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What causes individual tree death in tropical forests remains a major gap in our understanding of the biology of tropical trees and leads to significant uncertainty in predicting global carbon cycle dynamics. We measured individual characteristics (diameter at breast height, wood density, growth rate, crown illumination and crown form) and environmental conditions (soil fertility and habitat suitability) for 26 425 trees ≥ 10 cm diameter at breast height belonging to 416 species in a 52-ha plot in Lambir Hills National Park, Malaysia. We used structural equation models to investigate the relationships among the different factors and tree mortality. Crown form (a proxy for mechanical damage and other stresses) and prior growth were the two most important factors related to mortality. The effect of all variables on mortality (except habitat suitability) was substantially greater than expected by chance. Tree death is the result of interactions between factors, including direct and indirect effects. Crown form/damage and prior growth mediated most of the effect of tree size, wood density, fertility and habitat suitability on mortality. Large-scale assessment of crown form or status may result in improved prediction of individual tree death at the landscape scale.

Arbuscular mycorrhizal fungi (AMF) and ectomycorrhizal fungi (EMF) produce contrasting plant–soil feedbacks, but how these feedbacks are constrained by lithology is poorly understood. We investigated the hypothesis that lithological drivers of soil fertility filter plant resource economic strategies in ways that influence the relative fitness of trees with AMF or EMF symbioses in a Bornean rain forest containing species with both mycorrhizal strategies. Using forest inventory data on 1245 tree species, we found that although AMF-hosting trees had greater relative dominance on all soil types, with declining lithological soil fertility EMF-hosting trees became more dominant. Data on 13 leaf traits and wood density for a total of 150 species showed that variation was almost always associated with soil type, whereas for six leaf traits (structural properties; carbon, nitrogen, phosphorus ratios, nitrogen isotopes), variation was also associated with mycorrhizal strategy. EMF-hosting species had slower leaf economics than AMF-hosts, demonstrating the central role of mycorrhizal symbiosis in plant resource economies. At the global scale, climate has been shown to shape forest mycorrhizal composition, but here we show that in communities it depends on soil lithology, suggesting scale-dependent abiotic factors influence feedbacks underlying the relative fitness of different mycorrhizal strategies.

Although leaf toughness is an essential plant adaptation to herbivore pressure and environmental stress, the relationships of leaf toughness with leaf anatomy and photosynthetic traits, and its spatial variations within tropical rainforests, remain poorly understood. We measured these traits in 103 tree species belonging to 27 families from the canopy to understory using a canopy crane system in a tropical rainforest in Sarawak, Malaysia. We focused on the leaf anatomical trait of bundle-sheath extensions (BSEs) around the vascular bundle due to their diverse ecophysiological functions. We divided the trees into heterobaric species with BSEs and homobaric species lacking BSEs, to investigate the relationships of leaf toughness with tree height, leaf functional traits such as carbon (C) and nitrogen (N) content, thickness, leaf mass per area (LMA) and the maximum photosynthetic rate ( P max ). Leaf toughness, LMA, thickness and C and N contents increased with height regardless of BSE presence. Heterobaric leaves had greater toughness than homobaric leaves, whereas leaf thickness, LMA and C were similar between the two leaf types throughout the height gradient. We found that standardized toughness per thickness or C was greater in heterobaric species, as BSEs consist mainly of fibrous tissue. P max was higher for heterobaric than homobaric leaves in the upper canopy presumably due to the functions of BSEs, including water conductivity, but did not differ with plant type in the lower layers. In other words, heterobaric species efficiently exploit the advantages of tougher leaves and higher P max by having BSEs. The increased proportion of heterobaric species, with their tougher leaves and higher P max , in the upper canopy is consistent with adaptation to physically stressful conditions in the tropical rainforest canopy, including high herbivore pressure and strong light.

