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The relative importance of tree mortality risk factors remains unknown, especially in diverse tropical forests where species may vary widely in their responses to particular conditions. We present a new framework for quantifying the importance of mortality risk factors and apply it to compare 19 risks on 31 203 trees (1977 species) in 14 one-year periods in six tropical forests. We defined a condition as a risk factor for a species if it was associated with at least a doubling of mortality rate in univariate analyses. For each risk, we estimated prevalence (frequency), lethality (difference in mortality between trees with and without the risk) and impact ('excess mortality' associated with the risk, relative to stand-level mortality). The most impactful risk factors were light limitation and crown/trunk loss; the most prevalent were light limitation and small size; the most lethal were leaf damage and wounds. Modes of death (standing, broken and uprooted) had limited links with previous conditions and mortality risk factors. We provide the first ranking of importance of tree-level mortality risk factors in tropical forests. Future research should focus on the links between these risks, their climatic drivers and the physiological processes to enable mechanistic predictions of future tree mortality.

Accurately projecting global carbon dynamics requires understanding controls over temporal and spatial tropical forest biomass variation. Changes in tropical forest aboveground biomass per area are most commonly estimated using repeat forest plot censuses, but errors in these estimates can be relatively large, limiting our ability to detect real changes in forest carbon. One source of error is nonstandard heights of diameter measurement, which are common in tropical forests because buttressed trees are usually measured above buttresses. Nonstandard and temporally variable measurement heights affect biomass change estimates because tree trunks taper—higher measurements result in smaller diameters and, thus, smaller estimated biomass. Modeling trunk taper is a potential way to correct for biases associated with nonstandard and changing measurement heights. We measured trunk taper for 260 buttressed trees in five lowland tropical forests to assess variation in taper within and among sites. Trunks taper more slowly in trees with larger diameters above buttress, taller buttresses, and higher wood density; these variables together explain 29% of taper variation. We also used plot data to quantify the distribution of measurement heights—and potential for bias in biomass estimates. We found significant variation in measurement heights among plots, and over time within plots. At the site level, our general taper model (taper predicted from diameter above buttress, buttress height, and wood density) produced site-level biomass estimates within 3.4%, on average, of estimates from measured taper. We recommend using this model to reduce bias from nonstandard measurement heights in studies of tropical forest biomass variation.

1. Organisms of all species must balance their allocation to growth, survival and recruitment. Among tree species, evolution has resulted in different life-history strategies for partitioning resources to these key demographic processes. Life-history strategies in tropical forests have often been shown to align along a trade-off between fast growth and high survival, that is, the well-known fast–slow continuum. In addition, an orthogonal trade-off has been proposed between tall stature—resulting from fast growth and high survival—and recruitment success, that is, a stature−recruitment trade-off. However, it is not clear whether these two independent dimensions of life-history variation structure tropical forests worldwide. 2. We used data from 13 large-scale and long-term tropical forest monitoring plots in three continents to explore the principal trade-offs in annual growth, survival and recruitment as well as tree stature. These forests included relatively undisturbed forests as well as typhoon-disturbed forests. Life-history variation in 12 forests was structured by two orthogonal trade-offs, the growth−survival trade-off and the stature−recruitment trade-off. Pairwise Procrustes analysis revealed a high similarity of demographic relationships among forests. The small deviations were related to differences between African and Asian plots. 3. Synthesis. The fast–slow continuum and tree stature are two independent dimensions structuring many, but not all tropical tree communities. Our discovery of the consistency of demographic trade-offs and life-history strategies across different forest types from three continents substantially improves our ability to predict tropical forest dynamics worldwide.

The growth and survival of individual trees determine the physical structure of a forest with important consequences for forest function. However, given the diversity of tree species and forest biomes, quantifying the multitude of demographic strategies within and across forests and the way that they translate into forest structure and function remains a significant challenge. Here, we quantify the demographic rates of 1961 tree species from temperate and tropical forests and evaluate how demographic diversity (DD) and demographic composition (DC) differ across forests, and how these differences in demography relate to species richness, aboveground biomass (AGB), and carbon residence time. We find wide variation in DD and DC across forest plots, patterns that are not explained by species richness or climate variables alone. There is no evidence that DD has an effect on either AGB or carbon residence time. Rather, the DC of forests, specifically the relative abundance of large statured species, predicted both biomass and carbon residence time. Our results demonstrate the distinct DCs of globally distributed forests, reflecting biogeography, recent history, and current plot conditions. Linking the DC of forests to resilience or vulnerability to climate change, will improve the precision and accuracy of predictions of future forest composition, structure, and function.

Tree rings provide an invaluable long-term record for understanding how climate and other drivers shape tree growth and forest productivity. However, conventional tree-ring analysis methods were not designed to simultaneously test effects of climate, tree size, and other drivers on individual growth. This has limited the potential to test ecologically relevant hypotheses on tree growth sensitivity to environmental drivers and their interactions with tree size. Here, we develop and apply a new method to simultaneously model nonlinear effects of primary climate drivers, reconstructed tree diameter at breast height (DBH), and calendar year in generalized least squares models that account for the temporal autocorrelation inherent to each individual tree's growth. We analyze data from 3811 trees representing 40 species at 10 globally distributed sites, showing that precipitation, temperature, DBH, and calendar year have additively, and often interactively, influenced annual growth over the past 120 years. Growth responses were predominantly positive to precipitation (usually over ≥3-month seasonal windows) and negative to temperature (usually maximum temperature, over ≤3-month seasonal windows), with concave-down responses in 63% of relationships. Climate sensitivity commonly varied with DBH (45% of cases tested), with larger trees usually more sensitive. Trends in ring width at small DBH were linked to the light environment under which trees established, but basal area or biomass increments consistently reached maxima at intermediate DBH. Accounting for climate and DBH, growth rate declined over time for 92% of species in secondary or disturbed stands, whereas growth trends were mixed in older forests. These trends were largely attributable to stand dynamics as cohorts and stands age, which remain challenging to disentangle from global change drivers. By providing a parsimonious approach for characterizing multiple interacting drivers of tree growth, our method reveals a more complete picture of the factors influencing growth than has previously been possible.

All species must balance their allocation to growth, survival and recruitment. Among trees, evolution has resulted in different strategies of partitioning resources to these key demographic processes, i.e. demographic trade-offs. It is unclear whether the same demographic trade-offs structure tropical forests worldwide. Here, we used data from 13 large-scale and long-term tropical forest plots to estimate the principal trade-offs in growth, survival, recruitment, and tree stature at each site. For ten sites, two trade-offs appeared repeatedly. One trade-off showed a negative relationship between growth and survival, i.e. the well-known fast−slow continuum. The second trade-off distinguished between tall-statured species and species with high recruitment rates, i.e. a stature−recruitment trade-off. Thus, the fast-slow continuum and tree stature are two independent dimensions structuring most tropical tree communities. Our discovery of the consistency of demographic trade-offs and strategies across forest types in three continents substantially improves our ability to predict tropical forest dynamics worldwide.

