We report above-ground biomass (AGB), basal area, stem density 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- compared with neotropical 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 relationships with C : N ratio (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.

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.

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.

Plinio Sist 10,88 | Bonaventure Sonke 60 | J. Daniel Soto 21 | Cintia Rodrigues de Souza 24 | Juliana Stropp 89 | Martin J. P. Sullivan 35 | Ben Swanepoel 34 | Hans ter Steege 25,90 | John Terborgh 91,92 | Nicolas Texier 93 | Takeshi Toma 94 | Renato Valencia 95 | Luis Valenzuela 75 | Leandro Valle Ferreira 96 | Fernando Cornejo Valverde 97 | Tinde R. Van Andel 25 | Rodolfo Vasque 77 | Hans Verbeeck 61 | Pandi Vivek 22 | Abstract Aim:Large tropical trees form the interface between ground and airborne observations, offering a unique opportunity to capture forest properties remotely and to investigate their variations on broad scales. However, despite rapid development of metrics to characterize the forest canopy from remotely sensed data, a gap remains between aerial and field inventories. To close this gap, we propose a new pan-tropical model to predict plot-level forest structure properties and biomass from only the largest trees.Location: Pan-tropical.Time period: Early 21st century. Major taxa studied: Woody plants.Methods: Using a dataset of 867 plots distributed among 118 sites across the tropics, we tested the prediction of the quadratic mean diameter, basal area, Lorey's height, community wood density and aboveground biomass (AGB) from the ith largest trees. Results:Measuring the largest trees in tropical forests enables unbiased predictions of plot-and site-level forest structure. The 20 largest trees per hectare predicted quadratic mean diameter, basal area, Lorey's height, community wood density and AGB with 12, 16, 4, 4 and 17.7% of relative error, respectively. Most of the remaining error in biomass prediction is driven by differences in the proportion of total biomass held in medium-sized trees (50-70 cm diameter at breast height), which shows some continental dependency, with American tropical forests presenting the highest proportion of total biomass in these intermediate-diameter classes relative to other continents. Main conclusions:Our approach provides new information on tropical forest structure and can be used to generate accurate field estimates of tropical forest carbon stocks to support the calibration and validation of current and forthcoming space missions. It will reduce the cost of field inventories and contribute to scientific understanding of tropical forest ecosystems and response to climate change. K E Y W O R D Scarbon, climate change, forest structure, large trees, pan-tropical, REDD+, tropical forest ecology

Quantifying the relationship between tree diameter and height is a key component of efforts to estimate biomass and carbon stocks in tropical forests. Although substantial site‐to‐site variation in height–diameter allometries has been documented, the time consuming nature of measuring all tree heights in an inventory plot means that most studies do not include height, or else use generic pan‐tropical or regional allometric equations to estimate height.Using a pan‐tropical dataset of 73 plots where at least 150 trees had in‐field ground‐based height measurements, we examined how the number of trees sampled affects the performance of locally derived height–diameter allometries, and evaluated the performance of different methods for sampling trees for height measurement.Using cross‐validation, we found that allometries constructed with just 20 locally measured values could often predict tree height with lower error than regional or climate‐based allometries (mean reduction in prediction error = 0.46 m). The predictive performance of locally derived allometries improved with sample size, but with diminishing returns in performance gains when more than 40 trees were sampled. Estimates of stand‐level biomass produced using local allometries to estimate tree height show no over‐ or under‐estimation bias when compared with biomass estimates using field measured heights. We evaluated five strategies to sample trees for height measurement, and found that sampling strategies that included measuring the heights of the ten largest diameter trees in a plot outperformed (in terms of resulting in local height–diameter models with low height prediction error) entirely random or diameter size‐class stratified approaches.Our results indicate that even limited sampling of heights can be used to refine height–diameter allometries. We recommend aiming for a conservative threshold of sampling 50 trees per location for height measurement, and including the ten trees with the largest diameter in this sample.