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Funder: Agua Salud Project, Funder: U.S. Department of Energy; Id: http://dx.doi.org/10.13039/100000015, Funder: CAPES; Id: http://dx.doi.org/10.13039/501100002322, Data capturing multiple axes of tree size and shape, such as a tree's stem diameter, height and crown size, underpin a wide range of ecological research-from developing and testing theory on forest structure and dynamics, to estimating forest carbon stocks and their uncertainties, and integrating remote sensing imagery into forest monitoring programmes. However, these data can be surprisingly hard to come by, particularly for certain regions of the world and for specific taxonomic groups, posing a real barrier to progress in these fields. To overcome this challenge, we developed the Tallo database, a collection of 498,838 georeferenced and taxonomically standardized records of individual trees for which stem diameter, height and/or crown radius have been measured. These data were collected at 61,856 globally distributed sites, spanning all major forested and non-forested biomes. The majority of trees in the database are identified to species (88%), and collectively Tallo includes data for 5163 species distributed across 1453 genera and 187 plant families. The database is publicly archived under a CC-BY 4.0 licence and can be access from: https://doi.org/10.5281/zenodo.6637599. To demonstrate its value, here we present three case studies that highlight how the Tallo database can be used to address a range of theoretical and applied questions in ecology-from testing the predictions of metabolic scaling theory, to exploring the limits of tree allometric plasticity along environmental gradients and modelling global variation in maximum attainable tree height. In doing so, we provide a key resource for field ecologists, remote sensing researchers and the modelling community working together to better understand the role that trees play in regulating the terrestrial carbon cycle.

Temporal and spatial patterns in flowering phenology were assessed for eight tropical African tree species. Specifically, the frequency and seasonality of flowering at seven sites in central Africa were determined using field data, graphical analysis and circular statistics. Additionally, spatial variation in the timing of flowering across species range was investigated using herbarium data, analysing the relative influence of latitude, longitude and timing of the dry season with a Bayesian circular generalized linear model. Annual flowering was found for 20 out of the 25 populations studied. For 21 populations located at the north of the climatic hinge flowering was occurring during the dry season. The analysis of herbarium collections revealed a significant shift in the timing of flowering with latitude for E. suaveolens, and with the timing of the dry season for M. excelsa (and to a lesser extent L. alata), with the coexistence of two flowering peaks near the equator where the distribution of monthly rainfall is bimodal. For the other species, none of latitude, longitude or timing of the dry season had an effect on the timing of flowering. Our study highlights the need to identify the drivers of the flowering phenology of economically important African tree species., SCOPUS: ar.j, info:eu-repo/semantics/published

Caractériser la dynamique d’une forêt est essentiel pour la gestion forestière. Les houppiers des arbres forment un élément clé de cette dynamique ; mais, en forêt tropicale, les mesurer n’est pas simple. Cette étude teste l’utilisation d’images aériennes à haute résolution pour estimer la croissance diamétrique des arbres, en intégrant des mesures fines des houppiers détectés. Des ortho-images de 10 cm/pixel de résolution ont été obtenues à l’aide d’un drone à aile fixe sur une parcelle de 9 ha, installée dans la forêt de Yoko en République démocratique du Congo. Les inventaires menés sur les arbres de DHP ≥ 10 cm en 2008 et en 2016 ont permis d’avoir accès à différentes caractéristiques dendrométriques individuelles, dont le diamètre des arbres et leur tempérament, et de calculer des accroissements diamétriques. Des modèles linéaires mixtes ont été calibrés pour prédire l’accroissement de 163 arbres identifiés à la fois sur le terrain et sur les ortho-images en utilisant les variables quantifiées uniquement sur le terrain et/ou à partir de variables mesurées sur les ortho-images. Les images aériennes ont permis de détecter 23,4 % des arbres de DHP ≥ 10 cm inventoriés au sol, et représentant 75,1 % de la biomasse aérienne du peuplement. La probabilité de détection des arbres a varié en fonction de leur DHP : de 0,09 pour les arbres de DHP < 30 cm à 0,97 pour les arbres de DHP ≥ 60 cm. Les variables quantifiées par télédétection ajoutées aux variables de terrain ont permis d’améliorer significativement la prédiction de l’accroissement diamétrique. Les meilleurs modèles d’estimation des accroissements diamétriques contiennent notamment un terme caractérisant la dimension du houppier des arbres qui n’a pu être mesuré que par télédétection. Parmi les variables déterminées par télédétection, la superficie convexe du houppier est apparue la plus performante dans les modèles, et s’avère ainsi être la mesure la plus intéressante pour décrire la compétition entre les houppiers. Ces résultats ouvrent des perspectives pour construire de nouveaux outils d’acquisition de données au service de l’aménagement forestier.

Termite mounds have been poorly studied in tropical rainforest in contrast to those of savannahs where they create fertility hotspots and influence the spatial pattern of vegetation. An inventory of termite mounds and of 15 tree species with a diameter at breast height ≥ 10 cm was carried out in two 800-ha blocks, in two rainforest sites located in northern Congo. We used inhomogeneous and intertype K functions to study the spatial pattern of termite mounds and of tree species around mounds, respectively. We found that mounds in Loundoungou were over-dispersed within a radius of less than 70 m, while those in Mokabi were randomly spaced. Tree species within a 20-m radius around a mound were aggregated towards the mound, e.g. Entandrophragma cylindricum, randomly distributed, or even repulsed by the mound. The specific responses also differed in the two sites. These results suggest that (i) the mounds in Loundoungou were created by savannah termite species 3,000-2,000 years BP during the Late Holocene Rainforest Crisis and (ii) the mounds in Mokabi are characteristic of forest mounds. The impact of termite mounds on the spatial pattern of tree species is thus site-dependent, and these differences might be due to species seed dispersal strategies and to soil calcium concentrations.

