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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.

AimGlobal forests and their structural and functional features are shaped by many mechanisms that impact tree vital rates. Although many studies have tried to quantify how specific mechanisms influence vital rates, their relative importance among forests remains unclear. We aimed to assess the patterns of variation in vital rates among species and in space and time across forests to understand and provide a baseline for expectations of the relative importance of the different mechanisms in different contexts.Location21 forest plots worldwide.Time period1981-2021Major taxa studiedWoody plantsMethodsWe developed a conceptual and statistical framework (variance partitioning of multilevel models) that attributes the variability in growth, mortality, and recruitment to variation in species, space, and time, and their interactions, which we refer to asorganising principles(OPs). We applied it to data from 21 forest plots covering more than 2.9 million trees of approximately 6,500 species.ResultsDifferences among species, thespeciesOP, were a major source of variability in tree vital rates, explaining 28-33% of demographic variance alone, and in interaction withspace14-17%, totalling 40-43%. Models with small spatial grain sizes (quadrats at 5 × 5 m) retained most of the spatial OP, but a large proportion of variance remained unexplained (31-55%). The average variability among species declined with species richness across forests, indicating that diverse forests featured smaller interspecific differences in vital rates.Main conclusionsDecomposing variance in vital rates into the proposed OPs showed that taxonomy is crucial to predictions and understanding of tree demography. Our framework has a high potential for identifying the structuring mechanisms of global forest dynamics as it highlights the most promising avenues for future research both in terms of understanding the relative contributions of mechanisms to forest demography and diversity and for improving projections of forest ecosystems.

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.

Les forêts tropicales de la République démocratique du Congo sont menacées par la déforestation et la dégradation forestière, qui affectent la structure et la diversité arborée. Cette étude a pour objectif de quantifier les pertes de biomasse et de diversité liées à la déforestation et à la dégradation, en se concentrant sur la réserve de biosphère de Yangambi. Quatre parcelles de 50 m x 50 m ont été inventoriées dans chacun des huit types d’utilisation des terres identifiés : la forêt mature mixte, la forêt mature à Gilbertiodendron dewevrei, les champs des cultures vivrières, les jachères (5 à 10 ans), les forêts secondaires (20 à 40 ans), les plantations de cacaoyers, les plantations de caféiers et les plantations de palmiers à huile. Les stocks de biomasse dans les forêts matures s’élèvent à environ 400 et 380 Mg/ha respectivement pour la forêt à G. dewevrei et la forêt mixte. Les résultats montrent que la forêt mature perd 50, 70 et 90 % de sa biomasse aérienne quand elle est convertie en plantations de cacaoyers, caféiers et palmiers, respectivement, et la quasi-totalité (99 %) lorsqu’elle est convertie en champs de cultures vivrières ; mais quand le champ est abandonné, la biomasse se reconstitue progressivement à 7,5 % après 5 à 10 ans et à 38,8 % après 20 à 40 ans. La diversité est aussi impactée par la conversion, et, bien que la richesse spécifique semble se reconstituer plus rapidement que la biomasse, la composition est fondamentalement modifiée. Ces résultats fournissent des informations essentielles pour quantifier les impacts des solutions « naturelles » pour lutter contre le changement climatique : la protection des forêts matures, l’amélioration de la gestion forestière et le reboisement ; même si l’approche doit être étendue à plus grande échelle.

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.

The responses of tropical forests to environmental change are critical uncertainties in predicting the future impacts of climate change. The positive phase of the 2015–2016 El Niño Southern Oscillation resulted in unprecedented heat and low precipitation in the tropics with substantial impacts on the global carbon cycle. The role of African tropical forests is uncertain as their responses to short-term drought and temperature anomalies have yet to be determined using on-the-ground measurements. African tropical forests may be particularly sensitive because they exist in relatively dry conditions compared with Amazonian or Asian forests, or they may be more resistant because of an abundance of drought-adapted species. Here, we report responses of structurally intact old-growth lowland tropical forests inventoried within the African Tropical Rainforest Observatory Network (AfriTRON). We use 100 long-term inventory plots from six countries each measured at least twice prior to and once following the 2015–2016 El Niño event. These plots experienced the highest temperatures and driest conditions on record. The record temperature did not significantly reduce carbon gains from tree growth or significantly increase carbon losses from tree mortality, but the record drought did significantly decrease net carbon uptake. Overall, the long-term biomass increase of these forests was reduced due to the El Niño event, but these plots remained a live biomass carbon sink (0.51 ± 0.40 Mg C ha −1 y −1 ) despite extreme environmental conditions. Our analyses, while limited to African tropical forests, suggest they may be more resistant to climatic extremes than Amazonian and Asian forests., info:eu-repo/semantics/published

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.

