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Syzygium is a large genus (1200–1500 species) of Old World tropical trees, currently divided into five subgenera containing widely different numbers of species. Syzygium subgenus Perikion was defined by the presence of fibre bundles in the hypanthium wall, although until now this feature has not been investigated or images published. Furthermore, discovery of fibre bundles in certain species outside Syzygium subgenus Perikion calls for reassessment of the subgenus. In this paper, a morphological analysis is presented for all species previously associated with Syzygium subgenus Perikion or known to have fibre bundles. Results indicate the need for description of a new subgenus, Syzygium subgenus Oborapi, characterized by a distinctly goblet-shaped calyx, presence of fibre bundles in the hypanthium/mesocarp, prominent black lenticels on the abaxial leaf surface, ascending ovule orientation and species diversity centred on the Sunda Shelf. Fibre bundles are photographed and documented for the first time from a range of species and at different magnifications. A preliminary list of species is presented for Syzygium subgenus Perikion and Syzygium subgenus Oborapi, with recommendations for further investigation.

Tree size shapes forest carbon dynamics and determines how trees interact with their environment, including a changing climate. Here, we conduct the first global analysis of among-site differences in how aboveground biomass stocks and fluxes are distributed with tree size. We analyzed repeat tree censuses from 25 large-scale (4-52 ha) forest plots spanning a broad climatic range over five continents to characterize how aboveground biomass, woody productivity, and woody mortality vary with tree diameter. We examined how the median, dispersion, and skewness of these size-related distributions vary with mean annual temperature and precipitation. In warmer forests, aboveground biomass, woody productivity, and woody mortality were more broadly distributed with respect to tree size. In warmer and wetter forests, aboveground biomass and woody productivity were more right skewed, with a long tail towards large trees. Small trees (1-10 cm diameter) contributed more to productivity and mortality than to biomass, highlighting the importance of including these trees in analyses of forest dynamics. Our findings provide an improved characterization of climate-driven forest differences in the size structure of aboveground biomass and dynamics of that biomass, as well as refined benchmarks for capturing climate influences in vegetation demographic models.

Pathogenic and mutualistic fungi have contrasting effects on seedling establishment, but it remains unclear whether density-dependent survival and growth are regulated by access to different types of mycorrhizal fungal networks supported by neighbouring adult trees. Here, we conducted an extensive field survey to test how mycorrhizal and pathogenic fungal colonization of arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) seedlings in a subtropical forest respond to density of neighbouring adult trees. In addition, we undertook a hyphal exclusion experiment to explicitly test the role of soil fungal networks in driving density-dependent effects on seedling growth and survival. Conspecific adult density was a strong predictor for the relative abundance of putative pathogens, which was greater in roots of AM than of ECM seedlings, while mycorrhizal fungal abundance and colonization were not consistently affected by conspecific adult density. Both ECM and AM fungal networks counteracted conspecific density-dependent mortality, but ECM fungi were more effective at weakening the negative effects of high seedling density than AM fungi. Our findings reveal a critical role of common fungal networks in mitigating negative density-dependent effects of pathogenic fungi on seedling establishment, which provides mechanistic insights into how soil fungal diversity shapes plant community structure in subtropical forests.

Funding Information UK Natural Environment Research Council. Grant Number: NE/M019160/1 US National Science Foundation. Grant Number: DEB‐2025282 NERC Human‐modified Tropical Forest Programme. Grant Number: NE/M017508/1 Biodiversity And Land‐use Impacts on Tropical Ecosystem Function (BALI). Grant Numbers: NE/K016253/1, NE/K016253/1 Sime Darby Foundation Stability of Altered Forest Ecosystems (SAFE) Project Sabah Biodiversity Council Institute for Tropical Biology and Conservation (ITBC) at the University of Malaysia, Sabah (UMS) Sabah Forest Research Centre (FRC) at Sepilok Sabah Forestry Department SEARRP, Yayasan Sabah (Maliau Basin Conservation Area) Maliau Basin and Danum Valley Management Committees Acknowledgements Fieldwork was supported by Unding Jami, Matheus Henrique Nunes, Rudi Saul Cruz Chino, Milenka Ximena Montoya, and South East Asia Rainforest Research Program (SEARRP) staff. Research was facilitated by Rob Ewers, Laura Kruitbos, Reuben Nilus, Glen Reynolds, and Charles Vairappan. Species identifications were made by Bernadus Bala Ola, Bill McDonald, Alexander Karolus, and MinSheng Khoo. This work also was supported by the UK Natural Environment Research Council (NERC; no. NE/M019160/1, to BB) and the US National Science Foundation (no. DEB‐2025282, to BB). This publication is a contribution from the NERC Human‐modified Tropical Forest Programme (no. NE/M017508/1, to YAT) and Biodiversity And Land‐use Impacts on Tropical Ecosystem Function (BALI) consortium (no. NE/K016253/1, to YM and no. NE/K016253/1, to YAT). The SAFE Project was funded by the Sime Darby Foundation and the UK NERC. The study areas are part of the Global Ecosystems Monitoring Network (GEM) via an ERC Advanced Investigator Award to YM (no. 321131). The project also was supported by the Stability of Altered Forest Ecosystems (SAFE) Project, the Sabah Biodiversity Council (SaBC, permits JKM/MBS.1000‐2/2 JLD.3‐126 and ‐154), the Institute for Tropical Biology and Conservation (ITBC) at the University of Malaysia, Sabah (UMS), the Sabah Forest Research Centre (FRC) at Sepilok, the Sabah Forestry Department, the SEARRP, Yayasan Sabah (Maliau Basin Conservation Area), and the Maliau Basin and Danum Valley Management Committees. Sean Gleason and several anonymous reviewers provided constructive feedback on the manuscript.

• Fine-scale topographic–edaphic gradients are common in tropical forests and drive species spatial turnover and marked changes in forest structure and function. We evaluate how hydraulic traits of tropical tree species relate to vertical and horizontal spatial niche specialization along such a gradient. • Along a topographic–edaphic gradient with uniform climate in Borneo, we measured six key hydraulic traits in 156 individuals of differing heights in 13 species of Dipterocarpaceae. We investigated how hydraulic traits relate to habitat, tree height and their interaction on this gradient. • Embolism resistance increased in trees on sandy soils but did not vary with tree height. By contrast, water transport capacity increased on sandier soils and with increasing tree height. Habitat and height only interact for hydraulic efficiency, with slope for height changing from positive to negative from the clay-rich to the sandier soil. Habitat type influenced trait–trait relationships for all traits except wood density. • Our data reveal that variation in the hydraulic traits of dipterocarps is driven by a combination of topographic–edaphic conditions, tree height and taxonomic identity. Our work indicates that hydraulic traits play a significant role in shaping forest structure across topographic–edaphic and vertical gradients and may contribute to niche specialization among dipterocarp species.