Tree growth and mortality are two central processes in mixed and structurally complex moist tropical forests, yet accurate estimates of the variables needed to model them remain sparse and scattered. It is thus still difficult to predict forest evolution at a local scale and build reliable management plans. To help fill this gap, for 1–7 years we annually monitored 21,180 trees belonging to 42 species exploited for timber production in Central Africa. We made new species-specific estimates of diameter increments and mortality rates, and investigated how tree growth varied with tree size and logging history. We compared our results with the legal values of diameter increments, mortality rates, and minimum cutting diameters used to build forest management plans in Cameroon. Diameter increment was found to vary with tree size for most of the species studied. The relationships between diameter increment and tree size were mostly humpback-shaped. The trees with diameters close to or lower than the reference minimum cutting diameter generally grew faster than the average. We also found that tree growth could slow for 1–2 years after timber exploitation and was then spurred for at least 5 years. The tree growth response to logging was nevertheless species-specific. This study provides new estimates of tree diameter increments and mortality rates that could help make more accurate forest projections and draw up sustainable management plans in Africa.

When ordinating plots of tropical rain forests using stand-level structural attributes such as biomass, basal area and the number of trees in different size classes, two patterns often emerge: a gradient from poorly to highly stocked plots and high positive correlations between biomass, basal area and the number of large trees. These patterns are inherited from the demographics (growth, mortality and recruitment) and size allometry of trees and tend to obscure other patterns, such as site differences among plots, that would be more informative for inferring ecological processes. Using data from 133 rain forest plots at nine sites for which site differences are known, we aimed to filter out these patterns in forest structural attributes to unveil a hidden pattern. Using a null model framework, we generated the anticipated pattern inherited from individual allometric patterns. We then evaluated deviations between the data (observations) and predictions of the null model. Ordination of the deviations revealed site differences that were not evident in the ordination of observations. These sites differences could be related to different histories of large-scale forest disturbance. By filtering out patterns inherited from individuals, our model analysis provides more information on ecological processes., Frontiers in Ecology and Evolution, 9, ISSN:2296-701X

International audience; In tropical forests, the high proportion of trees showing irregularities at the stem base complicates forest monitoring. For example, in the presence of buttresses, the height of the point of measurement (HPOM) of the stem diameter (DPOM) is raised from 1.3 m, the standard breast height, up to a regular part of the stem. While DPOM is the most important predictor for tree aboveground biomass (AGB) estimates, the lack of harmonized HPOM for irregular trees in forest inventory increases the uncertainty in plot-level AGB stock and stock change estimates. In this study, we gathered an original non-destructive three-dimensional (3D) data set collected with terrestrial laser scanning and close range terrestrial photogrammetry tools in three sites in central Africa. For the 228 irregularly shaped stems sampled, we developed a set of taper models to harmonize HPOM by predicting the equivalent diameter at breast height (DBH′) from a DPOM measured at any height. We analyzed the effect of using DBH′ on tree-level and plot-level AGB estimates. To do so, we used destructive AGB data for 140 trees and forest inventory data from eight 1-ha plots in the Republic of Congo. Our results showed that our best simple taper model predicts DBH′ with a relative mean absolute error of 3.7% (R2 = 0.98) over a wide DPOM range of 17–249 cm. Based on destructive AGB data, we found that the AGB allometric model calibrated with harmonized HPOM data was more accurate than the conventional local and pantropical models. At the plot level, the comparison of AGB stock estimates with and without HPOM harmonization showed an increasing divergence with the increasing share of irregular stems (up to −15%). The harmonization procedure developed in this study could be implemented as a standard practice for AGB monitoring in tropical forests as no additional forest inventory measurements is required. This would probably lead to important revisions of the AGB stock estimates in regions having a large number of irregular tree stems and increase their carbon sink estimates. The growing use of three-dimensional (3D) data offers new opportunities to extend our approach and further develop general taper models in other tropical regions.

The sustainability of the polycyclic logging system in tropical forests has been increasingly questioned for a variety of reasons, and particularly in central Africa as commercial species, mostly light-demanding long-lived pioneer species, usually fail to recover a stable number of large trees after exploitation. Several factors are known to affect tropical tree demographic processes, like tree growth, survival and recruitment. Tree growth has particularly been showed to depend on ecological conditions, tree genetics, and competition with surrounding vegetation, as well as tree size or ontogeny. Yet, due to the paucity of available data, the importance of such factors is unclear and usually ignored when estimating future timber yields. To fill this gap, we chose to evaluate the variability in growth of one African long-lived pioneer and commercially very important species: Triplochiton scleroxylon K. Schum, gathering a broad dataset composed of tree ring data recorded in one site in Cameroon and periodic field inventory data recorded in seven sites across central Africa. In total, we analyzed 13,225 records of annual tree diameter increments recorded over 920 trees from seven sites in Cameroon, Republic of the Congo and Central African Republic. We evaluated (i) to what extent the average growth of trees that reach harvestable dimensions differs from population average and (ii) to what extent past perturbations influence the growth of remaining trees. We found the diameter growth of T. scleroxylon to be remarkably variable and this study provided an unprecedented quantification of the magnitude of some key growth determinants. In unlogged forests, the diameter increment of T. scleroxylon ranged between 0.40 cm year−1 in Southern Cameroon and 0.83 cm year−1 in South-Eastern Cameroon. The diameter increment was weakly related to tree size but increased twofold from unlogged to logged forests. Perturbation caused by logging stimulates growth of T. scleroxylon for at least 10–15 years. Finally, harvestable timber stock of large-sized T. scleroxylon was found to be constituted by trees that grew in average twice faster than trees of the entire extant population. As more and more inventory data become available, quantifying these effects could be replicated for other timber species and in other sites, to improve the accuracy of future timber resource estimates and improve forest management guidelines.