All species must balance their allocation to growth, survival and recruitment. Among trees, evolution has resulted in different strategies of partitioning resources to these key demographic processes, i.e. demographic trade-offs. It is unclear whether the same demographic trade-offs structure tropical forests worldwide. Here, we used data from 13 large-scale and long-term tropical forest plots to estimate the principal trade-offs in growth, survival, recruitment, and tree stature at each site. For ten sites, two trade-offs appeared repeatedly. One trade-off showed a negative relationship between growth and survival, i.e. the well-known fast−slow continuum. The second trade-off distinguished between tall-statured species and species with high recruitment rates, i.e. a stature−recruitment trade-off. Thus, the fast-slow continuum and tree stature are two independent dimensions structuring most tropical tree communities. Our discovery of the consistency of demographic trade-offs and strategies across forest types in three continents substantially improves our ability to predict tropical forest dynamics worldwide.

In the tropics, antagonistic seed predation networks may have different properties than mutualistic pollination and seed dispersal networks, but the former have been considerably less studied. We tested whether the structure of antagonistic tripartite networks composed of host plants, insects developing within seeds and fruits, and their insect parasitoids could be predicted from plant phylogenetic distance and plant traits. We considered subsets of the networks (“subnetworks”) at three rainforest locations (Panama, Thailand, Papua New Guinea), based on insect families, plant families or plant functional groups. We recorded 3,197 interactions and observed a low percentage of realized interactions, especially in Panama, where insect host specificity was higher than in Thailand or New Guinea. Several factors may explain this, including insect faunal composition, incidence of dry fruits, high fruit production and high occurrence of Fabaceae at the Panamanian site. Host specificity was greater among seed-eaters than pulp-eaters and for insects feeding on dry fruits as opposed to insects feeding on fleshy fruits. Plant species richness within plant families did not influence insect host specificity, but site characteristics may be important in this regard. Most subnetworks were extremely specialized, such as those including Tortricidae and Bruchinae in Panama. Plant phylogenetic distance, plant basal area and plant traits (fruit length, number of seeds per fruit) had important effects on several network statistics in regressions weighted by sampling effort. A path analysis revealed a weak direct influence of plant phylogenetic distance on parasitoid richness, indicating limited support for the “nasty host hypothesis”. Our study emphasizes the duality between seed dispersal and seed predation networks in the tropics, as key plant species differ and host specificity tends to be low in the former and higher in the latter. This underlines the need to study both types of networks for sound practices of forest regeneration and conservation.

Symbiotic nitrogen (N)-fixing trees can provide large quantities of new N to ecosystems, but only if they are sufficiently abundant. The overall abundance and latitudinal abundance distributions of N-fixing trees are well characterised in the Americas, but less well outside the Americas. Here, we characterised the abundance of N-fixing trees in a network of forest plots spanning five continents, ~5,000 tree species and ~4 million trees. The majority of the plots (86%) were in America or Asia. In addition, we examined whether the observed pattern of abundance of N-fixing trees was correlated with mean annual temperature and precipitation. Outside the tropics, N-fixing trees were consistently rare in the forest plots we examined. Within the tropics, N-fixing trees were abundant in American but not Asian forest plots (~7% versus ~1% of basal area and stems). This disparity was not explained by mean annual temperature or precipitation. Our finding of low N-fixing tree abundance in the Asian tropics casts some doubt on recent high estimates of N fixation rates in this region, which do not account for disparities in N-fixing tree abundance between the Asian and American tropics. Synthesis. Inputs of nitrogen to forests depend on symbiotic nitrogen fixation, which is constrained by the abundance of N-fixing trees. By analysing a large dataset of ~4 million trees, we found that N-fixing trees were consistently rare in the Asian tropics as well as across higher latitudes in Asia, America and Europe. The rarity of N-fixing trees in the Asian tropics compared with the American tropics might stem from lower intrinsic N limitation in Asian tropical forests, although direct support for any mechanism is lacking. The paucity of N-fixing trees throughout Asian forests suggests that N inputs to the Asian tropics might be lower than previously thought.

In 2000, we established a 24-ha plot in Peninsular Thailand to investigate how forest composition, structure and dynamics vary with spatial heterogeneity in resource availability. Detailed soil and topographic surveys were used to describe four edaphic habitats in the plot. Disturbance history was inferred from historical records and floristic analysis. The plot included >119 000 trees ≥1 cm dbh in 578 species, and was recensused in 2010. Species distributions, floristic turnover, stand structure, demographic rates and biomass dynamics were strongly influenced by heterogeneity in soils, topography and disturbance history. Over 75% of species were aggregated on specific edaphic habitats leading to strong compositional turnover across the plot. Soil chemistry more strongly affected species turnover than topography. Forest with high biomass and slow dynamics occurred on well-drained, low fertility ridges. The distribution and size structure of pioneer species reflected habitat-specific differences in disturbance history. Overall, above-ground biomass (AGB) increased by 0.64 Mg ha−1 y−1, from 385 to 392 Mg ha−1, an increase that was entirely attributable to recovery after natural disturbance. Forest composition and stand structure, by reflecting local disturbance history, provide insights into the likely drivers of AGB change in forests. Predicting future changes in tropical forests requires improved understanding of how soils and disturbance regulate forest dynamics.

ForestGEO is a network of scientists and long-term forest dynamics plots (FDPs) spanning the Earth's major forest types. ForestGEO's mission is to advance understanding of the diversity and dynamics of forests and to strengthen global capacity for forest science research. ForestGEO is unique among forest plot networks in its large-scale plot dimensions, censusing of all stems ≥1 cm in diameter, inclusion of tropical, temperate and boreal forests, and investigation of additional biotic (e.g., arthropods) and abiotic (e.g., soils) drivers, which together provide a holistic view of forest functioning. The 71 FDPs in 27 countries include approximately 7.33 million living trees and about 12,000 species, representing 20% of the world's known tree diversity. With >1300 published papers, ForestGEO researchers have made significant contributions in two fundamental areas: species coexistence and diversity, and ecosystem functioning. Specifically, defining the major biotic and abiotic controls on the distribution and coexistence of species and functional types and on variation in species' demography has led to improved understanding of how the multiple dimensions of forest diversity are structured across space and time and how this diversity relates to the processes controlling the role of forests in the Earth system. Nevertheless, knowledge gaps remain that impede our ability to predict how forest diversity and function will respond to climate change and other stressors. Meeting these global research challenges requires major advances in standardizing taxonomy of tropical species, resolving the main drivers of forest dynamics, and integrating plot-based ground and remote sensing observations to scale up estimates of forest diversity and function, coupled with improved predictive models. However, they cannot be met without greater financial commitment to sustain the long-term research of ForestGEO and other forest plot networks, greatly expanded scientific capacity across the world's forested nations, and increased collaboration and integration among research networks and disciplines addressing forest science.

© 2018, The Author(s). Survival rates of large trees determine forest biomass dynamics. Survival rates of small trees have been linked to mechanisms that maintain biodiversity across tropical forests. How species survival rates change with size offers insight into the links between biodiversity and ecosystem function across tropical forests. We tested patterns of size-dependent tree survival across the tropics using data from 1,781 species and over 2 million individuals to assess whether tropical forests can be characterized by size-dependent life-history survival strategies. We found that species were classifiable into four ‘survival modes’ that explain life-history variation that shapes carbon cycling and the relative abundance within forests. Frequently collected functional traits, such as wood density, leaf mass per area and seed mass, were not generally predictive of the survival modes of species. Mean annual temperature and cumulative water deficit predicted the proportion of biomass of survival modes, indicating important links between evolutionary strategies, climate and carbon cycling. The application of survival modes in demographic simulations predicted biomass change across forest sites. Our results reveal globally identifiable size-dependent survival strategies that differ across diverse systems in a consistent way. The abundance of survival modes and interaction with climate ultimately determine forest structure, carbon storage in biomass and future forest trajectories.