Aluminium (Al) hyper-accumulation is a common trait expressed by tropical woody plants growing on acidic soils. Studies on Al accumulators have suggested that Al addition may enhance plant growth rates, but the functional significance of this trait and the mechanistic basis of the growth response are uncertain. This study aimed to test the hypothesis that differential growth responses to Al among populations of an Al accumulator species are associated with variation in biomass allocation and nutrient uptake. We conducted two experiments to test differential responses to the presence of Al in the growth medium for seedlings of the Al accumulator shrub Melastoma malabathricum collected from 18 populations across Peninsular Malaysia. Total dry mass and relative growth rate of dry mass were significantly greater for seedlings that had received Al in the growth medium than for control plants that did not receive Al, but growth declined in response to 5.0 mM Al addition. The increase in growth rate in response to Al addition was greater for a fast-growing than a slow-growing population. The increase in growth rate in response to Al addition occurred despite a reduction in dry mass allocation to leaves, at the expense of higher allocation to roots and stems, for plants grown with Al. Foliar concentrations of P, K, Mg and Ca increased in response to Al addition and the first axis of a PCA summarizing foliar nutrient concentrations among populations was correlated positively with seedling relative growth rates. Some populations of the Al hyper-accumulator M. malabathricum express a physiological response to Al addition which leads to a stimulation of growth up to an optimum value of Al in the growth medium, beyond which growth declines. This was associated with enhanced nutrient concentrations in leaves, which suggests that Al accumulation functions to optimize elemental stoichiometry and growth rate., The relative growth rate of dry mass was significantly greater for populations of M. malabathricum seedlings that had received Al in the growth medium than for control plants. The increase in growth rate in response to Al addition occurred despite a reduction in dry mass allocation to leaves, at the expense of higher allocation to roots and stems, for plants grown with Al. Foliar concentrations of P, K, Mg and Ca increased in response to Al addition and foliar nutrient concentrations among populations were correlated positively with seedling relative growth rates. Some populations of M. malabathricum express a physiological response to Al addition which leads to a stimulation of growth up to an optimum value of Al in the growth medium, beyond which growth declines.

Background: The spatial structure of tree species diversity may identify the relative importance of positive, negative and neutral interactions among species and the processes that maintain high sp...

Background: Tree species composition at the landscape scale is often tightly associated with underlying soil type in tropical forests. Changes in soil type may have effects on forest structure that drive changes in both light and soil resource availability, since light availability in the understorey tends to be lower in more fertile sites. Plant functional traits may determine species distributions across gradients of light and soil resource availability.Aims: To test whether tree species with contrasting distributions exhibit leaf traits that reflect adaptation to the resources most limiting in their native environment.Methods: We measured foliar nutrient concentrations, stomatal density, leaf δ13C values, leaf mass per area, and leaf lifespan for saplings of nine common dipterocarp species at Sepilok Forest Reserve, Malaysian Borneo, possessing varying associations to soil resource habitats.Results: Species specialised in their adult distribution to nutrient-poor sandstone soils had traits indicative o...

Background: Logging and habitat fragmentation of tropical rain forests may disrupt patterns of gene flow and genetic diversity. Consequently, inbreeding in tree populations may reduce fitness and increase extinction risks, especially among species that are predominantly outcrossing, dependent on biotic pollination and/or display limited seed dispersal such as species of the Dipterocarpaceae.Aims: To test the hypothesis that heterozygosity of individual progeny affects their likelihood of germination and the growth and survival of seedlings.Methods: Standardised measure of multilocus heterozygosity (sMLH) was estimated from seven microsatellite loci for individual progeny collected from 18 mother trees of the large dipterocarp Parashorea tomentella. The relationships among sMLH, germination and seedling growth and survival were determined for the progeny.Results: Seedling survival over 18 months increased with greater sMLH and fresh fruit weight. This result was expressed under all experimentally controlle...

Plant functional traits regulate ecosystem functions but little is known about how co‐occurring gradients of land use and edaphic conditions influence their expression. We test how gradients of logging disturbance and soil properties relate to community‐weighted mean traits in logged and old‐growth tropical forests in Borneo.We studied 32 physical, chemical and physiological traits from 284 tree species in eight 1 ha plots and measured long‐term soil nutrient supplies and plant‐available nutrients.Logged plots had greater values for traits that drive carbon capture and growth, whilst old‐growth forests had greater values for structural and persistence traits. Although disturbance was the primary driver of trait expression, soil nutrients explained a statistically independent axis of variation linked to leaf size and nutrient concentration. Soil characteristics influenced trait expression via nutrient availability, nutrient pools, and pH.Our finding, that traits have dissimilar responses to land use and soil resource availability, provides robust evidence for the need to consider the abiotic context of logging when predicting plant functional diversity across human‐modified tropical forests. The detection of two independent axes was facilitated by the measurement of many more functional traits than have been examined in previous studies.

Aluminium (Al) accumulation is a common trait expressed in at least 60 plant families and particularly prevalent in tropical woody plants. However, the functional significance and genetic or physiological controls on Al accumulation are currently unknown. We tested the hypothesis that differential expression of Al accumulation among wild populations of the Al-accumulating tropical shrub Melastoma malabathricum L. is associated with habitat-related variation in total and exchangeable soil Al concentrations. Mature leaves and seeds were sampled from 20 populations of M. malabathricum growing in six habitats across Peninsular Malaysia, and soil was collected from each site. The seeds were grown in hydroponic solutions comprising 50% Hoagland's solution amended with Al in the form of 1.0 mM AlCl3 to test the hypothesis that differential expression of foliar Al accumulation is an inherited trait. Foliar Al concentrations varied significantly among populations, but were not consistently different among plants growing in different habitats and showed no relationship to total or exchangeable Al concentrations in soils collected at the 20 sites. Mean foliar Al concentration in wild plants was positively correlated with foliar calcium (Ca) concentrations, and with total soil nitrogen (N), Ca and magnesium (Mg) concentrations, across the 20 populations, and Al addition increased foliar concentrations of phosphorus, Ca, Mg and potassium in seedlings. The differential expression of Al accumulation in M. malabathricum populations is uncoupled to local variation in soil Al concentrations, but may be sensitive to local soil-related variation in the availability of other macro-nutrients, in particular N, Ca and Mg. Further research on the factors controlling Al uptake should focus on the plasticity of this trait within populations of Al accumulators and interactions with micro-habitat variation in the availability of the macronutrient cations.