To predict the dynamics of tropical rainforest ecosystems in response to climate change, it is necessary to understand the drought tolerance and related mechanisms of trees in tropical rainforests. In this study, we assessed the ecophysiological responses of seedlings of six dipterocarp species ( Dipterocarpus pachyphyllus, Dryobalanops aromatica, Shorea beccariana, S. curtisii, S. parvifolia , and S. smithiana ) to experimental short-term drought conditions. The seedlings were initially grown in plastic pots with sufficient irrigation; irrigation was then stopped to induce drought. Throughout the soil-drying period, we measured various ecophysiological parameters, such as maximum photosynthetic and transpiration rates, stomatal conductance, water-use efficiency, predawn water potential, the maximum quantum yield of photosystem II ( F v /F m ), leaf water characteristics (using pressure-volume curves), leaf water content, and total sugar and starch contents. In all six dipterocarp species studied, the F v /F m values dropped sharply when the soil water content fell below 8%. However, there were interspecific differences in physiological responses to such a decrease in soil water content: S. parvifolia and S. beccariana actively controlled their stomata during drought to reduce water consumption via an isohydric response, but showed an increase ( S. parvifolia ) or no change ( S. beccariana ) in leaf drought tolerance; Di. pachyphyllus and Dry. aromatica maintained photosynthesis and transpiration close to the wilting point during drought without reducing water consumption via an anisohydric response, and also increased their leaf drought tolerance over the drying period; and S. curtisii and S. smithiana maintained their photosynthetic capacity without stomatal closure, but showed no change or a slight decrease in leaf drought tolerance. Our results indicate that extreme drought can cause the death of dipterocarp seedlings via various drought response, which could substantially impact the future distribution, population dynamics, and structure of tropical rainforests.

Mycorrhizae, a form of plant–fungal symbioses, mediate vegetation impacts on ecosystem functioning. Climatic effects on decomposition and soil quality are suggested to drive mycorrhizal distributions, with arbuscular mycorrhizal plants prevailing in low-latitude/high-soil-quality areas and ectomycorrhizal (EcM) plants in high-latitude/low-soil-quality areas. However, these generalizations, based on coarse-resolution data, obscure finer-scale variations and result in high uncertainties in the predicted distributions of mycorrhizal types and their drivers. Using data from 31 lowland tropical forests, both at a coarse scale (mean-plot-level data) and fine scale (20 × 20 metres from a subset of 16 sites), we demonstrate that the distribution and abundance of EcM-associated trees are independent of soil quality. Resource exchange differences among mycorrhizal partners, stemming from diverse evolutionary origins of mycorrhizal fungi, may decouple soil fertility from the advantage provided by mycorrhizal associations. Additionally, distinct historical biogeographies and diversification patterns have led to differences in forest composition and nutrient-acquisition strategies across three major tropical regions. Notably, Africa and Asia’s lowland tropical forests have abundant EcM trees, whereas they are relatively scarce in lowland neotropical forests. A greater understanding of the functional biology of mycorrhizal symbiosis is required, especially in the lowland tropics, to overcome biases from assuming similarity to temperate and boreal regions. Analysing data on the relative abundance of basal area of ectomycorrhizal trees from lowland tropical forests, the authors show that their distribution and abundance are independent of soil quality.

Community phylogenetic analysis is an effective approach to understanding the process of community formation. The phylogenetic tree of the species pool is reconstructed in the first step, and the phylogenetic tree obtained in the second step is used to analyze phylogenetic diversity. Sythetic trees have often been used in the construction of phylogenentic trees; however, in tropical rainforests with many closely related species, synthetic trees contain many unresolved nodes, which may affect the results of phylogenetic structure analysis. Here, we constructed a phylogenetic tree using DNA barcode sequences (rbcL, matK, trnH‐psbA) for 737 tree species from the rainforests of Borneo, which have a high‐species diversity and many closely related species. The phylogenetic tree had fewer polytomies and more branch length variations than the Phylocom synthetic trees. Comparison of community phylogenetic analyses indicated that values of the standardized effect size of mean pairwise distance (SES–MPD) were highly correlated between Phylocom and DNA barcode trees, but less so for the standardized effect size of mean nearest taxon distance (SES–MNTD), suggesting that caution is needed when using synthetic trees for communities containing many congeneric species, especially when using SES–MNTD. Simulation analysis suggested that spatial dependence on phylogenetic diversity is related to the phylogenetic signal of the species' habitat niche and the spatial structure of habitat, indicating the importance of detailed phylogeny in understanding community assembly processes. We determined DNA barcode sequences of 737 tree species in the Borneo rainforest, which has a high species diversity with many closely related species. Comparative analysis of community phylogeny suggested that caution should be exercised in the use of synthetic phylogenetic trees constructed with Phylocom and other tools for highly diverse forests. Simulation analysis was also conducted to demonstrate the significance of DNA barcode phylogeny in the study of community assembly processes.