Patterns of structural change associated with monodominant tropical forest complexes have remained enigmatic for decades. Here, we extend previous efforts in presenting a longitudinal, local-scale analysis of forest dynamics in central Africa. Using four 10-ha census plots measured across three time periods (959,312 stems ≥1 cm DBH), we analyzed changes in a number of biometrical attributes for four distinct forest types capturing the developmental gradient from mixed species forest to Gilbertiodendron dewevrei-dominated forest. We modeled above-ground biomass (AGB), basal area (BA), and stem density across all species, and diameter at breast height (DBH), recruitment, and mortality for Gilbertiodendron dewevrei. We hypothesized that trends in these attributes are consistent with a slow spread of Gilbertiodendron dewevrei into adjacent mixed species forest. We identified statistically significant increases in AGB and BA across sites and positive, though nonsignificant, increases in AGB and BA for most forest types. DBH and relative recruitment increased significantly for Gilbertiodendron dewevrei stems, while relative mortality did not. When looking from mixed species to transitional to monodominant forest types, we found a statistically significant pattern of developmental aggradation and net expansion of monodominant forest. We do not attribute this to atmospheric forcing but to a combination of (a) landscape-scale recovery or response to widespread disturbance (primarily historical fires), (b) Gilbertiodendron dewevrei’s ectomycorrhizal association, and (c) Gilbertiodendron dewevrei’s exceptional stress tolerance traits.

Pericopsis elata (a.k.a. African teak) is one of the most valuable timber species in Central Africa. Like other shade intolerant tropical tree species, P. elata could play a vital role in economic development, and ecological sustainability, but regenerates poorly following selective logging. Now endangered, there is a critical need for sustainable silvicultural systems to restore this once prominent timber species. To assess management options for P. elata we analyzed growth performance and survival in primary and secondary forest plots under burning and weeding treatments in Yoko Forest Reserve, Ubundu Democratic Republic of Congo. We transplanted nursery-grown seedlings of P. elata to experimental gaps and followed their growth and survival for 1 year. Seedlings in large canopy gaps 50 × 50 m were taller (mean difference; P = 0.006) and more likely to survive (mean difference; P

Structurally intact tropical forests sequestered about half of the global terrestrial carbon uptake over the 1990s and early 2000s, removing about 15 per cent of anthropogenic carbon dioxide emissions1,2,3. Climate-driven vegetation models typically predict that this tropical forest ‘carbon sink’ will continue for decades4,5. Here we assess trends in the carbon sink using 244 structurally intact African tropical forests spanning 11 countries, compare them with 321 published plots from Amazonia and investigate the underlying drivers of the trends. The carbon sink in live aboveground biomass in intact African tropical forests has been stable for the three decades to 2015, at 0.66 tonnes of carbon per hectare per year (95 per cent confidence interval 0.53–0.79), in contrast to the long-term decline in Amazonian forests6. Therefore the carbon sink responses of Earth’s two largest expanses of tropical forest have diverged. The difference is largely driven by carbon losses from tree mortality, with no detectable multi-decadal trend in Africa and a long-term increase in Amazonia. Both continents show increasing tree growth, consistent with the expected net effect of rising atmospheric carbon dioxide and air temperature7,8,9. Despite the past stability of the African carbon sink, our most intensively monitored plots suggest a post-2010 increase in carbon losses, delayed compared to Amazonia, indicating asynchronous carbon sink saturation on the two continents. A statistical model including carbon dioxide, temperature, drought and forest dynamics accounts for the observed trends and indicates a long-term future decline in the African sink, whereas the Amazonian sink continues to weaken rapidly. Overall, the uptake of carbon into Earth’s intact tropical forests peaked in the 1990s. Given that the global terrestrial carbon sink is increasing in size, independent observations indicating greater recent carbon uptake into the Northern Hemisphere landmass10 reinforce our conclusion that the intact tropical forest carbon sink has already peaked. This saturation and ongoing decline of the tropical forest carbon sink has consequences for policies intended to stabilize Earth’s climate.

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.