International audience; Tropical forests harbor the greatest terrestrial biodiversity and provide various ecosystem services. The increase of human activities on these forests, among which logging, makes the conservation of biodiversity and associated services strongly dependent on the sustainability of these activities. However the indicators commonly used to assess the impact of forest exploitation, namely species richness and biomass, provide a limited understanding of their sustainability. Here, we assessed the sustainability of common forest exploitation in the Guiana Shield studying the recovery of two ecosystem services i.e. carbon storage and wood stock, and an ecosystem function i.e. seed dispersal by animals. Specifically, we compared total and commercial biomass, as well as functional composition in seed size of animal-dispersed species in replicated forest plots before and 27 years after exploitation. Species richness is also studied to allow comparison. While species richness was not affected by forest exploitation, total and commercial biomass as well as seed size of animal-dispersed species decreased 27 years after exploitation, similarly to forests affected by hunting. These results show that ecosystem services and function likely did not recover even at the lowest intensity of forest exploitation studied, questioning the sustainability of the most common rotation-cycle duration applied in the tropics.

Tropical forests play a key role in regulating the terrestrial carbon cycle and climate change by storing a large amount of carbon. Yet, there is considerable uncertainty about the amount and spatial variation of aboveground biomass (AGB), especially in the relatively less studied African tropical forests. In this study, we explore the local-scale variation and determinants of plot-level AGB, between and within two types of forests, the Celtis and Manilkara forests, growing under the same climate but on different geological substrates in the northern Republic of Congo. In each forest site, all trees ≥10 cm diameter were censured in 36 × 1-ha plots and we measured tree height and crown size using a subsample of 18 × 1-ha of these plots. We developed height-diameter and crown-diameter allometric relationships and tested whether they differed between the two sites. For each 1-ha plot, we further estimated the AGB and calculated structural attributes (stem density and basal area), composition attributes (wood density) and architectural attributes (tree height and crown size), the latter being derived from site-specific allometric relationships. We found strong between-site differences in height-diameter and crown-diameter allometries. For a given diameter, trees were taller in the Celtis forest while they had larger crown in the Manilkara forest. Similar trends were found for the sixteen species present in both forest sites, suggesting an environmental control of tree allometry. Although there were some between-site differences in forest structure, composition and architecture, we did not detect any significant difference in mean AGB between the Celtis and the Manilkara forests. The AGB variation was related to the heterogeneous distribution of large trees, and influenced by basal area, height and crown dimensions, and to a lesser extent wood density. These forest attributes have strong practical implications on emerging remote-sensing technologies for carbon monitoring in tropical forests.

Africa is forecasted to experience large and rapid climate change1 and population growth2 during the twenty-first century, which threatens the world’s second largest rainforest. Protecting and sustainably managing these African forests requires an increased understanding of their compositional heterogeneity, the environmental drivers of forest composition and their vulnerability to ongoing changes. Here, using a very large dataset of 6 million trees in more than 180,000 field plots, we jointly model the distribution in abundance of the most dominant tree taxa in central Africa, and produce continuous maps of the floristic and functional composition of central African forests. Our results show that the uncertainty in taxon-specific distributions averages out at the community level, and reveal highly deterministic assemblages. We uncover contrasting floristic and functional compositions across climates, soil types and anthropogenic gradients, with functional convergence among types of forest that are floristically dissimilar. Combining these spatial predictions with scenarios of climatic and anthropogenic global change suggests a high vulnerability of the northern and southern forest margins, the Atlantic forests and most forests in the Democratic Republic of the Congo, where both climate and anthropogenic threats are expected to increase sharply by 2085. These results constitute key quantitative benchmarks for scientists and policymakers to shape transnational conservation and management strategies that aim to provide a sustainable future for central African forests. A large dataset of 6 million trees from 193 taxa is used to map the floristic and functional composition of central African forests and predict their vulnerability to climate change.

Aim:\ud \ud Tree crowns determine light interception, carbon and water exchange. Thus, understanding the factors causing tree crown allometry to vary at the tree and stand level matters greatly for the development of future vegetation modelling and for the calibration of remote sensing products. Nevertheless, we know little about large‐scale variation and determinants in tropical tree crown allometry. In this study, we explored the continental variation in scaling exponents of site‐specific crown allometry and assessed their relationships with environmental and stand‐level variables in the tropics.\ud \ud Location:\ud \ud Global tropics.\ud \ud Time period:\ud \ud Early 21st century.\ud \ud Major taxa :\ud \ud Woody plants.\ud \ud Methods:\ud \ud Using a dataset of 87,737 trees distributed among 245 forest and savanna sites across the tropics, we fitted site‐specific allometric relationships between crown dimensions (crown depth, diameter and volume) and stem diameter using power‐law models. Stand‐level and environmental drivers of crown allometric relationships were assessed at pantropical and continental scales.\ud \ud Results:\ud \ud The scaling exponents of allometric relationships between stem diameter and crown dimensions were higher in savannas than in forests. We identified that continental crown models were better than pantropical crown models and that continental differences in crown allometric relationships were driven by both stand‐level (wood density) and environmental (precipitation, cation exchange capacity and soil texture) variables for both tropical biomes. For a given diameter, forest trees from Asia and savanna trees from Australia had smaller crown dimensions than trees in Africa and America, with crown volumes for some Asian forest trees being smaller than those of trees in African forests.\ud \ud Main conclusions:\ud \ud Our results provide new insight into geographical variability, with large continental differences in tropical tree crown allometry that were driven by stand‐level and environmental variables. They have implications for the assessment of ecosystem function and for the monitoring of woody biomass by remote sensing techniques in the global tropics.

