass frequently alluded to when these forests are compared with neotropical forests.

Spatial distribution patterns and habitat associations of Fagaceae species in a Fagaceae-codominated hill forest in Sumatra were investigated. Ten Fagaceae species believed to be zoochorous (animal-dispersed seed) and five codominant canopy and emergent anemochorous (wind-dispersed seed) species from Anacardiaceae and Dipterocarpaceae were studied. Five Fagaceae species and all codominant anemochorous species were significantly aggregated while the other five Fagaceae species showed a random distribution pattern. The median distance of small saplings from the nearest reproductively mature tree tended to be shorter for aggregated species than spatially random species. This implied that some Fagaceae species dispersed over longer distances than anemochorous species. Relationships between four habitat variables and distribution of the target species were examined with torus-translation tests. Three Quercus and one Lithocarpus species showed positive habitat associations. Two Quercus species aggregated at the preferred habitat, but the others were randomly distributed. Thus tree species with specific habitat preference do not only aggregate at the preferred habitat. The three ridge-specialist Quercus species showed gradual changes in habitat association, which could reflect avoidance of competition among the species. Most of the Lithocarpus species showed little correlation with habitat variables. Coexistence of the three Quercus species partly reflected subtle differences in topographical preferences. Distribution of five of the six Lithocarpus species was unrelated to topography, so other mechanisms must be sought to account for the maintenance of coexistence in this species-rich genus.

The physical and chemical seed characteristics of 11 dipterocarp species (four Dipterocarpus, two Dryobalanops and five Shorea species) were quantified to examine variations among species. We measured dry weight, pericarp thickness and concentrations of nitrogen, total phenol, condensed tannin, lignin, lipid, starch and total sugar. Although there was low intraspecific variation among parent trees, each species had unique characteristics. We found significant correlations between chemical defensive compounds (i.e. lignin and total phenol) and nitrogen concentration. However, these relationships were no longer significant when an outlier was removed. Fruiting frequency was not significantly correlated with any chemical compound. Seeds of Shorea species had a higher concentration of lipid than seeds of Dipterocarpus. The implications of variation in the physical and chemical characteristics of dipterocarp seeds are discussed.

Seedlings of five species in the Dipterocarpaceae were grown in experimental plots in Sabah, Malaysia. These were sited both in gaps and understorey and on alluvial and sandstone soils. Half of all seedlings were provided with a complete fertilizer. Herbivore damage levels were recorded on over 25 000 individual leaves in four surveys over the course of 2 y. Rates of herbivory were lower on mature leaves (0.07–0.8% leaf area mo−1 among species) than new leaves (2.1–4.4% leaf area mo−1). There were no overall effects of light conditions, soil type, fertilizer treatment or time on rates of herbivory. The only consistent source of variation was that between species, with the three alluvial-specialist species suffering higher rates of damage than the two sandstone-specialists. Mature leaves of alluvial species received greater damage in sandstone soils, whereas sandstone species were damaged at equivalent rates on both soil types. New leaves were more damaged on their native soil type. Published herbivory rates vary in the timescales and methods of measurement. Nevertheless, the few comparable studies confirm that herbivory rates on seedlings in tropical rain forests are remarkably constant over time and across experimental treatments.