Quantifying carbon dynamics in forests is critical for understanding their role in long-term climate regulation 1–4 .Yet little is known about tree longevity in tropical forests 3,5–8 ,a factor that is vital for estimating carbon persistence 3,4 .Here we calculate mean carbon age (the period that carbon is fixed in trees 7 ) in different strata of African tropical forests using (1) growth-ring records with a unique timestamp accurately demarcating 66 years of growth in one site and (2) measurements of diameter increments from the African Tropical Rainforest Observation Network (23 sites). We find that in spite of their much smaller size, in understory trees mean carbon age (74 years) is greater than in sub-canopy (54 years) and canopy (57 years) trees and similar to carbon age in emergent trees (66 years). The remarkable carbon longevity in the understory results from slow and aperiodic growth as an adaptation to limited resource availability 9–11 .Our analysis also reveals that while the understory represents a small share (11%) of the carbon stock 12,13 ,it contributes disproportionally to the forest carbon sink (20%). We conclude that accounting for the diversity of carbon age and carbon sequestration among different forest strata is critical for effective conservation management 14–16 and for accurate modelling of carbon cycling 4 .© 2019, The Author(s), under exclusive licence to Springer Nature Limited., SCOPUS: le.j, info:eu-repo/semantics/published

Tropical forests are global centres of biodiversity and carbon storage. Many tropical countries aspire to protect forest to fulfil biodiversity and climate mitigation policy targets, but the conservation strategies needed to achieve these two functions depend critically on the tropical forest tree diversity-carbon storage relationship. Assessing this relationship is challenging due to the scarcity of inventories where carbon stocks in aboveground biomass and species identifications have been simultaneously and robustly quantified. Here, we compile a unique pan-Tropical dataset of 360 plots located in structurally intact old-growth closed-canopy forest, surveyed using standardised methods, allowing a multi-scale evaluation of diversity-carbon relationships in tropical forests. Diversity-carbon relationships among all plots at 1 ha scale across the tropics are absent, and within continents are either weak (Asia) or absent (Amazonia, Africa). A weak positive relationship is detectable within 1 ha plots, indicating that diversity effects in tropical forests may be scale dependent. The absence of clear diversity-carbon relationships at scales relevant to conservation planning means that carbon-centred conservation strategies will inevitably miss many high diversity ecosystems. As tropical forests can have any combination of tree diversity and carbon stocks both require explicit consideration when optimising policies to manage tropical carbon and biodiversity., 0, SCOPUS: ar.j, info:eu-repo/semantics/published

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.

Mbau forest covers much of the Congo, and shifts in its composition could have a large impact on the African tropics. The Ituri forest in east Congo is near a boundary between the monodominant mbau type and non-mbau mixed forest, and two 20-ha censuses of trees ≥ 1 cm diameter were carried out over 12 y to monitor forest change. Based on published diameter allometry, mbau forest had 535 Mg ha−1 biomass above ground and gained 1.1 Mg ha−1 y−1. Mixed forest had 399 Mg ha−1 and gained 3 Mg ha−1 y−1. The mbau tree (Gilbertiodendron dewevrei) increased its share of biomass from 4.1% to 4.4% in mixed forest; other common species also increased. Sapling density declined at both sites, likely because increased biomass meant shadier understorey, but the mbau tree increased in sapling density, suggesting it will become more important in the future. Tree mortality and growth rates were low relative to other tropical forests, especially in the mbau plots. Shifting toward G. dewevrei would represent a large gain in carbon in the mixed forest, but mbau is presently more important as a high-carbon stock: biomass lost during forest harvest could not recuperate for centuries due to slow community dynamics.

We test the hypotheses proposed by Gentry and Schnitzer that liana density and basal area in tropical forests vary negatively with mean annual precipitation (MAP) and positively with seasonality. Previous studies correlating liana abundance with these climatic variables have produced conflicting results, warranting a new analysis of drivers of liana abundance based on a different dataset. We compiled a pan-tropical dataset containing 28,953 lianas (Z2.5cmdiam.) from studies conducted at 13 Neotropical and 11 Paleotropical dry to wet lowland tropical forests. The ranges in MAP and dry season length (DSL) (number of months with mean rainfall o100mm) represented by these datasets were 860‐7250mm/yr and 0‐7mo, respectively. Pan-tropically, liana density and basal area decreased significantly with increasing annual rainfall and increased with increasing DSL, supporting the hypotheses of Gentry and Schnitzer. Our results suggest that much of the variation in liana density and basal area in the tropics can be accounted for by the relatively simple metrics of MAP and DSL.