Documenting the scale and intensity of fine-scale spatial genetic structure (FSGS), and the processes that shape it, is relevant to the sustainable management of genetic resources in timber tree species, particularly where logging or fragmentation might disrupt gene flow. In this study we assessed patterns of FSGS in three species of Dipterocarpaceae (Parashorea tomentella, Shorea leprosula and Shorea parvifolia) across four different tropical rain forests in Malaysia using nuclear microsatellite markers. Topographic heterogeneity varied across the sites. We hypothesised that forests with high topographic heterogeneity would display increased FSGS among the adult populations driven by habitat associations. This hypothesis was not supported for S. leprosula and S. parvifolia which displayed little variation in the intensity and scale of FSGS between sites despite substantial variation in topographic heterogeneity. Conversely, the intensity of FSGS for P. tomentella was greater at a more topographically heterogeneous than a homogeneous site, and a significant difference in the overall pattern of FSGS was detected between sites for this species. These results suggest that local patterns of FSGS may in some species be shaped by habitat heterogeneity in addition to limited gene flow by pollen and seed dispersal. Site factors can therefore contribute to the development of FSGS. Confirming consistency in species’ FSGS amongst sites is an important step in managing timber tree genetic diversity as it provides confidence that species specific management recommendations based on species reproductive traits can be applied across a species’ range. Forest managers should take into account the interaction between reproductive traits and site characteristics, its consequences for maintaining forest genetic resources and how this might influence natural regeneration across species if management is to be sustainable. ISSN:1932-6203

Aims We use dendroecological methods to test the hypothesis that variation in topographic position is related to radial growth and phenology for individuals of the endangered tropical tree Dalbergia nigra under uniform conditions of climate and irradiance, and to examine effects of seasonality on plant phenology and growth periodicity.

Spatial statistics are widely used in studies of ecological processes in plant communities, especially to provide evidence of neutral or non‐neutral mechanisms that might support species coexistence. The contribution of such statistics has been substantial, but their ability to identify any links between underlying processes and emergent patterns is not certain. We investigate the ability of a number of spatial statistics to distinguish theorized mechanisms of species coexistence (spatial and temporal niche differentiation, neutrality, the Janzen–Connell effect and heteromyopia) in a simulated plant community. We find that individual statistics differ substantially in their sensitivity to these mechanisms, with those based on nearest neighbour species identities being the most sensitive. These differences are largely robust to changes in the strength of the modelled mechanisms when simulated independently and in combination. The spatial signal of niche differentiation is always distinct in simulations that combine mechanisms. Synthesis. We describe full spatial signals of modelled coexistence mechanisms that are observed consistently across statistics and simulated strengths and combinations of mechanisms, and identify a set of spatial statistics that holds particular promise for empirical studies designed to investigate mechanisms of these kinds.

Recruitment limitation, measured by the failure of some species to arrive and establish in a site, is potentially an important process contributing to the diversity of plant communities. Limited seed production, seed dispersal, and seedling emergence due to abiotic and/or niche-dependent factors result in opportunities for less abundant species to persist. Whereas limitations affecting early phases of plant establishment are relatively well known, the less-examined limitations upon recruitment into more advanced life history stages may also contribute to the maintenance of diversity. We tested this hypothesis by decomposing recruitment limitation into seed limitation, emergence limitation (to seedlings), establishment limitation (saplings) and persistence limitation (adults) in high- and low-diversity communities. We found evidences of recruitment limitation in all ontogenetic stages, but no differences between the examined communities. Limitations in seed, seedling, saplings and adults were explained mainly by abundance in the previous stage. The negative effect of abundance of a given stage on the abundance of the next stage (negative density-dependent plant establishment) was more evident in the high diversity community. There was a reduction in species’ abundance and an increase in species richness from the seed to the adult stages, resulting in an increase in the evenness along ontogenetic stages in both communities; in all stages, the high-diversity community was richer than the low-diversity community. We found evidence that recruitment limitation operates at multiple stages and is density dependent in these two tropical forests. Our results also suggest that these effects may be stronger in the high-diversity community. Thus, recruitment limitation mediated by density dependence across multiple ontogenetic stages has to be considered in future studies of factors maintaining diversity in tropical forests.

International audience

Drought regimes can be characterized by the variability in the quantity of rainfall and the duration of rainless periods. However, most research on plant response to drought has ignored the impacts of rainfall variation, especially with regard to the influence of nonstructural carbohydrates (NSCs) in promoting drought resistance. To test the hypothesis that these components of drought differentially affect NSC dynamics and seedling resistance, we tracked NSC in plant tissues of tropical tree seedlings in response to manipulations of the volume and frequency of water applied. NSC concentrations decreased in woody tissues under infrequent-high watering but increased under no watering. A faster decline of growth relative to stomatal conductance in the no watering treatment was consistent with NSC accumulation as a result of an uncoupling of growth and photosynthesis, while usage of stored NSCs in woody tissues to maintain function may account for the NSC decline under infrequent-high watering. NSCs, and specifically stem NSCs, contributed to drought resistance under severe water deficits, while NSCs had a less clear role in drought resistance to variability in water availability. The contrasting response of NSCs to water variability and deficit indicates that unique processes support seedling resistance to these components of drought.

Summary: Tropical forest above-ground wood production (AGWP) varies substantially along environmental gradients. Some evidence suggests that AGWP may vary between regions and specifically that Asian forests have particularly high AGWP. However, comparisons across biogeographic regions using standardized methods are lacking, limiting our assessment of pan-tropical variation in AGWP and potential causes. We sampled AGWP in NW Amazon (17 long-term forest plots) and N Borneo (11 plots), both with abundant year-round precipitation. Within each region, forests growing on a broad range of edaphic conditions were sampled using standardized soil and forest measurement techniques. Plot-level AGWP was 49% greater in Borneo than in Amazonia (9.73 ± 0.56 vs. 6.53 ± 0.34 Mg dry mass ha -1 a -1 , respectively; regional mean ± 1 SE). AGWP was positively associated with soil fertility (PCA axes, sum of bases and total P). After controlling for the edaphic environment, AGWP remained significantly higher in Bornean plots. Differences in AGWP were largely attributable to differing height-diameter allometry in the two regions and the abundance of large trees in Borneo. This may be explained, in part, by the greater solar radiation in Borneo compared with NW Amazonia. Trees belonging to the dominant SE Asian family, Dipterocarpaceae, gained woody biomass faster than otherwise equivalent, neighbouring non-dipterocarps, implying that the exceptional production of Bornean forests may be driven by floristic elements. This dominant SE Asian family may partition biomass differently or be more efficient at harvesting resources and in converting them to woody biomass. Synthesis. N Bornean forests have much greater AGWP rates than those in NW Amazon when soil conditions and rainfall are controlled for. Greater resource availability and the highly productive dipterocarps may, in combination, explain why Asian forests produce wood half as fast again as comparable forests in the Amazon. Our results also suggest that taxonomic groups differ in their fundamental ability to capture carbon and that different tropical regions may therefore have different carbon uptake capacities due to biogeographic history. North Bornean forests have much greater AGWP rates than those in north-western Amazon when soil conditions and rainfall are controlled for. Greater resource availability and the highly productive dipterocarps may, in combination, explain why these Asian forests produce wood half as fast again as comparable forests in the Amazon. Our results also suggest that taxonomic groups differ in their fundamental ability to capture carbon and that different tropical regions may therefore have different carbon uptake capacities due to biogeographic history.