A prediction of the hypothesis that niche processes control plant species distributions across edaphic gradients is that pairwise comparisons of related species on contrasting soils should show consistent patterns of trait divergence. This hypothesis was tested in lowland Bornean rain forest by combining measurements of leaf traits, soil nutrients and the distribution of tree species in the Dipterocarpaceae across a nutrient gradient. Nine species were studied, comprising four related pairs in Shorea (phylogenetic independent contrasts; PICs) and one habitat generalist (Dipterocarpus sublamellatus). Lamina area, specific leaf area (SLA), leaf nitrogen (N) and phosphorus (P) were measured for ∼10 saplings of each species; habitat associations were defined as a continuous function of soil nutrients and categorically in relation to substrate. Species distributions and traits varied significantly with soil nutrients. When all species were pooled, SLA and leaf P increased significantly with species' distribution across the nutrient gradient; leaf N showed the same trend. Trait shifts with habitat were stronger and more pervasive when habitat was defined categorically – in all four PICs, rich-soil specialists had larger leaves, higher SLA, leaf N and P, and lower N:P ratios. Trait shifts with habitat at least partly reflect intrinsic differences between species. Within Shorea, variation in lamina area and N:P ratio were more strongly related to phylogeny than habitat, whereas the reverse held for SLA and leaf P. Phylogeny also influenced the extent of trait divergence between related species on different soils, and patterns of trait correlation within lineages. Results support the hypothesis that niche processes influence the distribution of species and traits in lowland tropical tree communities, and highlight the value of phylogenetic information for increasing the power of comparative studies.

To investigate whether seedlings of ten dipterocarp species differed significantly in terms of growth and mortality or whether species were not significantly different and could be considered ecologically similar, seedlings were grown, two per pot, in two experiments: (1) where the two seedlings were of equal height (30 cm); and (2) where one seedling was 10 cm shorter than the other. Seedlings were grown in a shade house with 15% above-canopy light in a 50:50 forest soil–sand mixture and were watered frequently; pots were placed so that seedling density was 130 seedlings m−2 of ground. In the first experiment there were 45 pairwise combinations of species when seedlings were 30 cm tall (AB, AC, AD,…. BC, BD…IJ; where A, B, C…J signify different species); each combination was replicated 10 times so there were 450 pots with 900 seedlings. In the second experiment there were 100 pairwise combinations of species and size e.g. Aa (30 cm A with 20 cm a), Ab (30 cm A with 20 cm b), each combination was replicated 10 times hence there were 1000 pots with 2000 seedlings. After 22 mo 79% of the initial 2900 seedlings survived; on average they had grown 42 cm (i.e. to 72 cm tall from their initial 30 cm). The most frequent outcome of competition-trials between different sized individuals (784 of 1000 trials) was that the initially taller seedling of each pair ‘won’ (it was the taller or surviving seedling). When 900 of these trials (setting aside, Aa, Bb, Cc etc.) were analysed as 45 comparisons between species with different sized individuals (Ab and aB are one interspecific comparison for these purposes), initial height determined the outcome in 23 cases (even in some competitions between light hardwood species and heavy hardwood species); in 6 cases a species (mostly light hardwoods) behaved as a ‘dominant’ – they usually won even if they were smaller initially. We found few significant differences between species in: initial seedling heights; leaf nitrogen concentrations; and specific leaf areas when they were grown in similar conditions, and these attributes were not correlated with growth rates. The similarity of seedlings of different species meant that often a height difference of just 10 cm was enough to determine the outcome of a pairwise competition-trial in high seedling densities and light equivalent to that in forest gaps.

Some tree species exhibit large year-to-year variation in seed production, a phenomenon known as masting (Kelly 1994, Kelly & Sork 2002). Even in tropical rain forests, in which the climate is suitable for plant growth all year round with little seasonal variation (Whitmore 1998), there are many reports of masting (Appanah 1993, Hart 1995, Newbery et al. 1998, Newstrom et al. 1994, Wheelwright 1986). In particular, Dipterocarpaceae, the dominant family in lowland mixed dipterocarp forests in South-East Asia, undergo mast fruiting following mass-flowering with strong interspecific synchronization in aseasonal western Malesia (Appanah 1985, 1993; Ashton 1989, Ashton et al. 1988, Curran et al. 1999, Janzen 1974, Medway 1972, Sakai et al. 1999, Whitmore 1998, Wood 1956). In mixed-dipterocarp forests, dipterocarp species contribute more than 70% of the canopy biomass (Bruenig 1996, Curran & Leighton 2000). Masting of dipterocarp species is therefore likely to have a major impact on animal populations, and also on the nutrient cycle in such forest ecosystems by causing fluctuations in the availability of resources (Sakai 2002).