Most canopy species in lowland tropical rain forests in Southeast Asia, represented by Dipterocarpaceae, undergo mast reproduction synchronously at community level during a general flowering event. Such events occur at irregular intervals of 2–10 years. Some species do not necessarily participate in every synchronous mast reproduction, however. This may be due to a lack of carbohydrate resources in the trees for masting. We tested the hypothesis that interspecific differences in the time required to store assimilates in trees for seed production are due to the frequency of masting and/or seed size in each species. We examined the relationship between reproductive frequency and the carbon accumulation period necessary for seed production, and between the seed size and the period, using radiocarbon analysis in 18 dipterocarp canopy species. The mean carbon accumulation period was 0.84 years before seed maturation in all species studied. The carbon accumulation period did not have any significant correlation with reproductive frequency or seed size, both of which varied widely across the species studied. Our results show that for seed production, dipterocarp masting species do not use carbon assimilates stored for a period between the masting years, but instead use recent photosynthates produced primarily in a masting year, regardless of the masting interval or seed size of each species. These findings suggest that storage of carbohydrate resources is not a limiting factor in the masting of dipterocarps, and that accumulation and allocation of other resources is important as a precondition for participation in general flowering.

Numerous studies have shown reduced performance in plants that are surrounded by neighbours of the same species 1 , 2 , a phenomenon known as conspecific negative density dependence (CNDD) 3 . A long-held ecological hypothesis posits that CNDD is more pronounced in tropical than in temperate forests 4 , 5 , which increases community stabilization, species coexistence and the diversity of local tree species 6 , 7 . Previous analyses supporting such a latitudinal gradient in CNDD 8 , 9 have suffered from methodological limitations related to the use of static data 10 – 12 . Here we present a comprehensive assessment of latitudinal CNDD patterns using dynamic mortality data to estimate species-site-specific CNDD across 23 sites. Averaged across species, we found that stabilizing CNDD was present at all except one site, but that average stabilizing CNDD was not stronger toward the tropics. However, in tropical tree communities, rare and intermediate abundant species experienced stronger stabilizing CNDD than did common species. This pattern was absent in temperate forests, which suggests that CNDD influences species abundances more strongly in tropical forests than it does in temperate ones 13 . We also found that interspecific variation in CNDD, which might attenuate its stabilizing effect on species diversity 14 , 15 , was high but not significantly different across latitudes. Although the consequences of these patterns for latitudinal diversity gradients are difficult to evaluate, we speculate that a more effective regulation of population abundances could translate into greater stabilization of tropical tree communities and thus contribute to the high local diversity of tropical forests. An analysis of tree survival data from forest sites worldwide shows that in the tropics, rare tree species experience stronger stabilizing density dependence than common species, wheras no correlation of stabilizing density dependence and abundance exists in the temperate zone.

Summary What causes individual tree death in tropical forests remains a major gap in our understanding of the biology of tropical trees and leads to significant uncertainty in predicting global carbon cycle dynamics. We measured individual characteristics (diameter at breast height, wood density, growth rate, crown illumination and crown form) and environmental conditions (soil fertility and habitat suitability) for 26 425 trees ≥ 10 cm diameter at breast height belonging to 416 species in a 52‐ha plot in Lambir Hills National Park, Malaysia. We used structural equation models to investigate the relationships among the different factors and tree mortality. Crown form (a proxy for mechanical damage and other stresses) and prior growth were the two most important factors related to mortality. The effect of all variables on mortality (except habitat suitability) was substantially greater than expected by chance. Tree death is the result of interactions between factors, including direct and indirect effects. Crown form/damage and prior growth mediated most of the effect of tree size, wood density, fertility and habitat suitability on mortality. Large‐scale assessment of crown form or status may result in improved prediction of individual tree death at the landscape scale.

Resource allocation within trees is a zero-sum game. Unavoidable trade-offs dictate that allocation to growth-promoting functions curtails other functions, generating a gradient of investment in growth versus survival along which tree species align, known as the interspecific growth–mortality trade-off. This paradigm is widely accepted but not well established. Using demographic data for 1,111 tree species across ten tropical forests, we tested the generality of the growth–mortality trade-off and evaluated its underlying drivers using two species-specific parameters describing resource allocation strategies: tolerance of resource limitation and responsiveness of allocation to resource access. Globally, a canonical growth–mortality trade-off emerged, but the trade-off was strongly observed only in less disturbance-prone forests, which contained diverse resource allocation strategies. Only half of disturbance-prone forests, which lacked tolerant species, exhibited the trade-off. Supported by a theoretical model, our findings raise questions about whether the growth–mortality trade-off is a universally applicable organizing framework for understanding tropical forest community structure. Using demographic data for 1,111 tree species across ten tropical forests, the authors test the generality of the growth–mortality trade-off, finding that it holds in undisturbed but not disturbed forests.