The effects of forest fires on tree recruitment dynamics in tropical forests is important for predicting forest dynamics and ecosystem function in Southeast Asia. To our knowledge, no studies have examined the effects of fire intensity on community-level recruitment patterns in tropical forests due to the rarity of long-term observation datasets in fire-impacted tropical forests and the difficulty of quantifying fire intensity. We addressed two questions: (1) is tree recruitment among species affected by fire intensity? and if so, (2) are there specific plant functional traits associated with these responses? We used data from a long-term forest dynamics plot at the Huai Kha Khaeng (HKK) Wildlife Sanctuary in Thailand. The HKK plot occurs in a strongly seasonal tropical environment and has experienced several fires since its establishment in 1994. We found 46 tree species (52% of the 89 species analysed) showed evidence of reduced recruitment rates with increasing fire intensities during the most recent fire in 2005. Tree species in this flammable landscape have various leaf and wood functional traits associated with fire. Spatial and temporal variability in fire activity may lead to alterations in long-term taxonomic and functional composition of the forest due to selection on fire-related traits.

Summary Our understanding of the factors that shape reproductive phenology in Southeast Asian tropical forests comes exclusively from studies of everwet, general flowering forests. Therefore we examined the phenology of an evergreen dipterocarp forest in a climate exhibiting seasonal drought in Southeast Asia, with the aim of evaluating what evidence it brings to bear on the hypothesis that reproductive phenology in contemporary dipterocarp forests is the product of evolutionarily conserved responses to seasonal drought. We hypothesized that (i) dipterocarp forests that experience seasonal drought would exhibit annual reproduction and seasonality similar to that of forests with similar climates in other parts of the world, (ii) within species, flowering frequency would be climate-dependent, not a fixed trait, and (iii) climatic events that cue flowering in everwet forests would also cue flowering in seasonally dry forests. From 2001-2009, we monitored flowering and fruiting monthly for 1344 trees (>300 spp.) in a forest in Khao Chong, Thailand that experiences an annual 2-4 month dry season, assessing frequency, duration, seasonality, and synchrony of reproduction. We also examined phenological records for trees in three dipterocarp-dominated forests in everwet climates. Reproductive phenology of the Khao Chong forest was more similar to tropical forests with similar rainfall seasonality in other parts of the world than it was to dipterocarp-dominated forests in everwet regions of Southeast Asia. The forest exhibited annual reproduction, with peak flowering occurring at the end of the dry season, and peak fruiting occurring early in the wet season. Approximately half of all species and individuals reproduced annually, including several species that are known to exhibit “general flowering” in everwet climates. Short dry spells appeared to cue flowering for early flowering species, while later flowering species may have responded to either low precipitation or temperature, consistent with data from everwet forests. General flowering behaviour also showed significant phylogenetic signal. Synthesis. Our results support the hypothesis that the phenological behaviour of both seasonal and everwet dipterocarp forests may have initially evolved in response to a seasonal drought and that the general flowering phenomenon is a product of evolutionary conservatism. This article is protected by copyright. All rights reserved.

Maintaining tree diversity Negative interaction among plant species is known as conspecific negative density dependence (CNDD). This ecological pattern is thought to maintain higher species diversity in the tropics. LaManna et al. tested this hypothesis by comparing how tree species diversity changes with the intensity of local biotic interactions in tropical and temperate latitudes (see the Perspective by Comita). Stronger local specialized biotic interactions seem to prevent erosion of biodiversity in tropical forests, not only by limiting populations of common species, but also by strongly stabilizing populations of rare species, which tend to show higher CNDD in the tropics. Science , this issue p. 1389 ; see also p. 1328

We propose a new classification of rain forest plants into eight fruit syndromes, based on fruit morphology and other traits relevant to fruit‐feeding insects. This classification is compared with other systems based on plant morphology or traits relevant to vertebrate fruit dispersers. Our syndromes are based on fruits sampled from 1,192 plant species at three Forest Global Earth Observatory plots: Barro Colorado Island (Panama), Khao Chong (Thailand), and Wanang (Papua New Guinea). The three plots differed widely in fruit syndrome composition. Plant species with fleshy, indehiscent fruits containing multiple seeds were important at all three sites. However, in Panama, a high proportion of species had dry fruits, while in New Guinea and Thailand, species with fleshy drupes and thin mesocarps were dominant. Species with dry, winged seeds that do not develop as capsules were important in Thailand, reflecting the local importance of Dipterocarpaceae. These differences can also determine differences among frugivorous insect communities. Fruit syndromes and colors were phylogenetically flexible traits at the scale studied, as only three of the eight seed syndromes, and one of the 10 colors, showed significant phylogenetic clustering at either genus or family levels. Plant phylogeny was, however, the most important factor explaining differences in overall fruit syndrome composition among individual plant families or genera across the three study sites.

Insect seed predators are important agents of mortality for tropical trees, but little is known about the impact of these herbivores in rainforests. During 3 years at Khao Chong (KHC) in southern Thailand we reared 17,555 insects from 343.2 kg or 39,252 seeds/fruits representing 357 liana and tree species. A commented list of the 243 insect species identified is provided, with details about their host plants. We observed the following. (i) Approximately 43% of identified species can be considered pests. Most were seed eaters, particularly on dry fruits. (ii) Approximately 19% of parasitoid species (all Opiinae) for which we could determine whether their primary insect host was a pest or not (all Bactrocera spp. breeding in fruits) can be considered beneficials. (iii) The seeds/fruits of approximately 28% of the plant species in this forest were free of attack. Phyllanthaceae, Rubiaceae and Meliaceae were attacked relatively infrequently; in contrast, Annonaceae, Fabaceae, Sapindaceae and Myristicaceae were more heavily attacked. There was no apparent effect of plant phylogeny on rates of attack but heavily attacked tree species had larger basal area in the KHC plot than rarely attacked tree species. (iv) Insects reared from fleshy fruits were more likely to show relatively stable populations compared to insects reared from dry fruits, but this was not true of insects reared from dipterocarps, which appeared to have relatively stable populations throughout the study period. We tentatively conclude that insects feeding on seeds and fruits have little effect on observed levels of host abundance in this forest.

