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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 high mortality and low recruitment of the myrmecophytic Acacia drepanolobium are not consistent with the demographic rates of monodominant species. The positive conspecific density dependence observed from the spatial analysis is consistent with the defensive benefits conferred by symbiotic ants to trees when they grow close to each other. Patches of savanna dominated by Acacia drepanolobium occur throughout East Africa on nutrient-rich vertisols, also known as black cotton soils. We assessed the survival and recruitment for all freestanding trees with diameter at knee height (dkh) ≥ 10 mm in one of such mono-dominated patches (47 ha) at the Mpala Research Centre, Kenya, with the aim of identifying demographic traits that might explain the dominance of this species. Over a mean 6-year interval, mortality and recruitment rates in the habitat were 4.55%/year and 1.42%/year respectively, resulting in a net loss of 17.8% of the initial individuals. Of the 30 species recorded from the first census, 11 decreased in abundance, nine increased, and the remainder 10 did not change in abundance. The monodominant A. drepanolobium had a high mortality (4.69%/year), a low recruitment (1.31%/year), and a 19% population decline. There was no evidence of conspecific negative density dependence for this species. Rather, we found a statistically significant positive correlation between the number of conspecific neighbors and individual-level probability of survival, consistent with the “shared defense” benefits that symbiotic ant colonies occupying multiple trees can confer to these latter in a small neighborhood. Thus, mortality of A. drepanolobium was higher in areas where it occurred in lower densities, which resulted in an increase in the spatial aggregation of conspecifics. Mortality increased with dkh size classes and was mostly caused by elephants and stem-boring beetles. The demographic rates during the study period in theory are inconsistent with those of monodominant species. The protection against herbivory conferred by mutualistic ants associated with this species remains the most probable explanation of its dominance in this habitat.

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 need to address wildlife conservation outside of protected areas has become more urgent than ever before to meet environmental and socio‐economic goals. However, there is limited knowledge about how woody plants respond to herbivory within landscapes shared by wildlife and domestic herbivores in African savanna, thus management decisions might be based on inaccurate information and ultimately be ineffective. We compared woody vegetation dynamics between two adjacent ranches with different management objectives and subjected to varying levels of herbivory by both wildlife and domesticated mammals using 421 square plots of 400 m2 nested on three transects, each 3 km long and purposively selected to minimize bio‐physical differences. Both species and structural diversity were significantly higher (p

QUESTIONS: We examined the spatial patterns of dominant canopy species in a tropical forest to investigate: (a) what is the niche occupancy of canopy species with respect to topographic gradients; (b) what are the dominant ecological processes that explain their distribution; (c) what are the interactions among the most prevalent canopy species; and (d) what are the interactions between canopy species adults and juveniles trees? LOCATION: Rabi permanent CTFS‐ForestGEO plot, Gabon. METHODS: We selected the four most abundant canopy species and one timber species. We used Berman's test to determine the effect of three topographic variables on the distribution of each species and univariate analysis to model the spatial pattern of each species using either an inhomogeneous Poisson process or an inhomogeneous Cox process. We also used a bivariate form of the pair correlation function (PCF) to determine the spatial interaction between species and the correlation among conspecific adult and juvenile trees. RESULTS: Four of the five species had aggregated spatial patterns while Lophira alata showed spatial randomness. Most of the variance in the local tree density was explained by within‐population dispersal processes rather than environmental factors. Bivariate PCF tests showed significant segregation between species associations. Two species exhibited aggregation at small distances between young and adult trees, while others showed either complete spatial randomness at small inter‐tree distances or segregation at large distances. CONCLUSIONS: This study showed that the spatial pattern in the majority of canopy species was aggregation. Seed dispersal limitation mainly explained the observed aggregation pattern. Habitat filtering, as evidenced by the influence of topographic variables on niche occupancy, marginally, yet significantly, explained this pattern. The different spatial patterns of the principal species permit their coexistence. Spatial segregation among adult and juvenile trees reveals a strong pattern of either species‐specific seed predation or pathogens.

Stand structure and tree species diversity patterns were examined plot-wide and among four topographically defined habitats (plateau, cliff, low plain and depressions) in a 120-ha permanent plot in an Acacia-dominated savanna in Mpala Ranch, central Kenya. The four habitats were defined by clustering the 3000 quadrats of 20 × 20 m in the plot based on their altitude, slope and convexity. Structural and floristic differences among the four habitats were examined and species-habitat associations were tested for the 30 most abundant species using torus translation randomization tests. The plot included 113 337 trees in 62 species with diameter at knee height ≥ 2 cm (18.4 species ha−1), 41 genera and 23 families. Fabaceae with the genus Acacia were the dominant family, followed by Euphorbiaceae and Ebenaceae. Tree density and basal area were twice as high on low plain and depressions than on the plateau. Species richness was highest in the cliff and was seven times higher than in the adjacent plateau. Half of the species assessed showed significant positive associations with one habitat and 21 showed significant negative associations with at least one habitat. The variation in stand structure and tree species diversity within the Mpala plot shows that topography is among the important drivers of local species distribution and hence the maintenance of tree diversity in savannas.