Background: Connections between mature trees and seedlings via ectomycorrhizal (EcM) hyphal networks existing in dipterocarp-dominated tropical rain forests of South-east Asia could have strong implications for seedling growth and survival and the maintenance of high diversity in such forests. Aim: To test whether EcM hyphal network connections are important for the growth and survival of dipterocarp seedlings. Methods: We conducted four independent experiments that prevented contact of experimental seedlings with an EcM network by using a series of fine meshes and/or plastic barriers. We measured the growth and survival (and foliar δ13C in one experiment) of seedlings of six dipterocarp species over intervals ranging from 11 to 29 months. Results: Seedling growth (diameter, height or leaf number) was unaffected by exclusion from the EcM network in three experiments and there were no differences in foliar δ13C values in the fourth. Seedling survival was reduced following exclusion from the EcM network in one experiment. Our results give little support to the hypothesis that dipterocarp seedlings growing in the shaded forest understorey benefit from being connected, through a common EcM network, to surrounding trees. Conclusions: We suggest that our negative results, in contrast to studies conducted in low diversity boreo-temperate or tropical forests, are due to these high diversity forests lacking host species-specific EcM fungi and therefore providing little opportunity for adaptive support of seedlings via hyphal networks.

This study was supported by the FP7-PEOPLE-2013-IEF Marie-Curie Action – SPATFOREST. Tree data from BCI were provided by the Center for Tropical Forest Science of the Smithsonian Tropical Research Institute and the primary granting agencies that have supported the BCI plot tree census. Data for the liana censuses were supported by the US National Science Foundation grants: DEB-0613666, DEB-0845071, and DEB-1019436 (to SAS). Soil data was funded by the National Science Foundation grants DEB021104, DEB021115, DEB0212284 and DEB0212818 supporting soils mapping in the BCI plot. 1. Prediction of carbon dynamics in response to global climate change requires an understanding of the processes that govern the distribution of carbon stocks. Above ground biomass (AGB) in tropical forests is regulated by variation in soil fertility, climate, species composition and topography at regional scales, but the drivers of fine-scale variation in tropical forest AGB are poorly understood. The factors that control the growth and mortality of individual trees may be obscured by the low resolution of studies at regional scales. 2. In this paper, we evaluate the effects of soil nutrients, topography and liana abundance on the fine-scale spatial distribution of AGB and density of trees for a lowland tropical moist forest in Panama using additive regression models. 3. Areas with larger values of AGB were negatively associated with the presence of lianas, which may reflect competition with lianas and/or the association of lianas with disturbed or open canopy patches within forests. AGB was positively associated with soils possessing higher pH and K concentrations, reflecting the importance of below-ground resource availability on AGB independently of stem density. 4. Synthesis: Our results shed new light the factors that influence fine-scale tree AGB and carbon stocks in tropical forests: liana abundance is the strongest predictor, having a negative impact on tree AGB. The availability of soil nutrients was also revealed as an important driver of fine-scale spatial variation in tree AGB. Postprint

Questions Species-habitat associations may contribute to the maintenance of species richness in tropical forests, but previous research has been conducted almost exclusively in lowland forests and has emphasized the importance of topography and edaphic conditions. Is the distribution of woody plant species in a Peruvian cloud forest determined by microhabitat conditions? What is the role of environmental characteristics and forest structure in habitat partitioning in a tropical cloud forest? Location Cloud Forest, north Peruvian Andes. Methods We examined species-habitat associations in three 1-ha plots using the torus-translation method. We used three different criteria to define habitats for habitat partitioning analyses, based on microtopography, forest structure and both sets of factors. The number of species associated either positively or negatively with each habitat was assessed. Results Habitats defined on the basis of environmental conditions and forest structure discriminated a greater number of positive and negative associations at the scale of our analyses in a tropical cloud forest. Conclusions Both topographic conditions and forest structure contribute to small-scale microhabitat partitioning of woody plant species in a Peruvian tropical cloud forest. Nevertheless, canopy species were most correlated with the distribution of environmental variables, while understorey species displayed associations with forest structure. © 2012 International Association for Vegetation Science.

Summary 1. Attempts to infer underlying ecological process from observed patterns in ecology have been widespread, but have generally relied on first-order (non-spatial) community characteristics such as the species abundance distribution (SAD). This measure has become an important test of several theories of species coexistence, but has proved unsuccessful in distinguishing between them. 2. Spatially explicit data are increasingly available for a range of ecological communities, and analysis methods for these data are well developed. However, relatively little work has investigated the potential of these data for similar inference about mechanisms of coexistence. 3. In this study, we systematically investigate the spatial and non-spatial signals of simulated ecological processes. We include neutral, niche, lottery, Janzen–Connell and heteromyopia models, deriving and comparing first- and second-order measures for the patterns they generate. 4. We find that the SAD is unable to distinguish reliably between underlying models, with random variation in its shape concealing any systematic differences. 5. A new second-order summary measure of spatial structure derived in this paper, in contrast, proves highly sensitive to the type of ecological interaction being modelled, and is robust to random variation. 6. Synthesis. A simple summary measure of the spatial structure of plant communities is presented and found to be a more powerful indicator of underlying process in simulated data than a widely used first-order measure, the SAD. The potential for answering important ecological questions using spatial statistics, particularly concerning mechanisms of coexistence in diverse communities, appears to be great.