Forest biomass is an essential indicator for monitoring the Earth’s ecosystems and climate. It is a critical input to greenhouse gas accounting, estimation of carbon losses and forest degradation, assessment of renewable energy potential, and for developing climate change mitigation policies such as REDD+, among others. Wall-to-wall mapping of aboveground biomass (AGB) is now possible with satellite remote sensing (RS). However, RS methods require extant, up-to-date, reliable, representative and comparable in situ data for calibration and validation. Here, we present the Forest Observation System (FOS) initiative, an international cooperation to establish and maintain a global in situ forest biomass database. AGB and canopy height estimates with their associated uncertainties are derived at a 0.25 ha scale from field measurements made in permanent research plots across the world’s forests. All plot estimates are geolocated and have a size that allows for direct comparison with many RS measurements. The FOS offers the potential to improve the accuracy of RS-based biomass products while developing new synergies between the RS and ground-based ecosystem research communities., Measurement(s)above-ground biomass • organic materialTechnology Type(s)tree species census • measurement methodFactor Type(s)geographic location • tree speciesSample Characteristic - Environmentforest biomeSample Characteristic - LocationEarth (planet) Machine-accessible metadata file describing the reported data: 10.6084/m9.figshare.9850571

1.Architectural traits that determine the light captured in a given environment are an important aspect of the life‐history strategies of tropical tree species. In this study, we examined how interspecific variation in architectural traits is related to the functional traits of 45 coexisting tree species in central Africa. 2.At the tree level, we measured tree diameter, total height and crown dimensions for an average of 30 trees per species (range 14–72, total 968 trees) distributed over a large range of diameters (up to 162 cm). Using log‐log models, we fitted species‐specific allometric relationships between tree diameter, height and crown dimensions. At the species level, we derived architectural traits (height and crown dimensions) at 15 cm and maximum diameters from species‐specific allometries. The architectural traits were then related to functional traits, including light requirements, wood density, leaf habit, and dispersal mode. 3.Among the 45 coexisting tree species, we identified strong variations in height and crown allometries, along with architectural traits derived from these species‐specific allometries. There was a positive correlation among architectural traits, suggesting that large‐statured canopy species were taller and had larger and deeper crowns than small‐statured understory species at all ontogenic stages. The relationships between architectural and functional traits highlighted a continuum of species between the large‐statured canopy species and the small‐statured understory species. In this moist and seasonal forest, large‐statured canopy species tended to be light‐demanding, wind‐dispersed, deciduous and large contributors to forest biomass (high basal area), while small‐statured understory species tended to be shade‐tolerant, animal‐dispersed, evergreen and most abundant in terms of stem density. 4.Our results highlighted strong architectural differences among coexisting tropical tree species in central Africa. The relationships between architectural and functional traits provided insights into the life‐history strategy of tropical tree species.

Soil has been proposed as a driver explaining the development of monodominant forests in the tropics, for example, Gilbertiodendron dewevrei forests (GDF) in central Africa. The aim of this study was to compare the physical and chemical properties of soils under GDF with those under an adjacent mixed forest (AMF), while controlling for topography. To this end, we set up sixteen 0.25-ha plots according to forest type and topography (plateau vs. bottomland), in the Yoko forest reserve, Democratic Republic of Congo. In each plot, we measured litter thickness and collected a total of 80 soil samples at depths of 0–5, 5–10, 10–20, 20–40 and 120–150 cm, for standard physical and chemical analyses. When controlling for topography and soil texture, we found that most of the chemical properties of soils under GDF did not differ from those of soils under AMF, particularly acidity, cation concentration, total N and the C:N ratio. The litter layer was 2.3 times thicker under GDF than under AMF stands, and, for a given texture, soils under GDF had a slightly higher organic C concentration in the 0–5 cm soil layer. This study suggests that G. dewevrei stands modify organic matter dynamics, which may be important in maintaining its monodominance.

This study evaluates the characteristics of soil seed bank in two types of central African rainforests: Celtis forest on clay soils and Manilkara forest on sandy soils. In each study site, 30 samples were collected per soil layers (litter, 0–5 cm, 5–10 cm and 10–20 cm depth). The species diversity and abundance of the soil seed bank were estimated after soil samples were brought to germination. We globally observed 297 seedlings of 53 species for the Celtis forest and 222 seedlings of 39 species for the Manilkara forest. The total densities of germinated seeds were 330 seedlings m−2 and 247 seedlings m−2, respectively. Herbaceous species dominated with percentages of 41.0 and 45.3%, respectively in the Manilkara forest and the Celtis forest. Both forest types displayed a regeneration potential through the soil seed bank. However, this potential seems higher in the Celtis forest. Pioneer taxa were more abundant in the soil seed bank of the Celtis forest (13 woody pioneer species) than the Manilkara forest (9 woody pioneer species). The values of Sorensen similarity index between the standing tree vegetation and the soil seed bank in each site were relatively low: 11.0% for the Celtis forest and 8.8% for the Manilkara forest. But these similarity values were higher when only the pioneer species were considered: 46.8% in the Celtis forest and 38.9% in the Manilkara forest. The highest species richness were obtained in the first two soil layers (0–10 cm depth) while 21.8% and 21.4% of the species were exclusively found in the deepest layer (10–20 cm) in the Celtis forest and the Manilkara forest, respectively. Among the timber species found in the forest, only three were observed in the soil seed bank of the two sites: Nauclea diderrichii, Erythrophleum suaveolens and Staudtia kamerunensis. N. diderrichii was particularly abundant in the soil seed stock of the two sites (14.4–34.4 seeds m−2). Results suggested that to improve regeneration of timber species, planting in open forest habitats with seedlings coming from tree nursery should be more efficient.