Advances in forest carbon mapping have the potential to greatly reduce uncertainties in the global carbon budget and to facilitate effective emissions mitigation strategies such as REDD+ (Reducing Emissions from Deforestation and Forest Degradation). Though broad-scale mapping is based primarily on remote sensing data, the accuracy of resulting forest carbon stock estimates depends critically on the quality of field measurements and calibration procedures. The mismatch in spatial scales between field inventory plots and larger pixels of current and planned remote sensing products for forest biomass mapping is of particular concern, as it has the potential to introduce errors, especially if forest biomass shows strong local spatial variation. Here, we used 30 large (8–50 ha) globally distributed permanent forest plots to quantify the spatial variability in aboveground biomass density (AGBD in Mg ha–1) at spatial scales ranging from 5 to 250 m (0.025–6.25 ha), and to evaluate the implications of this variability for calibrating remote sensing products using simulated remote sensing footprints. We found that local spatial variability in AGBD is large for standard plot sizes, averaging 46.3% for replicate 0.1 ha subplots within a single large plot, and 16.6% for 1 ha subplots. AGBD showed weak spatial autocorrelation at distances of 20–400 m, with autocorrelation higher in sites with higher topographic variability and statistically significant in half of the sites. We further show that when field calibration plots are smaller than the remote sensing pixels, the high local spatial variability in AGBD leads to a substantial "dilution" bias in calibration parameters, a bias that cannot be removed with standard statistical methods. Our results suggest that topography should be explicitly accounted for in future sampling strategies and that much care must be taken in designing calibration schemes if remote sensing of forest carbon is to achieve its promise.

Hülsmann and Hartig suggest that ecological mechanisms other than specialized natural enemies or intraspecific competition contribute to our estimates of conspecific negative density dependence (CNDD). To address their concern, we show that our results are not the result of a methodological artifact and present a null-model analysis that demonstrates that our original findings—(i) stronger CNDD at tropical relative to temperate latitudes and (ii) a latitudinal shift in the relationship between CNDD and species abundance—persist even after controlling for other processes that might influence spatial relationships between adults and recruits.

Aim Insects feeding on seeds and fruits represent interesting study systems, potentially able to lower the fitness of their host plants. In addition to true seed eaters, a suite of insects feed on the fleshy parts of fruits. We examined the likelihood of community convergence in whole insect assemblages attacking seeds/fruits in three tropical rainforests. Location Three ForestGEO permanent forest plots within different biogeographical regions: Barro Colorado Island (Panama), Khao Chong (Thailand) and Wanang (Papua New Guinea). Methods We surveyed 1,186 plant species and reared 1.1 ton of seeds/fruits that yielded 80,600 insects representing at least 1,678 species. We assigned seeds/fruits to predation syndromes on the basis of plant traits relevant to insects, seed/fruit appearance and mesocarp thickness. Results We observed large differences in insect faunal composition, species richness and guild structure between our three study sites. We hypothesize that the high species richness of insect feeding on seeds/fruits in Panama may result from a conjunction of low plant species richness and high availability of dry fruits. Insect assemblages were weakly influenced by seed predation syndromes, both at the local and regional scale, and the effect of host phylogeny varied also among sites. At the driest site (Panama), the probability of seeds of a plant species being attacked depended more on seed availability than on the measured seed traits of that plant species. However when seeds were attacked, plant traits shaping insect assemblages were difficult to identify and not related to seed availability. Main conclusions We observed only weak evidence of community convergence at the intercontinental scale among these assemblages. Our study suggests that seed eaters may be most commonly associated with dry fruits at relatively dry tropical sites where fleshy fruits may be less prevalent.

The important role of tropical forests in the global carbon cycle makes it imperative to assess changes in their carbon dynamics for accurate projections of future climate-vegetation feedbacks. Forest monitoring studies conducted over the past decades have found evidence for both increasing and decreasing growth rates of tropical forest trees. The limited duration of these studies restrained analyses to decadal scales, and it is still unclear whether growth changes occurred over longer time scales, as would be expected if CO2 -fertilization stimulated tree growth. Furthermore, studies have so far dealt with changes in biomass gain at forest-stand level, but insights into species-specific growth changes - that ultimately determine community-level responses - are lacking. Here, we analyse species-specific growth changes on a centennial scale, using growth data from tree-ring analysis for 13 tree species (~1300 trees), from three sites distributed across the tropics. We used an established (regional curve standardization) and a new (size-class isolation) growth-trend detection method and explicitly assessed the influence of biases on the trend detection. In addition, we assessed whether aggregated trends were present within and across study sites. We found evidence for decreasing growth rates over time for 8-10 species, whereas increases were noted for two species and one showed no trend. Additionally, we found evidence for weak aggregated growth decreases at the site in Thailand and when analysing all sites simultaneously. The observed growth reductions suggest deteriorating growth conditions, perhaps due to warming. However, other causes cannot be excluded, such as recovery from large-scale disturbances or changing forest dynamics. Our findings contrast growth patterns that would be expected if elevated CO2 would stimulate tree growth. These results suggest that commonly assumed growth increases of tropical forests may not occur, which could lead to erroneous predictions of carbon dynamics of tropical forest under climate change.

Some soils under tropical forests in western Thailand are derived from skarn complexes of hydrothermally metamorphosed granitic, calcareous and ultramafic rocks. We used data from six large, long-term forest ecological research plots to compare the soils derived from skarn with forest soils derived from granites and sedimentary parent materials elsewhere Thailand. The soils derived from skarn are Vermic Phaeozems and have deep, dark, worm-worked topsoils and bimodal particle size distributions of coarse sand and grit in clay or fine loam matrices. They are eutrophic with respect to both labile and non-labile forms of the mineral nutrients. The soils derived from other parent materials are mostly Acrisols. Analyses of variance for the cationic nutrients taken independently clearly distinguished the Phaeozems from the Acrisols. The two groups are also stoichiometrically distinct with respect to the main cationic nutrients, as depicted graphically by nutrient roses and as quantified as M+:TEB ratios. The cationic stoichiometric proportions also differentiated between the Acrisols on different plots and parent materials; and between the Phaeozems in our study and eutrophic soils in lowland forests elsewhere in the lowland tropics, with the Phaeozems having lower exchangeable Ca and Mg contents but higher exchangeable K. Subsoil cationic stoichiometric profiles appear to derive from parent materials, but those of the topsoil may be modified by selective biotic recycling. Nonetheless, inherited lithogenic stoichiometric ratios are still apparent in our topsoils. The forests on skarn in West Thailand are varied and overlap with those on Acrisols and other soils. This confirms earlier findings that climate and disturbance history have more influence than soils on the regional distribution of forest types in Thailand, although soils can be important at more local scales.

Global change is impacting forests worldwide, threatening biodiversity and ecosystem services including climate regulation. Understanding how forests respond is critical to forest conservation and climate protection. This review describes an international network of 59 long-term forest dynamics research sites (CTFS-ForestGEO) useful for characterizing forest responses to global change. Within very large plots (median size 25ha), all stems 1cm diameter are identified to species, mapped, and regularly recensused according to standardized protocols. CTFS-ForestGEO spans 25 degrees S-61 degrees N latitude, is generally representative of the range of bioclimatic, edaphic, and topographic conditions experienced by forests worldwide, and is the only forest monitoring network that applies a standardized protocol to each of the world's major forest biomes. Supplementary standardized measurements at subsets of the sites provide additional information on plants, animals, and ecosystem and environmental variables. CTFS-ForestGEO sites are experiencing multifaceted anthropogenic global change pressures including warming (average 0.61 degrees C), changes in precipitation (up to +/- 30% change), atmospheric deposition of nitrogen and sulfur compounds (up to 3.8g Nm(-2)yr(-1) and 3.1g Sm(-2)yr(-1)), and forest fragmentation in the surrounding landscape (up to 88% reduced tree cover within 5km). The broad suite of measurements made at CTFS-ForestGEO sites makes it possible to investigate the complex ways in which global change is impacting forest dynamics. Ongoing research across the CTFS-ForestGEO network is yielding insights into how and why the forests are changing, and continued monitoring will provide vital contributions to understanding worldwide forest diversity and dynamics in an era of global change.