Fine-scale spatial genetic structure (FSGS) within plant populations is an emergent property of the recruited adult trees, influenced by pollen and seed mediated gene flow, selection and demographic processes. This study aims to increase our understanding of the individual species traits that contribute to the generation of FSGS in the Dipteropcarpaceae, which is an ecologically and economically important family of tropical trees that dominate lowland forests in Southeast Asia. We examined FSGS in three cooccurring dipterocarp species at a single site in Borneo. Shorea xanthophylla, Parashorea tomentella and Dipterocarpus grandiflorus share limited seed dispersal but differ markedly in flower size, pollinator body size and pollen dispersal. Here we explore the role of pollen dispersal limitation in shaping FSGS in these three species. Using six microsatellite loci, we explore patterns of FSGS and landscape genetic structure and compare these across species. Significant FSGS was observed in S. xanthophylla and P. tomentella, both of which are known to have relatively limited pollen dispersal, but no clear signal of FSGS was observed in D. grandiflorus. Significantly greater FSGS was observed in P. tomentella (Sp = 0.012) than S. xanthophylla (Sp = 0.007) despite greater pollen dispersal in P. tomentella. Bayesian clustering analysis revealed significant structure in P. tomentella at the scale of the forest reserve (4000 ha). We discuss the alternative explanations for the observed patterns of FSGS emphasising the complexity of the mechanisms that can generate FSGS in long-lived trees. The extent of species-specific pollen dispersal is one factor that can contribute to differences in FSGS across species, but is not the only determinant. The observed patterns of FSGS and landscape scale genetic structure in S. xanthophylla and P. tomentella illustrate the potential of gap regeneration to counter act pollen dispersal and contribute to increased aggregation of related individuals (FSGS). © 2010 Elsevier GmbH. All rights reserved.

Summary 1. Understanding why some alien plant species become invasive when others fail is a fundamental goal in invasion ecology. We used detailed historical planting records of alien plant species introduced to Amani Botanical Garden, Tanzania and contemporary surveys of their invasion status to assess the relative ability of phylogeny, propagule pressure, residence time, plant traits and other factors to explain the success of alien plant species at different stages of the invasion process. 2. Species with native ranges centred in the tropics and with larger seeds were more likely to regenerate, whereas naturalization success was explained by longer residence time, faster growth rate, fewer seeds per fruit, smaller seed mass and shade tolerance. 3. Naturalized species spreading greater distances from original plantings tended to have more seeds per fruit, whereas species dispersed by canopy-feeding animals and with native ranges centred on the tropics tended to have spread more widely in the botanical garden. Species dispersed by canopyfeeding animals and with greater seed mass were more likely to be established in closed forest. 4. Phylogeny alone made a relatively minor contribution to the explanatory power of statistical models, but a greater proportion of variation in spread within the botanical garden and in forest establishment was explained by phylogeny alone than for other models. Phylogeny jointly with variables also explained a greater proportion of variation in forest establishment than in other models. Phylogenetic correction weakened the importance of dispersal syndrome in explaining compartmental spread, seed mass in the forest establishment model, and all factors except for growth rate and residence time in the naturalization model. 5. Synthesis . This study demonstrates that it matters considerably how invasive species are defined when trying to understand the relative ability of multiple variables to explain invasion success. By disentangling different invasion stages and using relatively objective criteria to assess species status, this study highlights that relatively simple models can help to explain why some alien plants are able to naturalize, spread and even establish in closed tropical forests.

Summary 1. This article reviews the application of some summary statistics from current theory of spatial point processes for extracting information from spatial patterns of plants. Theoretical measures and issues connected with their estimation are described. Results are illustrated in the context of specific ecological questions about spatial patterns of trees in two forests. 2. The pair correlation function, related to Ripley’s K function, provides a formal measure of the density of neighbouring plants and makes precise the general notion of a ‘plant’s-eye’ view of a community. The pair correlation function can also be used to describe spatial relationships of neighbouring plants with different qualitative properties, such as species identity and size class. 3. The mark correlation function can be used to describe the spatial relationships of quantitative measures (e.g. biomass). We discuss two types of correlation function for quantitative marks. Applying these functions to the distribution of biomass in a temperate forest, it is shown that the spatial pattern of biomass is uncoupled from the spatial pattern of plant locations. 4. The inhomogeneous pair correlation function enables first-order heterogeneity in the environment to be removed from second-order spatial statistics. We illustrate this for a tree species in a forest of high topographic heterogeneity and show that spatial aggregation remains after allowing for spatial variation in density. An alternative method, the master function, takes a weighted average of homogeneous pair correlation functions computed in subareas; when applied to the same data and compared with the former method, the spatial aggregations are smaller in size. 5. Synthesis. These spatial statistics, especially those derived from pair densities, will help ecologists to extract important ecological information from intricate spatially correlated plants in populations and communities.

Background: Disturbance and small-scale variation in environmental conditions are potential factors that influence structure and diversity in tropical forest communities. In the coastal lowland vegetation in Brazil, forests that differ in flooding regimes could differ in these metrics, although long-term data are usually lacking. Aims: Our aim was to determine if a stand in an unflooded forest and a stand in a flooded forest had distinct structural characteristics and patterns of change over a 16-year period. Methods: Individuals ≥4.44 cm in diameter at breast height were recorded on two 0.3-ha plots in 1991 and 2007. The differences in stem density, species richness, community-level mortality and recruitment, and stem growth rates were assessed. Results: The unflooded forest plot had a higher density of smaller individuals, but a substantially lower basal area than the flooded forest plot. Basal area increased by 4% and 20% over 16 years in the flooded and unflooded plots, respectively. Species richness ...

Orthodox, desiccation-tolerant seeds lose desiccation tolerance during germination. Here, we quantify the timing of the loss of desiccation tolerance, and explore the implications of this event for seed mortality and the shape of germination progress curves for pioneer tree species. For the nine species studied, all seeds in a seedlot lost desiccation tolerance after the same fixed proportion of their time to germination, and this proportion was fairly constant across the species (0.63–0.70). The loss of desiccation tolerance after a fixed proportion of the time to germination has the implication that the maximum number of seeds in a seedlot that can be killed by a single drying event during germination (Mmax) increases with an increasing time to 50% germination (t50) and an increasing slope of the germination progress curve. Consequently, to prevent the seed population from becoming highly vulnerable to desiccation-induced mortality, species with a greater t50 would be expected to have a shallower germination progress curve. In conclusion, these data suggest that the loss of desiccation tolerance during germination may constitute a significant, but previously unexplored, source of mortality for seeds in seasonal environments with unpredictable rainfall.