Large tropical trees and a few dominant species were recently identified as the main structuring elements of tropical forests. However, such result did not translate yet into quantitative approaches which are essential to understand, predict and monitor forest functions and composition over large, often poorly accessible territories. Here we show that the above-ground biomass (AGB) of the whole forest can be predicted from a few large trees and that the relationship is proved strikingly stable in 175 1-ha plots investigated across 8 sites spanning Central Africa. We designed a generic model predicting AGB with an error of 14% when based on only 5% of the stems, which points to universality in forest structural properties. For the first time in Africa, we identified some dominant species that disproportionally contribute to forest AGB with 1.5% of recorded species accounting for over 50% of the stock of AGB. Consequently, focusing on large trees and dominant species provides precise information on the whole forest stand. This offers new perspectives for understanding the functioning of tropical forests and opens new doors for the development of innovative monitoring strategies., SCOPUS: ar.j, info:eu-repo/semantics/published

This chapter examines the pantropical patterns of liana abundance and species diversity and their correlates with climatic characteristics to gain insight into which processes are important for the distribution of tropical lianas. The analyses follow from the standard sampling protocol of liana diversity and abundance used by Alwyn Gentry in the 1980s and 1990s. The chapter examines how climatic conditions are associated with liana density and diversity, by analyzing old-growth continental forests at 850mm yr-1.To broaden the discussion of determinants of liana density and diversity beyond climatic factors, the authors comment on differences between temperate and tropical forests and between continental and island regions. The chapter focuses on sites in the Global Liana Database (GLD) from temperate forests in Australia, subtropical forest in Argentina, and an island in the Caribbean, and also reviews the available literature.

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.

Valuable timber tree species frequently show poor regeneration after selective logging in tropical forests. Small size of logging gaps, lack of soil disturbance, and limited seed availability have each been blamed for observed regeneration failures. We investigated seed germination and seedling performance using a split-plot factorial design involving light availability and litter removal for six Central African timber tree species, hypothesizing that canopy gaps and litter removal would improve seedling establishment, and that less shade-tolerant species would show stronger responses to both factors. Contrary to our expectations, significantly more germinants established on intact litter than on exposed mineral soil 3 mo after seeding. After 18 mo, seedling survival, height and diameter growth, leaf area, and rooting depth were all much higher in gap plots than in the understory for all species, with the exception of Gilbertiodendron dewevrei, a highly shade-tolerant species whose survival was higher in the understory. Leaf production was negatively influenced by litter removal in the least shade-tolerant species, Nauclea diderrichii, with weak or positive effects in other species. G. dewevrei, while displaying a low-light threshold for growth, exhibited a surprisingly high growth response to increasing light comparable to more shade-intolerant species, a response that may help explain its local competitive dominance in the region. Due to the rapid closure of small gaps, we suggest that shade-intolerant species such as N. diderrichii, Khaya anthotheca, and Entandrophragma utile might benefit from more intensive silvicultural practices that create larger canopy gaps.

The abundance of forest tree species may be locally limited by the inability of species to disperse to all sites suitable for germination and establishment. This phenomenon of "dispersal limitation" has been suggested to promote the maintenance of high species diversity in tropical forests by slowing down competitive exclusion. We present the first direct experimental evaluation of dispersal limitation in tropical forests, and of its importance relative to other factors affecting recruitment. Seed addition, litter removal and light availability effects on seedling establishment were evaluated in a full factorial experiment on two African mahogany species, Entandrophragma utile and Khaya anthotheca. Seedling recruitment after 18 months, was -22 times greater in seed addition treatments (with addition rates corresponding to the tail of seed shadows of large reproductive trees) compared to controls. This seed addition effect was an order of magnitude greater than either gap or litter removal effects, with similar results observed in both logged and primary forest locations. We conclude that dispersal limitation strongly limits seedling establishment for the two species studied. An important "applied corollary" of this result is that seed supplementation may substantially increase local abundance of valuable or rare species in disturbed or managed tropical forests.