Long-term surveys of entire communities of species are needed to measure fluctuations in natural populations and elucidate the mechanisms driving population dynamics and community assembly. We analysed changes in abundance of over 4000 tree species in 12 forests across the world over periods of 6-28 years. Abundance fluctuations in all forests are large and consistent with population dynamics models in which temporal environmental variance plays a central role. At some sites we identify clear environmental drivers, such as fire and drought, that could underlie these patterns, but at other sites there is a need for further research to identify drivers. In addition, cross-site comparisons showed that abundance fluctuations were smaller at species-rich sites, consistent with the idea that stable environmental conditions promote higher diversity. Much community ecology theory emphasises demographic variance and niche stabilisation; we encourage the development of theory in which temporal environmental variance plays a central role.

Long-distance seed dispersal (LDD) is considered a crucial determinant of tree distributions, but its effects depend on demographic processes that enable seeds to establish into adults and that remain poorly understood at large spatial scales. We estimated rates of seed arrival, germination, and survival and growth for a canopy tree species (Miliusa horsfieldii), in a landscape ranging from evergreen forest, where the species' abundance is high, to deciduous forest, where it is extremely low. We then used an individual-based model (IBM) to predict sapling establishment and to compare the relative importance of seed arrival and establishment in explaining the observed distribution of seedlings. Individuals in deciduous forest, far from the source population, experienced multiple benefits (e.g., increased germination rate and seedling survival and growth) from being in a habitat where conspecifics were almost absent. The net effect of these spatial differences in demographic processes was significantly higher estimated sapling establishment probabilities for seeds dispersed long distances into deciduous forest. Despite the high rate of establishment in this habitat, Miliusa is rare in the deciduous forest because the arrival of seeds at long distances from the source population is extremely low. Across the entire landscape, the spatial pattern of seed arrival is much more important than the spatial pattern of establishment for explaining observed seedling distributions. By using dynamic models to link demographic data to spatial patterns, we show that LDD plays a pivotal role in the distribution of this tree in its native habitat.

Many tropical tree species have population structures that exhibit strong recruitment failure. While the presence of adult trees indicates that appropriate regeneration conditions occurred in the past, it is often unclear why small individuals are absent. Knowing how, when and where these tree species regenerate provides insights into their life history characteristics. Based on tree age distributions inferences can be made on past forest dynamics and information is obtained that is important for forest management. We used tree-ring analyses to obtain tree ages and reconstruct >200 years of estimated establishment rates in a sparsely regenerating population of Afzelia xylocarpa (Fabaceae), a light-demanding and long-lived canopy tree species. We sampled all 85 Afzelia trees >5 cm diameter at breast height (dbh) in a 297-ha plot in a seasonal tropical forest in the Huai Kha Khaeng (HKK) Wildlife Sanctuary, western Thailand. The age distribution of the sampled Afzelia trees revealed two distinct recruitment peaks centred around 1850 and 1950. The presence of distinct age cohorts provides a strong indication of disturbance-mediated recruitment. Additionally we found three lines of evidence supporting this interpretation. (1) Similarly aged trees were spatially aggregated up to ~500 m, a scale larger than single tree-fall gaps. (2) High juvenile growth rates (5–10 mm dbh year-1) of extant small and large trees indicate that recruitment took place under open conditions. (3) A significant positive correlation between tree age and local canopy height indicates that trees recruited in low-canopy forest patches. Likely causes of these severe canopy disturbances include windstorms and ground fires, which are common in the region. In addition, successful establishment seems to be favoured by wetter climate conditions, as the estimated establishment rate was correlated to the Palmer Drought Severity Index (PDSI). Thus, the co-occurrence of canopy disturbance and favourable climatic conditions may provide a window of opportunity for Afzelia establishment. Our results indicate that forest patches with occurrence of large Afzelia trees have undergone high-severity canopy disturbance prior to establishment, suggesting that these disturbances have shaped forests at HKK. Tree-ring analyses provide a powerful tool to understanding tropical tree establishment patterns. Rare, high-severity canopy disturbances may play a key role in the regeneration of long-lived tropical canopy tree species with recruitment failure, potentially in interaction with climate variability to determine variation in establishment success over decades or centuries.

Climate change effects on growth rates of tropical trees may lead to alterations in carbon cycling of carbon-rich tropical forests. However, climate sensitivity of broad-leaved lowland tropical trees is poorly understood. Dendrochronology (tree-ring analysis) provides a powerful tool to study the relationship between tropical tree growth and annual climate variability. We aimed to establish climate-growth relationships for five annual-ring forming tree species, using ring-width data from 459 canopy and understory trees from a seasonal tropical forest in western Thailand. Based on 183/459 trees, chronologies with total lengths between 29 and 62 years were produced for four out of five species. Bootstrapped correlation analysis revealed that climate-growth responses were similar among these four species. Growth was significantly negatively correlated with current-year maximum and minimum temperatures, and positively correlated with dry-season precipitation levels. Negative correlations between growth and temperature may be attributed to a positive relationship between temperature and autotrophic respiration rates. The positive relationship between growth and dry-season precipitation levels likely reflects the strong water demand during leaf flush. Mixed-effect models yielded results that were consistent across species: a negative effect of current wet-season maximum temperatures on growth, but also additive positive effects of, for example, prior dry-season maximum temperatures. Our analyses showed that annual growth variability in tropical trees is determined by a combination of both temperature and precipitation variability. With rising temperature, the predominantly negative relationship between temperature and growth may imply decreasing growth rates of tropical trees as a result of global warming.

The relationship between species richness and ecosystem function, as measured by productivity or biomass, is of long-standing theoretical and practical interest in ecology. This is especially true for forests, which represent a majority of global biomass, productivity and biodiversity. Here, we conduct an analysis of relationships between tree species richness, biomass and productivity in 25 forest plots of area 8–50 ha from across the world. The data were collected using standardized protocols, obviating the need to correct for methodological differences that plague many studies on this topic. We found that at very small spatial grains (0.04 ha) species richness was generally positively related to productivity and biomass within plots, with a doubling of species richness corresponding to an average 48% increase in productivity and 53% increase in biomass. At larger spatial grains (0.25 ha, 1 ha), results were mixed, with negative relationships becoming more common. The results were qualitatively similar but much weaker when we controlled for stem density: at the 0.04 ha spatial grain, a doubling of species richness corresponded to a 5% increase in productivity and 7% increase in biomass. Productivity and biomass were themselves almost always positively related at all spatial grains. Synthesis. This is the first cross-site study of the effect of tree species richness on forest biomass and productivity that systematically varies spatial grain within a controlled methodology. The scale-dependent results are consistent with theoretical models in which sampling effects and niche complementarity dominate at small scales, while environmental gradients drive patterns at large scales. Our study shows that the relationship of tree species richness with biomass and productivity changes qualitatively when moving from scales typical of forest surveys (0.04 ha) to slightly larger scales (0.25 and 1 ha). This needs to be recognized in forest conservation policy and management.