The distribution and spatial pattern of plants in tropical forests have important implications for how species interact with each other and their environments. In this article we use a large-scale permanent census plot to address if the coexistence of 16 co-occurring species of Myristicaceae is aided by topographic and light gradient niche partitioning. We used a second order spatial pattern analysis based on Ripley’s K function to describe species’ distributions and associations among species, and a torus translation procedure to test for associations with three topographically defined habitats. A majority of species displayed spatial aggregation and over half had one or more significant habitat associations. Four species were associated with the ridge habitat, four species with slope habitat, and two with valley habitat. Seven other species showed no habitat association. Within each habitat group, species exhibited a variety of distributions in relation to light availability. However, habitat associations were largely unexplained by differential rates of mortality, growth or recruitment over a 5-year interval. We conclude that although in principle partitioning of the topographic and light environments may double or treble the number of species able to coexist, there is no evidence that partitioning of physical habitats can explain the coexistence of all 16 of these closely related species.

Estimates of the sex ratio and cost of reproduction in plant populations have implications for resource use by animals, reserve design, and mechanisms of species coexistence, but may be biased unless all potentially reproductive individuals are censused over several flowering seasons. To investigate mechanisms maintaining dioecy in tropical forest trees, we recorded the flowering activity, sexual expression, and reproductive effort of all 2209 potentially reproductive individuals within 16 species of Myristicaceae over 4 years on a large forest plot in Amazonian Ecuador. Female trees invested .10 times more biomass than males in total reproduction. Flowering sex ratios were male-biased in four species in � 1 year, and cumulative 4-year sex ratios were male-biased in two species and for the whole family, but different mechanisms were responsible for this in different species. Annual growth rates were equivalent for both sexes, implying that females can compensate for their greater reproductive investment. There was no strict spatial segregation of the sexes, but females were more often associated with specific habitats than males. We conclude that male-biased sex ratios are not manifested uniformly even after exhaustive sampling and that the mechanisms balancing the higher cost of female reproduction are extremely variable.

The extent to which plant communities are determined by resource availability is a central theme in ecosystem science, but patterns of small-scale variation in resource availability are poorly known. Studies of carbon (C) and nutrient cycling provide insights into factors limiting tree growth and forest productivity. To investigate rates of tropical forest litter production and decomposition in relation to nutrient availability and topography in the absence of confounding large-scale variation in climate and altitude we quantified nutrient fluxes via litterfall and leaf litter decomposition within three distinct floristic associations of tropical rain forest growing along a soil fertility gradient at the Sepilok Forest Reserve (SFR), Sabah, Malaysia. The quantity and nutrient content of small litter decreased along a gradient of soil nutrient availability from alluvial forest (most fertile) through sandstone forest to heath forest (least fertile). Temporal variation in litterfall was greatest in the sandstone forest, where the amount of litter was correlated negatively with rainfall in the previous month. Mass loss and N and P release were fastest from alluvial forest litter, and slowest from heath forest litter. All litter types decomposed most rapidly in the alluvial forest. Stand-level N and P use efficiencies (ratios of litter dry mass to nutrient content) were greatest for the heath forest followed by the sandstone ridge, sandstone valley and alluvial forests, respectively. We conclude that nutrient supply limits productivity most in the heath forest and least in the alluvial forest. Nutrient supply limited productivity in sandstone forest, especially on ridge and hill top sites where nutrient limitation may be exacerbated by reduced rates of litter decomposition during dry periods. The fluxes of N and P varied significantly between the different floristic communities at SFR and these differences may contribute to small-scale variation in species composition.

26 Topic: Research Notes Key Author: Matthew I. Daws Correspondence Address: Seed Conservation Department Royal Botanic Gardens, Kew Wakehurst Place Ardingly, West Sussex RH17 6TN UK E-mail: m.daws@rbgkew.org.uk Authors: M.I. Daws, D. Orr, D.F.R.P. Burslem and C.E. Mullins Effect of high temperature on chalazal plug removal and germination in Apeiba tibourbou Aubl. Pages: 221-225 We tested the effects of a two minute wet heat treatment at a range of temperatures on germination of Apeiba tibourbou seeds. Exposure to >50°C for two minutes resulted in an increase in germination from c. 0 to 80% and resulted in the removal of the chalazal cap. Physical removal of the chalazal cap or mechanical scarification of the seed coat allowed water uptake and subsequent germination. Our results indicate that A. tibourbou seeds exhibit physical dormancy that can be overcome with hot water treatment, chalazal cap removal or mechanical scarification of the testa.

Lianas (woody vines) contribute substantially to the diversity and structure of most tropical forests, yet little is known about the importance of habitat specialization in maintaining tropical liana diversity and the causes of variation among forests in liana abundance and species composition. We examined habitat associations, species diversity, species composition, and community structure of lianas at Sepilok Forest Reserve, Sabah, Malaysia in northeastern Borneo among three soil types that give rise to three distinct forest types of lowland tropical rain forest: alluvial, sandstone hill, and kerangas (heath) forest. Alluvial soils are more nutrient rich and have higher soil moisture than sandstone soils, whereas kerangas soils are the most nutrient poor and drought prone. Lianas ≥0.5-cm in diameter were measured, tagged, and identified to species in three square 0.25-ha plots in each forest type. The number of lianas ≥0.5 cm did not differ significantly among forest types and averaged 1348 lianas ha−1, but mean liana stem diameter, basal area, estimated biomass, species richness, and Fisher’s \(\upalpha\) diversity index were all greater for plots in alluvial than sandstone or kerangas forests. Liana species composition also differed greatly among the three habitats, with 71% of species showing significant positive or negative habitat associations. Sandstone forests were intermediate to alluvial and kerangas forests in most aspects of liana community structure and composition, and fewer species showed significant habitat associations with this forest type. Ranking of forest types with respect to liana density, biomass, and diversity matches the ranking in soil fertility and water availability (alluvial > sandstone hill > kerangas). These results suggest that edaphic factors play an important role in maintaining liana species diversity and structuring liana communities.

This study examined whether topography-induced gradients in water potential and leaf litter depth contribute to species coexistence in tropical forests through species-specific effects on seedling emergence and mortality. Seedling emergence and mortality were followed for a period of 12 months in 36 (1 × 2 m) plots on Barro Colorado Island (BCI), Panama. Plots with and without litter were distributed on slope and plateau sites in three catchments. In the absence of manipulations, the lower litter depth on slopes resulted in approximately four times as many emergent seedlings than on plateau sites. However, litter depth had little effect on seedling community composition. By the end of the first dry-season, post-emergence, there were no significant differences in surviving seedling numbers between any treatments. There were differences in the emergent seedling community between slope and plateau sites within the same catchment as well as differences in composition between catchments, suggesting that both niche partitioning and dispersal limitation might play a role in structuring seedling community composition. During the wet-season seedling mortality was highest on slope sites although this pattern was reversed during the dry-season. In both seasons mortality was higher for small-seeded species. These results demonstrate that gradients in water potential related to topography impact on patterns of seedling emergence and mortality although processes in the first year after emergence may be insufficient to explain observed habitat preferences of adult plants.