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

We report above-ground biomass (AGB), basal area, stemdensity and wood mass density estimates from 260 sample plots (mean size: 1.2 ha) in intact closed-canopy tropical forests across 12 African countries. Mean AGB is 395.7 Mg dry mass ha-1 (95% CI: 14.3), substantially higher than Amazonian values, with the Congo Basin and contiguous forest region attaining AGB values (429 Mg ha-1) similar to those of Bornean forests, and significantly greater than East or West African forests. AGB therefore appears generally higher in palaeo- comparedwithneotropical forests.However, mean stem density is low(426±11 stems ha-1 greater than or equal to 100 mm diameter) compared with both Amazonian and Bornean forests (cf. approx. 600) and is the signature structural feature of African tropical forests. While spatial autocorrelation complicates analyses, AGB shows a positive relationship with rainfall in the driest nine months of the year, and an opposite association with the wettest three months of the year; a negative relationship with temperature; positive relationship with clay-rich soils; and negative relationshipswith C:Nratio (suggesting a positive soil phosphorus- AGB relationship), and soil fertility computed as the sum of base cations. The results indicate that AGB is mediated by both climate and soils, and suggest that the AGB of African closed-canopy tropical forests may be particularly sensitive to future precipitation and temperature changes. © 2013 The Authors., 0, SCOPUS: ar.j, info:eu-repo/semantics/published

The response of terrestrial vegetation to a globally changing environment is central to predictions of future levels of atmospheric carbon dioxide1, 2. The role of tropical forests is critical because they are carbon-dense and highly productive3, 4. Inventory plots across Amazonia show that old-growth forests have increased in carbon storage over recent decades5, 6, 7, but the response of one-third of the world's tropical forests in Africa8 is largely unknown owing to an absence of spatially extensive observation networks9, 10. Here we report data from a ten-country network of long-term monitoring plots in African tropical forests. We find that across 79 plots (163 ha) above-ground carbon storage in live trees increased by 0.63 Mg C ha-1 yr-1 between 1968 and 2007 (95% confidence interval (CI), 0.22¿0.94; mean interval, 1987¿96). Extrapolation to unmeasured forest components (live roots, small trees, necromass) and scaling to the continent implies a total increase in carbon storage in African tropical forest trees of 0.34 Pg C yr-1 (CI, 0.15¿0.43). These reported changes in carbon storage are similar to those reported for Amazonian forests per unit area6, 7, providing evidence that increasing carbon storage in old-growth forests is a pan-tropical phenomenon. Indeed, combining all standardized inventory data from this study and from tropical America and Asia5, 6, 11 together yields a comparable figure of 0.49 Mg C ha-1 yr-1 (n = 156; 562 ha; CI, 0.29¿0.66; mean interval, 1987¿97). This indicates a carbon sink of 1.3 Pg C yr-1 (CI, 0.8¿1.6) across all tropical forests during recent decades. Taxon-specific analyses of African inventory and other data12 suggest that widespread changes in resource availability, such as increasing atmospheric carbon dioxide concentrations, may be the cause of the increase in carbon stocks13, as some theory14 and models2, 10, 15 predict.

In Amazonian tropical forests, recent studies have reported increases in aboveground biomass and in primary productivity, as well as shifts in plant species composition favouring fast-growing species over slow-growing ones. This pervasive alteration of mature tropical forests was attributed to global environmental change, such as an increase in atmospheric CO2 concentration, nutrient deposition, temperature, drought frequency, and/or irradiance. We used standardized, repeated measurements of over 2 million trees in ten large (16–52 ha each) forest plots on three continents to evaluate the generality of these findings across tropical forests. Aboveground biomass increased at seven of our ten plots, significantly so at four plots, and showed a large decrease at a single plot. Carbon accumulation pooled across sites was significant (+0.24 MgC ha−1 y−1, 95% confidence intervals [0.07, 0.39] MgC ha−1 y−1), but lower than reported previously for Amazonia. At three sites for which we had data for multiple census intervals, we found no concerted increase in biomass gain, in conflict with the increased productivity hypothesis. Over all ten plots, the fastest-growing quartile of species gained biomass (+0.33 [0.09, 0.55] % y−1) compared with the tree community as a whole (+0.15 % y−1); however, this significant trend was due to a single plot. Biomass of slow-growing species increased significantly when calculated over all plots (+0.21 [0.02, 0.37] % y−1), and in half of our plots when calculated individually. Our results do not support the hypothesis that fast-growing species are consistently increasing in dominance in tropical tree communities. Instead, they suggest that our plots may be simultaneously recovering from past disturbances and affected by changes in resource availability. More long-term studies are necessary to clarify the contribution of global change to the functioning of tropical forests., Author Summary Recent studies have reported major changes in mature tropical forests, with increases in both forest biomass and net primary productivity, as well as shifts in plant species composition that favour fast-growing species over slow-growing ones. These pervasive alterations were attributed to global environmental change, and may result in dramatic shifts in the functioning of tropical forest ecosystems. We reassessed these findings using a dataset of large permanent forest plots on three continents. We found that tree biomass increased at seven of our ten plots, and showed a large decrease at a single plot. Overall, this increase was significant, albeit lower than reported previously for Amazonian forests. At three sites for which we had data for multiple census intervals, we found no concerted increase in biomass gain, in conflict with the increased productivity hypothesis. With the exception of one plot, slow-growing species gained more biomass than either fast-growing species or the tree community as a whole. Hence, our results do not support the hypothesis that fast-growing species are consistently increasing in dominance in tropical tree communities. Overall, our results suggest that our plots may be simultaneously recovering from past disturbances and affected by changes in resource availability., An analysis of changes in biomass and primary productivity across ten large tropical forest plots revealed that aboveground biomass increased significantly, fast-growing species increased significantly in dominance at a single site, and slow-growing species increased significantly at half of our sites.