The integration of ecology and evolutionary biology requires an understanding of the evolutionary lability in species' ecological niches. For tropical trees, specialization for particular soil resource and topographic conditions is an important part of the habitat niche, influencing the distributions of individual species and overall tree community structure at the local scale. However, little is known about how these habitat niches are related to the evolutionary history of species. We assessed the relationship between taxonomic rank and tree species' soil resource and topographic niches in eight large (24-50 ha) tropical forest dynamics plots. Niche overlap values, indicating the similarity of two species' distributions along soil or topographic axes, were calculated for all pairwise combinations of co-occurring tree species at each study site. Congeneric species pairs often showed greater niche overlap (i.e., more similar niches) than non-congeneric pairs along both soil and topographic axes, though significant effects were found for only five sites based on Mantel tests. No evidence for taxonomic effects was found at the family level. Our results indicate that local habitat niches of trees exhibit varying degrees of phylogenetic signal at different sites, which may have important ramifications for the phylogenetic structure of these communities.

Disturbances play an important role in forest dynamics across the globe. Researchers have mainly focused on the temporal context of disturbances, but have largely ignored the spatial patterns of tree recruitment they create. Geostatistical tools enable the analysis of spatial patterns and variability in tropical forest disturbance histories. Here, we examine the potential of combining dendroecological analysis and spatial statistics to reconstruct the disturbance history of a seasonal dry evergreen tropical forest plot at the Huai Kha Khaeng Wildlife Sanctuary (HKK), western Thailand. We used tree-ring-derived age estimates for 70 individuals of the shade-intolerant pioneer species Melia azederach (Meliaceae) and tree locations across a 316-ha study plot to identify the timing and spatial extent of past disturbances. Although the age distribution for Melia suggested that regeneration had been continuous over the past 60 yr, spatial analyses (mark correlation function and kriging) demonstrated the presence of three spatially discrete age cohorts. Two of these cohorts suggested a severe disturbance ~20 yr before present. A third cohort appears to have established ~50 years ago. Using historical records, we conclude that fire disturbance is the most likely disturbance factor affecting HKK. Nevertheless, we do not rule out other disturbance factors. The combined application of tree-ring analysis and spatial statistics as applied in this study could be readily applied to reconstruct disturbance histories in other tropical regions where tree species with annual growth rings are present.

The seasonally dry tropical forests of continental Southeast Asia occur in landscape-scale mosaics composed of several distinct forest types. The current understanding of the role of disturbances, such as fire and wind storms, in shaping these landscapes and the individual forest types within them, is limited by a lack of historical data. In this chapter we present the first dendroecological reconstruction of the historical dynamics of a mixed deciduous forest in western Thailand and compare it to an earlier study of the historical dynamics of an adjacent area of seasonal evergreen forest. The tree-ring data from the mixed deciduous forest demonstrate pulses of widespread gap recruitment since the 1850s for more than a dozen tree species. Recruitment and growth release data indicate complex patterns of disturbance both within and between forest types. However, there is strong evidence for extensive disturbances that synchronously impacted both the mixed deciduous and seasonal evergreen forests. Notably, several of the reconstructed landscape-scale disturbances are associated with intense regional drought events driven by ENSO variability in the Pacific Ocean. Interactions between the complex disturbance history of the landscape, which generate environmental conditions necessary for recruitment, and the wide range of reproductive phenologies, which generate the propagules for recruitment, provide a mechanism for coexistence for the many tree species that co-occur in these seasonal tropical forests.

Standardised transect counts of butterflies in old-growth rainforests in different biogeographical regions are lacking. Such data are needed to mitigate the influence of methodological and environmental factors within and between sites and, ultimately, to discriminate between long-term trends and short-term stochastic changes in abundance and community composition. 2. We compared butterfly assemblages using standardised Pollard Walks in the understory of closed-canopy lowland tropical rainforests across three biogeographi- cal regions: Barro Colorado Island (BCI), Panama; Khao Chong (KHC), Thailand; and Wanang (WAN), Papua New Guinea. 3. The length and duration of transects, their spatial autocorrelation, and number of surveys per year represented important methodological factors that strongly influ- enced estimates of butterfly abundance. Of these, the effect of spatial autocorrelation was most difficult to mitigate across study sites. 4. Butterfly abundance and faunal composition were best explained by air temper- ature, elevation, rainfall, wind velocity, and human disturbance at BCI and KHC. In the absence of weather data at WAN, duration of transects and number of forest gaps accounted for most of the explained variance, which was rather low in all cases (

Rising atmospheric carbon dioxide [CO₂] can accelerate tree growth by stimulating photosynthesis and increasing intrinsic water-use efficiency (iWUE). Little evidence exists, however, for the long-term growth and gas-exchange responses of mature trees in tropical forests to the combined effects of rising [CO₂] and other global changes such as warming. Using tree rings and stable isotopes of carbon and oxygen, we investigated long-term trends in the iWUE and stem growth (basal area increment, BAI) of three canopy tree species in a tropical monsoon forest in western Thailand (Chukrasia tabularis, Melia azedarach, and Toona ciliata). To do this, we modelled the contribution of ontogenetic effects (tree diameter or age) and calendar year to variation in iWUE, oxygen isotopes, and BAI using mixed-effects models. Although iWUE increased significantly with both tree diameter and calendar year in all species, BAI at a given tree diameter was lower in more recent years. For one species, C. tabularis, differences in crown dominance significantly influence stable isotopes and growth. Tree ring Δ¹⁸O increased with calendar year in all species, suggesting that increasing iWUE may have been driven by relatively greater reductions in stomatal conductance - leading to enrichment in Δ¹⁸O - than increases in photosynthetic capacity. Plausible explanations for the observed declines in growth include water stress resulting from rising temperatures and El Nino events, increased respiration, changes in allocation, or more likely, a combination of these factors.

Spatial and seasonal variation in soil respiration rates were investigated in a tropical dry forest in Thailand. The spatial variation was examined at 50 points within a 2-ha plot in the forest floor during the dry and wet seasons. The seasonal and diurnal variations in soil respiration were measured at 16 and 5 points, respectively. The mean soil respiration rate during the wet season was 1041 ± 542 mg CO2 m−2 h−1 (mean ± SD), which is about twice that during the dry season. Soil respiration rate was negatively correlated with soil water content during the wet season. A polynomial equation using seasonal data describes soil respiration and water content: soil respiration rate increased with soil water content, but started to drop when soil water content exceeded 21%. The diurnal variation in soil respiration rate during the wet season was positively correlated with soil temperature, whereas during the wet season it was not correlated with soil temperature. The diurnal variation in soil respiration rate during the dry season showed a midday depression. The estimation of soil carbon flux with polynomial equations should incorporate different functions for the wet and dry seasons in tropical dry forests.