Summary 1 The difference in gap size requirements observed for neotropical pioneer species is a classic example of habitat partitioning among tropical tree species. Previous research has shown that the three species Miconia argentea , Cecropia insignis and Trema micrantha differ in the minimum gap size that they occupy as adult plants in semi-deciduous tropical forest in Panama. Here we test whether these differences result from variation in seed germination and seedling establishment requirements, or from differences in susceptibility to herbivores. 2 We planted seeds of Miconia argentea and Trema micrantha into artificially created small (25 m 2 ), medium (64 m 2 ) and large (225 m 2 ) gaps in secondary forest, and transplanted seedlings of all three species into small and large gaps inside and outside mesh cages that excluded invertebrate herbivores. 3 Miconia seedlings emerged and established successfully across a broad range of gap sizes, and transplanted seedlings grew slowly but survived well in all gap sizes. By contrast, emergence and survival of Trema seedlings occurred only in the large gaps and transplants died in small gaps. Survival of Cecropia seedlings was intermediate between Trema and Miconia in the small gaps and growth was greater than either species in the large gaps. 4 Herbivore exclosures reduced foliar herbivory on all three species. The median lifespan of Trema seedlings in small gaps was reduced when the seedlings were exposed to herbivores, but survival of the other species was not affected by the exclosure treatment in either gap size. The growth of Miconia seedlings was lower outside exclosures in both gap sizes. 5 Differences in germination and establishment could generate the contrasting gap size distributions observed between Miconia and Trema , but they are partially offset by measured differences in seed bank density. However, patterns of seedling growth and survival reinforce differences among species at the establishment phase and provide a mechanism for the observed differences in gap size distribution at the sapling stage. ‘Cross-overs’ in species growth rates result in specialization to different environments and reduce their survival in unfavourable conditions. We found no evidence for a definitive influence of herbivory as a mechanism driving differences in gap size distribution among species.

Rates of tree growth in tropical forests reflect variation in life history strategies, contribute to the determination of species' distributional limits, set limits to timber harvesting and control the carbon balance of the stands. Here, we review the resources that limit tree growth at different temporal and spatial scales, and the different growth rates and responses of functional groups defined on the basis of regeneration strategy, maximum size, and species' associations with particular edaphic and climatic conditions. Variation in soil water availability determines intra- and inter-annual patterns of growth within seasonal forests, whereas irradiance may have a more important role in aseasonal forests. Nutrient supply limits growth rates in montane forests and may determine spatial variation in growth of individual species in lowland forests. However, its role in determining spatial variation in stand-level growth rates is unclear. In terms of growth rate, we propose a functional classification of tropical tree species which contrasts inherently fast-growing, responsive species (pioneer, large-statured species), from slow-growing species that are less responsive to increasing resource availability (shade-bearers, small-statured species). In a semi-deciduous forest in Ghana, pioneers associated with high-rainfall forests with less fertile soils, had significantly lower growth rates than pioneers that are more abundant in low-rainfall forests with more fertile soils. These results match patterns found in seedling trials and suggest for pioneers that species' associations with particular environmental conditions are useful indicators of maximum growth rate. The effects of variation in resource availability and of inherent differences between species on stand-level patterns of growth will not be independent if the functional group composition of tropical forests varies along resource gradients. We find that there is increasing evidence of such spatial shifts at both small and large scales in tropical forests. Quantifying these gradients is important for understanding spatial patterns in forest growth rates.

The effects of topographic position on water regime in a semideciduous tropical forest on Barro Colorado Island in Panama were assessed by measuring soil matric potential using the filter paper technique and by using measured soil water release characteristics to convert a long-term (20 years) gravimetric water content data-set to matric potential. These were also compared against predictions from a simple water balance model. Soil matric potentials on slope sites were significantly higher than on plateau sites throughout the measurement interval and slopes experienced a shorter duration of drought during the annual dry-season. Measured values of matric potential agreed with those predicted from converting the gravimetric measurements using water release characteristics. Annual duration of drought predicted by the simple water balance model agreed with values determined from the converted long term water content data-set and was able to predict the annual duration of drought on plateau sites. On slope sites, the water balance systematically and significantly overestimated the duration of drought obtained from the water content data-set, suggesting that slope sites were supplied with water from upslope. Predictions of annual drought duration from sites with higher annual rainfall than Barro Colorado Island (BCI), suggest that while plateau sites on BCI experience a water regime consistent with annual rainfall, slopes experience a water regime more similar to that of forests with much higher rainfall. We conclude that such large variations in water regime over small spatial scales may play a role in maintaining high species richness through providing opportunities for niche specialisation and by buffering slopes against possible climate change.

Foliar elemental concentrations are predictors of life-history variation and contribute to spatial patterns in biogeochemical cycling. We examined the contributions of habitat association, local soil environment, and elemental interactions to variation in foliar elemental concentrations in tropical trees using methods that account for phylogeny. We sampled top-soils and leaves of 58 tropical trees in heath forest (HF) on nutrient-poor sand and mixed dipterocarp forest (MDF) on nutrient-rich clay soils. A phylogenetic generalized least squares method was used to determine how foliar nutrient and aluminium (Al) concentrations varied in response to habitat distribution, soil chemistry and other elemental concentrations. Foliar nitrogen (N) and Al concentrations were greater for specialists of MDF than for specialists of HF, while foliar calcium (Ca) concentrations showed the opposite trend. Foliar magnesium (Mg) concentrations were lower for generalists than for MDF specialists. Foliar element concentrations were correlated with fine-scale variation in soil chemistry in phylogenetically controlled analyses across species, but there was limited within-species plasticity in foliar elemental concentrations. Among Al accumulators, foliar Al concentration was positively associated with foliar Ca and Mg concentrations, and negatively associated with foliar phosphorus (P) concentrations. The Al-accumulation trait and relationships between foliar elemental and Al concentrations may contribute to species habitat partitioning and ecosystem-level differences in biogeochemical cycles.