Tropical forests are major providers of natural resources and ecosystem services but their ecological functions are at threat, due to increasing human pressure linked to economic development. The identification of priority areas for conservation is crucial for land use planning to ensure the protection of biodiversity and ecological function. Intact Forest Landscapes (IFLs), as defined by Greenpeace and World Resources Institute (WRI), are areas of the forest ecosystems not subjected to human activities. They have been identified by mapping human disturbances through remote sensing. Contrary to similar global-scale concepts, IFLs have been integrated into the standards of the certification body Forest Stewardship Council (FSC) and therefore have practical implications for forest management policies. The Motion 65, approved in the general assembly of FSC in 2014, mandates the protection of IFLs located in FSC certified logging concessions. Until the implementation of national standards, forestry operations are banished from 80% of the IFL area within each forest management unit. To trace the history and evaluate the suitability of IFLs in the Central African context, we searched for documents related to the IFL method, and related approaches focusing on the identification of areas devoid of human disturbances. The IFL method is simple and cost-effective and allows for a global assessment of the influence of human infrastructures and industrial exploitation on forests. However, the method does not consider the situation below the canopy and those forest components not visible by satellites. For example, hunting, one of the main threats faced by wildlife in Central African forests today, cannot be detected with satellite imagery. On the other hand, other anthropogenic activities which remote sensing may detect may be compatible with forest ecosystem conservation. To better tailor the IFL approach to Central African forests, we recommend (i) the consideration of wildlife communities in the intactness analysis, (ii) a thorough evaluation of the impacts of human activities on forest ecosystems, and (iii) the integration of local stakeholders and governments in the design of land management strategies to respond to social, economic and environmental needs.

This viewpoint paper presents a reaction to the article by Brandt et al. (2016). It highlights the complexities inherent to the attribution of deforestation impacts to policy interventions when using remote-sensing data. This critique argues that in the context of the Congo a suite of factors (i.e., population density in particular) other than those considered by Brandt et al. (e.g., type of forest, distance from roads and markets) play essential roles in determining the fates of forests. It also contends that care is needed when making decisions regarding which units will be included in the comparison group so that contextual factors and on-the-ground information are properly considered (e.g., when logging operations are inactive or when a concession is used for 'conservation' purposes). Finally, it proposes that a focus on an analysis of deforestation rates for a given level of timber production might be a metric that more accurately represents one aspect of the consequences of forest management, which should also consider the appraisal of trade-offs associated with a larger set of social, financial and ecological objectives.

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.

L'impact de l'exploitation forestière sur le renouvellement du stock de bois d'œuvre reste largement méconnu en Afrique centrale du fait du manque de données sur la dynamique des populations d’arbres exploités. C’est en particulier le cas pour l’azobé, Lophira alata, un bois d’œuvre de grande valeur commerciale. L'objectif de cette étude était double : quantifier l'effet du type forestier et de l’exploitation sur la croissance de l’azobé et simuler l'évolution du stock exploitable et du taux de reconstitution après exploitation. Nous avons effectué pendant trois ans un suivi annuel de la croissance et de la mortalité dans trois types forestiers au Cameroun : en forêt sempervirente, en forêt mixte et en forêt semi-caducifoliée. Le recrutement a été étudié uniquement en forêt mixte, sous deux conditions : en milieu exploité et non exploité. Nous avons calibré, avec ces données, un modèle matriciel de Usher. Douze scénarios d’exploitation ont été simulés en faisant varier le diamètre minimum d’exploitation de 60, 70 et 80 cm, et l’intensité de prélèvement des arbres de 100, 80, 60 et 40 %. La croissance de l’azobé est influencée à la fois par le type forestier et l'exploitation. Les arbres de forêt sempervirente ont une croissance plus faible jusqu’à 50 cm de diamètre, alors que la croissance maximale prédite pour ce type forestier est la plus élevée. L'exploitation a par ailleurs stimulé la croissance. Enfin, l’exploitation de l'azobé ne respecte pas le principe du rendement soutenu : ses taux de croissance à long terme varient entre 0,54 et 0,83. Pour garantir la durabilité de son exploitation au Cameroun, une sylviculture dynamisant la croissance des futurs arbres exploitables, ainsi que leur régénération, s’avère indispensable.

Forest degradation in the tropics is often associated with roads built for selective logging. The protection of Intact Forest Landscapes (IFL) that are not accessible by roads is high on the biodiversity conservation agenda, a challenge for logging concessions certified by the Forest Stewardship Council (FSC). A frequently advocated conservation objective is to maximise the retention of "roadless space", a concept that is based on distance to the nearest road from any point. We developed a novel use of the empty space function - a general statistical tool based on stochastic geometry and random sets theory - to calculate roadless space in a part of the Congo Basin where there has recently been rapid expansion of road networks. We compared the temporal development of roadless space in certified and non-certified logging concessions inside and outside areas declared as IFL in the year 2000. Between 1999 and 2007, rapid road network expansion led to a marked loss of roadless space in IFL. After 2007, this trajectory levelled out in most areas, due to an equilibrium between newly built roads and abandoned roads that became revegetated. However, concessions within IFL that have been certified by FSC since around 2007 showed continued decreases in roadless space, thus reaching a level comparable to all other concessions. Only national parks remained road-free. We recommend that forest management policies make the preservation of large connected forest areas a top priority by effectively monitoring - and limiting - the occupation of space by roads that are permanently accessible.

1. Irregularly shaped trees including trees with buttresses, flutes or stilt roots are frequent in tropical forests. The lack of an international standard to measure the diameter of such trees leads to high uncertainties in biomass estimation, tree growth and carbon budget monitoring. 2. In this study, we developed a new method based on terrestrial close-range photogrammetry for measuring and modelling irregular stems. This approach is cheap and easy to implement in the field as it only requires a camera and a graduated rod. We validated the approach with destructive cross-section measurements along the stem of three buttressed trees. To demonstrate the broader utility of this method, we extended the validated approach to 43 additional trees belonging to two species: Celtis mildbraedii (Ulmaceae) and Entandophragma cylindricum (Meliaceae). Based on the three dimensional models, we computed shape indices for each tree, and we analysed the stem morphology of the two species. Finally, we analysed some standardized predictors for the estimation of above-ground biomass. 3. We found a high concordance between diameters derived from the photogrammetric process and destructive diameter measurements along the stem for the three calibration trees. We found that C. mildbraedii develop much stronger irregularities than E. cylindricum. We also identified a large intraspecific variation in trunk morphology for E. cylindricum. The basal area at 1·3 m height (Darea130) seems to be a more robust predictor for biomass estimates (lowest Akaike information criterion and relative squared error) than diameter measured above buttresses (DAB) or diameter at breast height estimated from available taper model. Finally, Darea130 might be estimated with a good precision [root mean square error (RMSE) < 5%] with linear model based on the field measurements DAB and the perimeter of the convex hull of the buttresses at 1·3 m height (Dconvhull130). 4. In this study, we showed the high potential of the photogrammetry for measuring and modelling irregular stems. Photogrammetry could then be used as a non-destructive measurement tool to produce correction factors for standardizing the diameter of irregular stems at a reference height which is a key issue in tree growth monitoring and biomass change estimation. (Resume d'auteur)