BACKGROUND AND AIMS: Wood density is a key variable for understanding life history strategies in tropical trees. Differences in wood density and its radial variation were related to the shade-tolerance of six canopy tree species in seasonally dry tropical forest in Thailand. In addition, using tree ring measurements, the influence of tree size, age and annual increment on radial density gradients was analysed. METHODS: Wood density was determined from tree cores using X-ray densitometry. X-ray films were digitized and images were measured, resulting in a continuous density profile for each sample. Mixed models were then developed to analyse differences in average wood density and in radial gradients in density among the six tree species, as well as the effects of tree age, size and annual increment on radial increases in Melia azedarach. KEY RESULTS: Average wood density generally reflected differences in shade-tolerance, varying by nearly a factor of two. Radial gradients occurred in all species, ranging from an increase of (approx. 70%) in the shade-intolerant Melia azedarach to a decrease of approx. 13% in the shade-tolerant Neolitsea obtusifolia, but the slopes of radial gradients were generally unrelated to shade-tolerance. For Melia azedarach, radial increases were most-parsimoniously explained by log-transformed tree age and annual increment rather than by tree size. CONCLUSIONS: The results indicate that average wood density generally reflects differences in shade-tolerance in seasonally dry tropical forests; however, inferences based on wood density alone are potentially misleading for species with complex life histories. In addition, the findings suggest that a 'whole-tree' view of life history and biomechanics is important for understanding patterns of radial variation in wood density. Finally, accounting for wood density gradients is likely to improve the accuracy of estimates of stem biomass and carbon in tropical trees.

Summary 1Patterns of water availability are frequently implicated in local and regional tree species distributions. A major floristic and climatic transition from aseasonal to seasonal evergreen tropical forest is the Kangar–Pattani Line (KPL) in the Indo-Sundaic region of Southeast Asia. We hypothesize that differences in species’ drought tolerance will correspond with their distribution with respect to the KPL. Using a common garden study, we assess the role of differences in physiological drought tolerance traits to geographic distributions for 24 tropical tree species in relation to rainfall seasonality. 2Inherent differences in desiccation tolerance of seedlings were quantified as water potentials and relative water contents (RWCs) below which the plant could no longer support living tissue, and plant water relation parameters were measured using pressure–volume analysis. The relationships among these traits were examined using bivariate trait relationships and a principal components analysis (PCA). The physiological traits contributing most to lethal water potential and RWC were assessed using multiple regression analysis. 3Distribution-related differences in all desiccation tolerance and pressure–volume traits were detected both with and without phylogenetic correction. Widespread species that occurred in seasonally dry forests were able to maintain living tissue at more negative water potentials and lower RWCs than were species restricted to aseasonal forests. Likewise, widespread species demonstrated more negative water potentials at turgor loss, more negative saturated osmotic potential, lower symplastic water fraction and higher bulk modulus of elasticity values. Turgor loss point (TLP) and bulk modulus of elasticity were the best predictors of lethal water potential while symplastic water fraction and bulk modulus of elasticity were included as predictors of lethal RWC. 4Inherent differences in physiological traits contributing to drought tolerance are associated with differences in tropical tree species distributions in relation to rainfall seasonality. These results, combined with lack of support for hypothesized historical factors, strongly implicate climate as a determinant of tree species distributions around the KPL.

We used tree-rings to reconstruct long-term patterns of suppression, release and growth among five sympatric canopy tree species representing the full range of shade tolerance in a seasonal tropical forest in western Thailand. We expected that the frequency and duration of suppression and release events would be positively correlated with shade tolerance. All five species showed evidence of major and moderate growth releases. As expected, Melia azederach, an extreme heliophile, had the fewest releases. However, among the other species the number of major releases was consistent across the range of shade tolerance. The most significant difference among the species was the number of moderate releases recorded. There was a general positive correlation between the number of moderate releases and shade tolerance; however, Chukrasia tabularis, a relatively shade-intolerant species, had an anomalously high number of moderate releases. The study species also showed considerable variation in canopy accession strategies. The least common canopy accession strategy was establishment in the understorey and growth into the canopy in the absence of any gaps. However, with the exception of Melia, all four study species had one or more individuals that successfully reached the canopy using each of the four canopy accession strategies. These results highlight the importance of periodic or episodic bouts of gap formation on canopy tree recruitment and the utility of tree-rings for reconstructing long-term growth patterns in tropical trees.

Tropical forests vary substantially in the densities of trees of different sizes and thus in above-ground biomass and carbon stores. However, these tree size distributions show fundamental similarities suggestive of underlying general principles. The theory of metabolic ecology predicts that tree abundances will scale as the -2 power of diameter. Demographic equilibrium theory explains tree abundances in terms of the scaling of growth and mortality. We use demographic equilibrium theory to derive analytic predictions for tree size distributions corresponding to different growth and mortality functions. We test both sets of predictions using data from 14 large-scale tropical forest plots encompassing censuses of 473 ha and > 2 million trees. The data are uniformly inconsistent with the predictions of metabolic ecology. In most forests, size distributions are much closer to the predictions of demographic equilibrium, and thus, intersite variation in size distributions is explained partly by intersite variation in growth and mortality.

Disturbances influence forest dynamics across a range of spatial and temporal scales. In tropical forests most studies have focused on disturbances occurring at small spatial and temporal scales (i.e., gap dynamics). This is primarily due to the difficulty of reconstructing long-term disturbance histories of forests in which most tree species lack annual growth rings. Consequently, the role of past disturbances in tropical forests is poorly understood. We used a combination of direct and indirect methods to reconstruct the his- torical disturbance regime and stand development patterns in mature and regenerating seasonal dry evergreen forest (SDEF) in the Huai Kha Khaeng Wildlife Sanctuary in western Thailand. Direct estimates of long-term establishment and growth patterns were obtained from 12 tree species that form annual growth rings as a consequence of the region's strong intra-annual rainfall seasonality. Indirect estimates of establishment patterns were obtained from analyses of stand structure and individual tree architecture and application of age- estimation models to 10 dominant canopy-tree species using demographic data from a large- scale, permanent forest-dynamics plot. The combination of direct and indirect methodologies revealed a complex disturbance history in the seasonal evergreen forest over the past 250 years. In the mid-1800s, 200-300 ha of forest were destroyed by a catastrophic disturbance, which led to the synchronous es- tablishment of many of the trees that presently dominate the forest canopy. Since then wide- spread disturbances of variable intensity have occurred at least three times (1910s, 1940s, and 1960s). These disturbances created discrete temporal pulses of establishment in small to large gaps in the forest matrix across several square kilometers. Background mortality and gap formation were evident in every decade since 1790, but these varied in intensity and frequency. The SDEF retains a distinct structural and floristic legacy from the catastrophic dis- turbance of the mid-1800s. The single-age cohort that established after the disturbance has developed a complex three-dimensional structure as a consequence of differences in in- terspecific growth patterns of the canopy-tree species and subsequent disturbances of mod- erate and low intensity. While no single methodological approach provided a complete picture of the disturbance history and stand development patterns of the seasonal evergreen forest, taken together they offered new insights into the long-term dynamics of a primary tropical forest. In particular, the study highlighted the role of disturbance at multiple spatial and temporal scales and varying intensities in determining the structure and composition of a complex, species-rich tropical forest and raises important questions about the role of rare, catastrophic events on tropical forest dynamics.