Summary 1 We evaluate the effects of large-scale disturbance on tropical tree communities by examining the population dynamics of all individuals > 4.9 cm in diameter at breast height (d.b.h.) of 12 tree species over 30 years (1964–94) in lowland tropical rain forest on Kolombangara, Solomon Islands. 2 During the study period Kolombangara was struck by four cyclones between 1967 and 1970. The last cyclone caused most damage to canopy structure. Mortality in the 6-month interval spanning the first cyclone was 7.0% of all trees, while mean annual mortality for all other intervals (including those spanning other cyclones) was 1.4–2.2% year−1. Mortality varied between species but was independent of topography and geographical location. 3 Recruitment increased from very low rates (median 0.0% year−1) before the first cyclone to median values of 1.6–3.2% year−1 during 1971–79, i.e. following a lag period of 3.5–8 years after the first cyclone. Recruitment rates were higher on plots showing greater mortality rates during this cyclone. Recruitment and mortality rates were still higher in 1994 than they had been before the 1967–70 cyclones. 4 Mean annual mortality rates were positively correlated with mean annual recruitment rates across species. This relationship reflects a continuum of life-history characteristics and contributes to constancy in the relative abundance of the 12 species when the same sets of plots are compared over all measurement intervals up to 30 years. 5 We conclude that cyclone impacts have only short-term effects on the relative abundance of common tree species on Kolombangara, and do not therefore prevent the establishment of an equilibrium rank abundance hierarchy or create spatial variation in tree species composition. Differences in forest composition across Kolombangara are more likely to have been caused by differential anthropogenic disturbance linked to settlement patterns.

In 1964 a census of all trees > 9.7 cm diameter at breast height (DBH) was conducted on 22 plots totalling 13.2 ha in lowland tropical evergreen rain forest on Kolombangara, Solomon Islands. Over the following 30 yr (1964–1994), populations of all individuals > 4.85 cm DBH of the 12 most common tree species and amounts of disturbance have been monitored on a declining number of these plots (in 1994, nine plots totalling 5.4 ha were still being recorded). Between November 1967 and April 1970, Kolombangara was struck by four cyclones, although only two of these caused substantial amounts of damage to the canopy structure. Multivariate analysis has identified six forest types on Kolombangara (Greig-Smith et al. 1967). The species richness and diversity of trees in the 1964 census, turnover rates of the populations monitored over 1964–1975, and the amount of disturbance sustained during a cyclone in 1970, were all positively correlated across five of the forest types. The sixth forest type was a consistent outlier in these analyses and is believed to have been seriously disturbed by humans about a century ago. The floristics, turnover and disturbance data support Connell's intermediate disturbance hypothesis. The most species-rich forest types contained a higher proportion of fast-growing individuals and species that are early-successional and which have low density timber. Properties of these species rendered them more susceptible to damage when struck by the 1970 cyclone. They showed higher turnover rates because disturbance-dependent species are also characterised by higher mortality and recruitment rates. Thus, periodic cyclones appear to favour the maintenance of differences in species diversity and composition between forest types.

The new species Rafflesia parvimaculata is described from Pahang, Peninsular Malaysia. This species is characterized by its numerous small white warts on the perigone lobes, and also by its slender, unbranched, capitate ramenta that are white in color and densely arranged inside the floral perigone tube. These unique characters distinguish R. parvimaculata from other Rafflesia species. The discovery of this new species brings the total number of Rafflesia species described from Peninsular Malaysia to five

Summary • High foliar concentrations of aluminium (Al) have been reported in numerous plant species, but progress on the understanding of the functional significance of this trait is constrained by the absence of a quantitative analysis of its distribution among plant lineages and across biomes. • We constructed a global dataset of foliar Al and nutrient concentrations for 1044 plant species from literature sources and new data collections in Brunei Darussalam. • Our results provide statistical support for the existence of Al accumulators and non-Al accumulators in global, regional and local floras based on foliar Al concentrations. A value of 1 mg Al g−1 leaf dry mass is a suitable threshold to distinguish between these two groups in a sample of species that lacks any geographical reference. However, a higher threshold foliar Al concentration is required to distinguish between Al accumulators in tropical (2.3–3.9 mg Al g−1 leaf dry mass) than in temperate (1.1 mg Al g−1 leaf dry mass) floras. There was a phylogenetic signal in the foliar concentrations of Al, but phylogeny did not explain the difference in the mean foliar Al concentration between tropical and temperate floras in a phylogenetically controlled analysis. • Phylogeny and soil chemistry are potential factors driving Al accumulation in certain groups of plants.

BACKGROUND AND AIMS In neotropical forests, very small-seeded pioneer species (

Ecologists frequently measure and compare mortality rates and other count-dependent rates of change. The simplest measures employ mortality counts for predetermined populations over a defined census interval (e.g. Harper 1977; Putz & Milton 1983; Connell et al. 1984; Hubbell & Foster 1990; Turner 1990; Osunkoya et al. 1992). More complex formulations are required to allow comparison over varying time periods, because these measures require a knowledge or assumption of how probabilities of death change over time. In many ecological applications this probability is taken to be constant and can therefore be used to define a rate. In this paper we examine some potentially misleading discrepancies that occur in the recent ecological literature. In its simplest form a constant mortality is modelled by exponential population decline

1 Two bioassays of growth limitation were carried out for seedlings of four shadetolerant tree species (Antidesma cuspidatum, Calophyllum tetrapterum, Dipterocarpus kunstleri and Garcinia scortechinii) growing in P-deficient soil taken from lowland dipterocarp forest in Singapore, as a test of the hypothesis that growth would be limited by the availability of phosphorus. 2 Seedlings of only one species, Antidesma cuspidatum, showed increased growth in response to increased nutrient supply and in that case the limiting nutrient was not P. A majority of seedlings of Antidesma, Calophyllum and Garcinia in this experiment possessed VA mycorrhizas. 3 For seedlings of Antidesma, addition of magnesium led to an increase in the concentration of Mg in all fractions and a positive relation between Mg concentrations and dry mass yield. Addition of potassium and calcium resulted in reductions in concentrations of these elements in the leaves of Antidesma. 4 Seedlings of Antidesma, Calophyllum and Dipterocarpus responded to P by altering distribution of dry mass between different plant parts; the pattern of response varied between species. Phosphorus taken up in excess of requirements for vegetative growth was transferred to plant stems rather than leaves. 5 The outcome of pot bioassays may be dependent on factors such as pot size, irradiance and soil moisture conditions; therefore conclusions drawn here need to be tested by field fertilization experiments.

C. V. S. GUNATILLEKE, I. A. U. N. GUNATILLEKE, G. A. D. PERERA, D. F. R. P. BURSLEM*, P. M. S. ASHTONt and P. S. ASHTONt Department of Botany, University of Peradeniya, Peradeniya, Sri Lanka, *Department of Plant and Soil Science, University of Aberdeen, Cruickshank Building, St. Machar Drive, Aberdeen AB24 3UU, UK, tSchool of Forestry and Environmental Studies, Yale University, New Haven, CT 06511 and $Arnold Arboretum, Harvard University, 22 Divinity Avenue, Cambridge, MA 02138, USA