Logging roads can trigger tropical forest degradation by reducing the integrity of the ecosystem and providing access for encroachment. Therefore, road management is crucial in reconciling selective logging and biodiversity conservation. Most logging roads are abandoned after timber harvesting; however, little is known about their long-term impacts on forest vegetation and accessibility, especially in Central Africa. In 11 logging concessions in the Congo Basin, we field-sampled a chronosequence of roads that, judging from satellite images, had been abandoned between 1985 and 2015. We assessed recovery of timber resources, tree diversity and above-ground biomass in three zones: the road track, the road edge (where forest had been cleared during road construction) and the adjacent logged forest. The density of commercial timber species

We present the first cross continental comparison of the flowering and fruiting phenology of tropical forests across Africa. Flowering events of 5,446 trees from 196 species across 12 sites, and fruiting events of 4,595 trees from 191 species, across 11 sites were monitored over periods of 6 to 29 years, and analysed to describe phenology at the continental level. To study phenology we used Fourier analysis to identify the dominant cycles of flowering and fruiting for each individual tree and we identified the time of year African trees bloom and bear fruit and their relationship to local seasonality. Reproductive strategies were diverse and no single regular cycle was found in >50% of individuals across all 12 sites. Additionally, we found annual flowering and fruiting cycles to be the most common. Sub-annual cycles were the next most common for flowering whereas supra-annual patterns were the next most common for fruiting. We also identify variation in different subsets of species, with species exhibiting mainly annual cycles most common in West and West-Central African tropical forests, while more species at sites in East-Central and Eastern African forests showed cycles ranging from sub-annual to supra-annual. Despite many trees showing strong seasonality, at most sites some flowering and fruiting occurred all year round. Environmental factors with annual cycles are likely to be important drivers of seasonal periodicity in trees across Africa, but proximate triggers are unlikely to be constant across the continent

Although the importance of large trees regarding biodiversity and carbon stock in old-growth forests is undeniable, their annual contribution to biomass production and carbon uptake remains poorly studied at the stand level. To clarify the role of large trees in biomass production, we used data of tree growth, mortality, and recruitment monitored during 20 yr in 10 4-ha plots in a species-rich tropical forest (Central African Republic). Using a random block design, three different silvicultural treatments, control, logged, and logged + thinned, were applied in the 10 plots. Annual biomass gains and losses were analyzed in relation to the relative biomass abundance of large trees and by tree size classes using a spatial bootstrap procedure. Although large trees had high individual growth rates and constituted a substantial amount of biomass, stand-level biomass production decreased with the abundance of large trees in all treatments and plots. The contribution of large trees to annual stand-level biomass production appeared limited in comparison to that of small trees. This pattern did not only originate from differences in abundance of small vs. large trees or differences in initial biomass stocks among tree size classes, but also from a reduced relative growth rate of large trees and a relatively constant mortality rate among tree size classes. In a context in which large trees are increasingly gaining attention as being a valuable and a key structural characteristic of natural forests, the present study brought key insights to better gauge the relatively limited role of large trees in annual stand-level biomass production. In terms of carbon uptake, these results suggest, as already demonstrated, a low net carbon uptake of old-growth forests in comparison to that of logged forests. Tropical forests that reach a successional stage with relatively high density of large trees progressively cease to be carbon sinks as large trees contribute sparsely or even negatively to the carbon uptake at the stand level.

Tree cover is a key variable for ecosystem functioning, and is widely used to study tropical ecosystems. But its determinants and their relative importance are still a matter of debate, especially because most regional and global analyses have not considered the influence of agricultural practices. More information is urgently needed regarding how human practices influence vegetation structure. Here we focused in Central Africa, a region still subjected to traditional agricultural practices with a clear vegetation gradient. Using remote sensing data and global databases, we calibrated a Random Forest model to correlatively link tree cover with climatic, edaphic, fire and agricultural practices data. We showed that annual rainfall and accumulated water deficit were the main drivers of the distribution of tree cover and vegetation classes (defined by the modes of tree cover density), but agricultural practices, especially pastoralism, were also important in determining tree cover. We simulated future tree cover with our model using different scenarios of climate and land-use (agriculture and population) changes. Our simulations suggest that tree cover may respond differently regarding the type of scenarios, but land-use change was an important driver of vegetation change even able to counterbalance the effect of climate change in Central Africa.

To understand the functioning of montane forests, this study was conducted in the highlands of the Kahuzi-Biega National Park in the Democratic Republic of the Congo. The relationship between the altitude and the floristic stability of woody layers and regeneration capability of canopy species after many years of disturbance was studied. Ten 1-ha plots were established from 1935m to 2760m a.s.l. In each plot we inventoried the trees =10cm of diameter at breast height (DBH), separating a canopy layer (10% of the tallest trees) and an understorey layer (all the other trees). In each plot, we nested a 0.1 ha subplot to inventory the saplings between 1 and 10 cm DBH. We found that the Jaccard index of dissimilarity between the understorey layer and the canopy layer decreases with the altitude. The proportion of species which are well represented in the three layers increases with the altitude. The number of pioneer species decreases with the altitude while that of non-pioneer and shade tolerant species increases. These findings suggest that altitude influences the stability of highland forests, higher altitude being more stable than lower ones in the case of this study.