Population structure and spatial patterns were examined for four species of canopy dipterocarps (Anisoptera costata, Dipterocarpus alatus, Hopea odorata, Vatica cinerea) in a 50 ha plot in seasonal dry evergreen forest at the Huai Kha Khaeng Wildlife Sanctuary in western Thailand. Spatial dispersion was assessed with Morisita’s index for quadrat sizes ranging from 61 m2 to 25 ha; spatial attraction and repulsion between species and size classes were measured with Iwao’s index. Only Vatica had a negative exponential diameter distribution suggesting continuous recruitment. The other species had either normal (Hopea) or irregular diameter distributions with peaks in the large size classes (Anisoptera, Dipterocarpus). All four species were significantly clumped at most quadrat sizes. At the local scale, saplings and poles of Hopea and Anisoptera were negatively associated with adults at quadrat sizes

Dependency on topographical habitat was examined for Lauraceae tree species in a lower montane forest using a large-scale research plot established at Doi Inthanon National Park, northern Thailand. Twenty species of 10 genera of Lauraceae were recorded in a 7.5-ha part of the plot; Lauraceae accounted for 18% of the total basal area. Lauraceae was the most species-rich and most abundant family in the plot. In a cluster analysis based on the matrix of spatial associations between species, two clusters were recognized. Members of one cluster seemed to associate with lower-elevation habitats, and members of the other associated with habitats on ridges. By subdividing the study plot into 20 m × 20 m squares, a discriminant analysis could be applied to the presence–absence data for the 17 species that had sufficient population density. The predictor variables used were the relative elevation, slope inclination, slope direction (transformed to deviation from SSW) and slope convexity for each of the squares. The discriminant models were tested statistically by applying the random shift technique. The models were significant for 11 of the species (65% of the species examined) and were associated with the topographical condition of the habitat. Stepwise selection of the predictor variables for these 11 species revealed that relative elevation and slope convexity were the most important factors for predicting the presence or absence of the Lauraceae species. Both these variables were considered to indicate the hydrological condition of the habitat.

Overhunting in tropical forests reduces populations of vertebrate seed dispersers. If reduced seed dispersal has a negative impact on tree population viability, overhunting could lead to altered forest structure and dynamics, including decreased biodiversity. However, empirical data showing decreased animal-dispersed tree abundance in overhunted forests contradict demographic models which predict minimal sensitivity of tree population growth rate to early life stages. One resolution to this discrepancy is that seed dispersal determines spatial aggregation, which could have demographic consequences for all life stages. We tested the impact of dispersal loss on population viability of a tropical tree species, Miliusa horsfieldii, currently dispersed by an intact community of large mammals in a Thai forest. We evaluated the effect of spatial aggregation for all tree life stages, from seeds to adult trees, and constructed simulation models to compare population viability with and without animal-mediated seed dispersal. In simulated populations, disperser loss increased spatial aggregation by fourfold, leading to increased negative density dependence across the life cycle and a 10-fold increase in the probability of extinction. Given that the majority of tree species in tropical forests are animal-dispersed, overhunting will potentially result in forests that are fundamentally different from those existing now.

The high species richness of tropical forests has long been recognized, yet there remains substantial uncertainty regarding the actual number of tropical tree species. Using a pantropical tree inventory database from closed canopy forests, consisting of 657,630 trees belonging to 11,371 species, we use a fitted value of Fisher's alpha and an approximate pantropical stem total to estimate the minimum number of tropical forest tree species to fall between ∼40,000 and ∼53,000, i.e. at the high end of previous estimates. Contrary to common assumption, the Indo-Pacific region was found to be as species-rich as the Neotropics, with both regions having a minimum of ∼19,000-25,000 tree species. Continental Africa is relatively depauperate with a minimum of ∼4,500-6,000 tree species. Very few species are shared among the African, American, and the Indo-Pacific regions. We provide a methodological framework for estimating species richness in trees that may help refine species richness estimates of tree-dependent taxa., 0, SCOPUS: ar.j, info:eu-repo/semantics/published

Forests are major components of the global carbon cycle, providing substantial feedback to atmospheric greenhouse gas concentrations. Our ability to understand and predict changes in the forest carbon cycle—particularly net primary productivity and carbon storage— increasingly relies on models that represent biological processes across several scales of biological organization, from tree leaves to forest stands. Yet, despite advances in our understanding of productivity at the scales of leaves and stands, no consensus exists about the nature of productivity at the scale of the individual tree, in part because we lack a broad empirical assessment of whether rates of absolute treemass growth (and thus carbon accumulation) decrease, remain constant, or increase as trees increase in size and age. Here we present a global analysis of 403 tropical and temperate tree species, showing that for most species mass growth rate increases continuously with tree size. Thus, large, old trees do not act simply as senescent carbon reservoirs but actively fix large amounts of carbon compared to smaller trees; at the extreme, a single big tree can add the same amount of carbon to the forest within a year as is contained in an entire mid-sized tree. The apparent paradoxes of individual tree growth increasing with tree size despite declining leaf-level and stand-level productivity can be explained, respectively, by increases in a tree’s total leaf area that outpace declines in productivity per unit of leaf area and, among other factors, age-related reductions in population density. Our results resolve conflicting assumptions about the nature of tree growth,inform efforts to undertand and model forest carbon dynamics, and have additional implications for theories of resource allocation and plant senescence. Fil: Blundo, Cecilia Mabel. Universidad Nacional de Tucumán. Facultad de Ciencias Naturales e Instituto Miguel Lillo. Instituto de Ecología Regional; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Grau, Hector Ricardo. Universidad Nacional de Tucumán. Facultad de Ciencias Naturales e Instituto Miguel Lillo. Instituto de Ecología Regional; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Malizia, Agustina. Universidad Nacional de Tucumán. Facultad de Ciencias Naturales e Instituto Miguel Lillo. Instituto de Ecología Regional; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Malizia, L. R.. Universidad Nacional de Jujuy. Facultad de Ciencias Agrarias; Argentina

While the importance of local-scale habitat niches in shaping tree species turnover along environmental gradients in tropical forests is well appreciated, relatively little is known about the influence of phylogenetic signal in species' habitat niches in shaping local community structure. We used detailed maps of the soil resource and topographic variation within eight 24-50 ha tropical forest plots combined with species phylogenies created from the APG III phylogeny to examine how phylogenetic beta diversity (indicating the degree of phylogenetic similarity of two communities) was related to environmental gradients within tropical tree communities. Using distance-based redundancy analysis we found that phylogenetic beta diversity, expressed as either nearest neighbor distance or mean pairwise distance, was significantly related to both soil and topographic variation in all study sites. In general, more phylogenetic beta diversity within a forest plot was explained by environmental variables this was expressed as nearest neighbor distance versus mean pairwise distance (3.0-10.3 % and 0.4-8.8 % of variation explained among plots, respectively), and more variation was explained by soil resource variables than topographic variables using either phylogenetic beta diversity metric. We also found that patterns of phylogenetic beta diversity expressed as nearest neighbor distance were consistent with previously observed patterns of niche similarity among congeneric species pairs in these plots. These results indicate the importance of phylogenetic signal in local habitat niches in shaping the phylogenetic structure of tropical tree communities, especially at the level of close phylogenetic neighbors, where similarity in habitat niches is most strongly preserved.