Most ecological hypotheses about species coexistence hinge on species differences, but quantifying trait differences across species in diverse communities is often unfeasible. We examined the variation of demographic traits using a global tropical forest data set covering 4500 species in 10 large-scale tree inventories. With a hierarchical Bayesian approach, we quantified the distribution of mortality and growth rates of all tree species at each site. This allowed us to test the prediction that demographic differences facilitate species richness, as suggested by the theory that a tradeoff between high growth and high survival allows species to coexist. Contrary to the prediction, the most diverse forests had the least demographic variation. Although demographic differences may foster coexistence, they do not explain any of the 16-fold variation in tree species richness observed across the tropics.

The theory of metabolic ecology predicts specific relationships among tree stem diameter, biomass, height, growth and mortality. As demographic rates are important to estimates of carbon fluxes in forests, this theory might offer important insights into the global carbon budget, and deserves careful assessment. We assembled data from 10 oldgrowth tropical forests encompassing censuses of 367 ha and > 1.7 million trees to test the theory’s predictions. We also developed a set of alternative predictions that retained some assumptions of metabolic ecology while also considering how availability of a key limiting resource, light, changes with tree size. Our results show that there are no universal scaling relationships of growth or mortality with size among trees in tropical forests. Observed patterns were consistent with our alternative model in the one site where we had the data necessary to evaluate it, and were inconsistent with the predictions of metabolic ecology in all forests.

Forests are ubiquitous in the Democratic Republic of Congo; they touch the cultural and economic life of most of the population and have enormous global environmental significance. After years of conflicts and mismanagement, reconstruction is key to improving living conditions and consolidating peace. At the same time, better roads and trade bring risks—threatening forests and biodiversity by facilitating logging, land conversion, and the seizure of forest rights by vested interests. Anticipating these threats, in 2002, the transitional government started a Priority Reform Agenda. This report analyses the soundness of this Agenda, the progress achieved to date, and the priorities for the future. It emphasises the nature of forests as a public good; and the importance of the rule of law, transparency and public participation in managing natural resources. It highlights the multiplicity of claims on forests; calls for multipurpose participatory land-use planning; and emphasises the need to secure traditional user rights. Beyond the risks, the return of peace to the DRC also offers a unique opportunity to take a fresh look at the second-largest rainforest in the world, and to implement innovative strategies that give priority to the environment and to local people.

Tropical forests store 40–50 per cent of terrestrial vegetation carbon 1. However, spatial variations in aboveground live tree biomass carbon (AGC) stocks remain poorly understood, in particular in tropical montane forests 2. Owing to climatic and soil changes with increasing elevation 3, AGC stocks are lower in tropical montane forests compared with lowland forests 2. Here we assemble and analyse a dataset of structurally intact old-growth forests (AfriMont) spanning 44 montane sites in 12 African countries. We find that montane sites in the AfriMont plot network have a mean AGC stock of 149.4 megagrams of carbon per hectare (95% confidence interval 137.1–164.2), which is comparable to lowland forests in the African Tropical Rainforest Observation Network 4 and about 70 per cent and 32 per cent higher than averages from plot networks in montane 2,5,6 and lowland 7 forests in the Neotropics, respectively. Notably, our results are two-thirds higher than the Intergovernmental Panel on Climate Change default values for these forests in Africa 8. We find that the low stem density and high abundance of large trees of African lowland forests 4 is mirrored in the montane forests sampled. This carbon store is endangered: we estimate that 0.8 million hectares of old-growth African montane forest have been lost since 2000. We provide country-specific montane forest AGC stock estimates modelled from our plot network to help to guide forest conservation and reforestation interventions. Our findings highlight the need for conserving these biodiverse 9,10 and carbon-rich ecosystems.