Patterns in above-ground biomass of tropical forests over short altitudinal gradients are poorly known. The aim of this study was to investigate the variation of above-ground biomass with altitude in old-growth forests and determine the importance of changes in floristic composition as a cause of this variation. We used a dataset from 15 1-ha permanent plots established from lowland (200 m asl) to submontane forests (900 m asl) in the Ngovayang Massif, south-western Cameroon. We analysed variation over altitude in two specific functional traits, the potential maximum tree height and the wood density. Forest above-ground biomass decreased from 500–600 Mg ha−1 in lowland plots to around 260 Mg ha−1 at the highest altitudes. The contribution to above-ground biomass of large-tree species (dbh ≥ 70 cm) decreased with altitude, while the contribution of smaller trees was constant. Contribution of the Fabaceae subfamily Caesalpinioideae decreased with altitude, while those of Clusiaceae, Phyllanthaceae and Burseraceae increased. While potential maximum tree height significantly decreased, wood specific gravity displayed no trend along the gradient. Finally, the decrease in above-ground biomass along the short altitudinal gradient can be at least partially explained by a shift in species composition, with large-tree species being filtered out at the highest altitudes. These results suggest that global change could lead to significant shifts in the properties of montane forests over time.

Understanding how environmental factors could impact population dynamics is of primary importance for species conservation. Matrix population models are widely used to predict population dynamics. However, in species-rich ecosystems with many rare species, the small population sizes hinder a good fit of species-specific models. In addition, classical matrix models do not take into account environmental variability. We propose a mixture of regression models with variable selection allowing the simultaneous clustering of species into groups according to vital rate information (recruitment, growth and mortality) and the identification of group-specific explicative environmental variables. We develop an inference method coupling the R packages flexmix and glmnet. We first highlight the effectiveness of the method on simulated datasets. Next, we apply it to data from a tropical rain forest in the Central African Republic. We demonstrate the accuracy of the inhomogeneous mixture matrix model in successfully reproducing stand dynamics and classifying tree species into well-differentiated groups with clear ecological interpretations. Copyright (c) 2014 John Wiley & Sons, Ltd.

The greater part of the semi-deciduous moist forests of the Congo basin has been given to logging companies for exploitation. In the next decades, very few of these forests will remain intact. In this paper, we aimed to identify large-scale variations in the structure, composition and functioning of African moist forests that could serve as a baseline for both management and conservation purposes. Commercial forest inventory data were assembled for 49,711 0.5-ha plots, covering an area of more than six million hectares, crossing the borders of Cameroon, Central African Republic and Republic of Congo. Floristic composition was analyzed for a subset of 176 genera reliably identified in the field. Three key functional traits of tropical trees: regeneration guild, leaf phenology, and wood specific gravity, were collected at the species level from various sources, and assigned at the genus level. We first investigated the main variations in forest structure and composition, and identified seven forest types based on these variations. Differences in the percentage of pioneer and deciduous stems, and mean wood specific gravity were tested between forest types. Most of the study area was composed of a mosaic of the structural variations of the forests characterized by the occurrence of Celtis (Ulmaceae) species, which are mostly composed of frequent and abundant genera that formed the common floristic pool of the region. Secondary Musanga (Moraceae) forest is located in repeatedly disturbed areas, along roads and around main cities; mixed Manilkara (Sapotaceae) forest covers a huge area in the southern Central African Republic and in the northern Republic of Congo; and monodominant Gilbertiodendron (Fabaceae) forest is sparsely distributed along rivers. The contrasted structure, composition, and functioning of the forest types imply pronounced differences in population and ecosystem processes, and call for adapted management and conservation strategies.

AimIn this study we identified large-scale variation in tree species composition across tropical African forests and determined the underlying environmental and historical factors.LocationTropical forests from Senegal to Mozambique.MethodsDistribution data were gathered for 1175 tree species in 455 sample sites scattered across tropical Africa, including all types of tropical forests (wet, moist, dry, and lowland to moderate elevation montane forests). The value of elevation and 19 climatic variables extracted from the BIOCLIM data set were assigned to each sample site. We determined the variation in species composition using correspondence analysis and identified the environmental correlates. We defined floristic clusters according to species composition and identified the characteristic species using indicator analysis.ResultsWe identified a major floristic discontinuity located at the Albertine rift that separated the dry, moist and wet forests of West and Central Africa (the entire Guineo-Congolian Region) from the upland and coastal forests of East Africa. Except for the Albertine Rift, we found no evidence to support the other proposed floristic discontinuities (Dahomey Gap etc.). We detected two main environmental gradients across tropical African forests. The rainfall gradient was strongly correlated with the variation in tree species composition in West and Central Africa. The elevation/temperature gradient highlighted the major floristic differences within East Africa and between East Africa and the Guineo-Congolian Region, the latter being most probably due to the geological disruption and associated climatic history of the East African uplift.Main conclusionsWe found floristic evidence for three main biogeographical regions across the tropical African forests, and described six floristic clusters with particular environmental conditions within these regions: Coastal and Upland for East Africa, Dry and Wet-Moist for West Africa, and Moist and Wet for Central Africa.

Gérer durablement les forêts tropicales de production nécessite de connaître leur diversité, leur dynamique et l’impact de l’exploitation sur ces caractéristiques. Cette connaissance peut s’obtenir avec de grands dispositifs de suivi permanents dont les arbres sont régulièrement mesurés. Un seul dispositif de ce type existe en Afrique centrale, M’Baïki, installé en 1982 en République centrafricaine. Il a fait référence pour établir les règles d’aménagement d’un grand nombre de concessions forestières de la région. Plusieurs projets ont permis l’installation de nouveaux dispositifs, en privilégiant des zones situées sur des substrats géologiques différents. Deux dispositifs ont ainsi été installés dans des concessions forestières aménagées au nord de la République du Congo, sur alluvions de l’ère cénozoïque (Loundoungou) et sur grès de Carnot de l’ère mésozoïque (Mokabi). Chaque dispositif est constitué de deux blocs de 400 ha, contenant chacun un sentier d’environ 4 000 arbres d’une vingtaine d’espèces commerciales et deux parcelles de 9 ha suivies en plein. Tous les arbres supérieurs à 10 cm de diamètre sont mesurés. Un bloc sera exploité après quelques années de suivi, l’autre restera en témoin. L’installation et le premier inventaire d’un dispositif ont nécessité en moyenne 34 mois de travail avec une équipe de huit personnes pour un coût total moyen de 94500 €. Les peuplements des deux dispositifs, Loundoungou et Mokabi respectivement, ont les caractéristiques suivantes : 240 et 230 espèces d’arbres, densité de 346,8 (± 5,8) et 426,8 (± 3,6) arbres par hectare, biomasse de 432,6 (± 13,8) et 457,3 (± 9,3) Mg/ha. Les difficultés rencontrées lors de l’installation des dispositifs (emplacement, localisation des parcelles, constitution des sentiers, application du protocole d’inventaire, logistique...) sont analysées. Plusieurs recommandations sont émises pour améliorer l’installation de tels dispositifs et leur protocole d’inventaire.

Cette étude évalue l’abondance des graines d’Erythrophleum suaveolens dans la banque de semences du sol des forêts denses humides d’Afrique centrale. Les travaux ont été menés au Nord-Congo dans deux types forestiers : la forêt à Celtis sur des sols argilo-sableux à sablo-argileux et la forêt à Manilkara sur des sols sableux. Les tiges d’E. suaveolens (dhp ≥ 10 cm) ont été inventoriées dans deux blocs de 400 ha, et les structures diamétriques de leurs populations ont été comparées. En outre, 80 fosses (2 x 40 fosses par type de forêt) ont été creusées au pied de 20 arbres (10 par forêt), sur trois couches contiguës de 10 cm chacune, soit à une profondeur totale de 30 cm, et l’abondance des graines dans la banque de semences du sol a été évaluée. La dormance des graines récoltées a été testée par des essais de germination après traitement au H2SO4 et cinq graines prélevées jusqu'à une profondeur de 20 cm dans la forêt à Celtis ont été utilisées pour estimer leur âge par spectroscopie de masse par accélérateur (AMS). Dans les deux types forestiers, les structures diamétriques présentent une distribution gaussienne, illustrant un déficit de régénération. Alors que les densités de tiges (dhp ≥ 10 cm) sont proches, avec 0,85 et 1,05 tige/ha respectivement, dans la forêt à Celtis et la forêt à Manilkara. Les densités de graines sont significativement plus élevées dans la forêt à Celtis (8,55 graines/m2) que dans la forêt à Manilkara (0,15 graine/m2). Le pourcentage maximum de germination obtenu était de 19,1 % pour des graines n’ayant subi aucun traitement. Les lots traités à l’acide ont présenté de moindres taux de germination. Ces graines semblent pouvoir se conserver une dizaine d’années au moins dans la banque du sol. Les facteurs pouvant influencer les variations de densité des graines sont discutés et des recommandations sylvicoles sont formulées.

Forest degradation in the tropics is often associated with roads built for selective logging. The protection of intact forest landscapes (IFL) that are not accessible by roads is high on the biodiversity conservation agenda and a challenge for logging concessions certified by the Forest Stewardship Council (FSC). A frequently advocated conservation objective is to maximize the retention of roadless space, a concept that is based on distance to the nearest road from any point. We developed a novel use of the empty-space function - a general statistical tool based on stochastic geometry and random sets theory - to calculate roadless space in a part of the Congo Basin where road networks have been expanding rapidly. We compared the temporal development of roadless space in certified and uncertified logging concessions inside and outside areas declared IFL in 2000. Inside IFLs, road-network expansion led to a decrease in roadless space by more than half from 1999 to 2007. After 2007, loss leveled out in most areas to close to 0 due to an equilibrium between newly built roads and abandoned roads that became revegetated. However, concessions in IFL certified by FSC since around 2007 continuously lost roadless space and reached a level comparable to all other concessions. Only national parks remained mostly roadless. We recommend that forest-management policies make the preservation of large connected forest areas a top priority by effectively monitoring - and limiting - the occupation of space by roads that are permanently accessible.

Tropical forests account for over 50 % of the global forested area and forest carbon stock. Although the deforestation rate is tending to decline, forests are confronted with climate change, which could profoundly modify their functioning. The migration of species that took place during the Pleistocene is no longer possible because human activities have markedly altered tropical landscapes. Forest species will thus have to adapt (or not) particularly to the increased water stress. Forest management methods must incorporate new knowledge on the vulnerability of species and evolve in order to reduce potentially negative interactions between disturbances and the water deficit. A key challenge is to identify trade-offs between logging in water deficit situations and the increased forest fire risk. In drylands, factors related to climate change are meshed with other change factors, but innovations in the management of woodlands could ensure their long-term persistence.

International audience; Phenotypic traits and their associated trade-offs have been shown to have globally consistent effects on individual plant physiological functions1, 2, 3, but how these effects scale up to influence competition, a key driver of community assembly in terrestrial vegetation, has remained unclear4. Here we use growth data from more than 3 million trees in over 140,000 plots across the world to show how three key functional traits-wood density, specific leaf area and maximum height-consistently influence competitive interactions. Fast maximum growth of a species was correlated negatively with its wood density in all biomes, and positively with its specific leaf area in most biomes. Low wood density was also correlated with a low ability to tolerate competition and a low competitive effect on neighbours, while high specific leaf area was correlated with a low competitive effect. Thus, traits generate trade-offs between performance with competition versus performance without competition, a fundamental ingredient in the classical hypothesis that the coexistence of plant species is enabled via differentiation in their successional strategies5. Competition within species was stronger than between species, but an increase in trait dissimilarity between species had little influence in weakening competition. No benefit of dissimilarity was detected for specific leaf area or wood density, and only a weak benefit for maximum height. Our trait-based approach to modelling competition makes generalization possible across the forest ecosystems of the world and their highly diverse species composition.