Arbuscular mycorrhizal (AM) and ectomycorrhizal (EcM) associations are critical for host-tree performance. However, how mycorrhizal associations correlate with the latitudinal tree beta-diversity remains untested. Using a global dataset of 45 forest plots representing 2,804,270 trees across 3840 species, we test how AM and EcM trees contribute to total beta-diversity and its components (turnover and nestedness) of all trees. We find AM rather than EcM trees predominantly contribute to decreasing total beta-diversity and turnover and increasing nestedness with increasing latitude, probably because wide distributions of EcM trees do not generate strong compositional differences among localities. Environmental variables, especially temperature and precipitation, are strongly correlated with beta-diversity patterns for both AM trees and all trees rather than EcM trees. Results support our hypotheses that latitudinal beta-diversity patterns and environmental effects on these patterns are highly dependent on mycorrhizal types. Our findings highlight the importance of AM-dominated forests for conserving global forest biodiversity., The relationship of mycorrhizal associations with latitudinal gradients in tree beta-diversity is unexplored. Using a global dataset approach, this study examines how trees with arbuscular mycorrhizal and ectomycorrhizal associations contribute to latitudinal beta-diversity patterns and the environmental controls of these patterns.

In 2015 and 2016, the AfriSAR campaign was carried out as a collaborative effort among international space and National Park agencies (ESA, NASA, ONERA, DLR, ANPN and AGEOS) in support of the upcom...

The relative abundance patterns of tropical trees have been of interest since the expeditions of Alfred Russel Wallace, but little is known about how differences in relative abundance relate to reproductive patterns. Flowering is resource-dependent and fitness differences as well as differences in the quality of the abiotic and biotic neighborhood may contribute to the variation in reproductive status responsible for population-level flowering patterns. This variation determines the density and distance between flowering conspecifics and may alter relative abundance extremes among species during reproduction, factors known to influence pollination success. We collected flowering status data for a guild of twenty-three co-occurring tree species that flower in the understory of the Korup Forest Dynamics Plot in Cameroon. We examined how the occurrence and location of reproductive events were related to spatial patterns of adult abundance, focal tree size, neighborhood crowding, and habitat, while accounting for the influence of shared ancestry. Across species, the probability of flowering was higher for individuals of rarer species and for larger individuals but was unrelated to neighborhood crowding or habitat differences. Relative abundance extremes were reduced when only flowering individuals were considered, leading to a negative relationship between plot abundance and flowering probability at the species level that was not structured by shared ancestry. Spatially, flowering conspecifics tended to be overdispersed relative to all adult conspecifics. Rare species are predicted to suffer Allee effects or reduced fitness due to the difficulty of finding mates at low densities and frequencies. Here, however, rare species appear to maximize the size of their mate pool, compared to abundant species. If this partial ‘leveling of the playing field’ during reproduction is typical, it has consequences for our understanding of biodiversity maintenance and species coexistence in tropical forests.

Legumes provide an essential service to ecosystems by capturing nitrogen from the atmosphere and delivering it to the soil, where it may then be available to other plants. However, this facilitation by legumes has not been widely studied in global tropical forests. Demographic data from 11 large forest plots (16–60 ha) ranging from 5.25° S to 29.25° N latitude show that within forests, leguminous trees have a larger effect on neighbor diversity than non-legumes. Where soil nitrogen is high, most legume species have higher neighbor diversity than non-legumes. Where soil nitrogen is low, most legumes have lower neighbor diversity than non-legumes. No facilitation effect on neighbor basal area was observed in either high or low soil N conditions. The legume–soil nitrogen positive feedback that promotes tree diversity has both theoretical implications for understanding species coexistence in diverse forests, and practical implications for the utilization of legumes in forest restoration., Xu et al. examine the effect of leguminous trees on neighbor diversity across 11 plots in tropical forests around the world, and find that in high soil nitrogen conditions, most legume species have higher neighbor diversity than non-legumes, and vice versa where soil nitrogen is low. Their results have practical implications for the utilization of legumes in forest restoration.

Do hotspots of plant biodiversity translate into hotspots in the abundance and diversity of large mammalian herbivores? A common expectation in community ecology is that the diversity of plants and animals should be positively correlated in space, as with the latitudinal diversity gradient and the geographic mosaic of biodiversity. Whether this pattern ‘scales down’ to landscape-level linkages between the diversity of plants or the activities of highly mobile megafauna has received less attention. We investigated spatial associations between plants and large herbivores by integrating data from a plant-DNA-barcode phylogeny, camera traps, and a comprehensive map of woody plants across the 1.2-km2 Mpala Forest Global Earth Observatory (ForestGEO) plot, Kenya. Plant and large herbivore communities were strongly associated with an underlying soil gradient, but the richness of large herbivore species was negatively correlated with the richness of woody plants. Results suggest thickets and steep terrain create associational refuges for plants by deterring megaherbivores from browsing on otherwise palatable species. Recent work using dietary DNA metabarcoding has demonstrated that large herbivores often directly control populations of the plant species they prefer to eat, and our results reinforce the important role of megaherbivores in shaping vegetation across landscapes.

Background and aims – Cola, the second largest genus of the Malvaceae-Sterculioideae comprises 100–135 small to large tree species confined in nature to African forests, though cultivated elsewhere. Current species distribution ranges show that the genus is highly diverse in the seasonally wet forests along the Nigeria-Cameroon border, including the Korup National Park (KNP). In this paper we examine the diversity and abundance of Cola in KNP compared to other forests for which comparable data are available. We also describe two novelties in the genus.Methods – We used inventory data from a 50-ha permanent plot in southern KNP where all Cola trees and saplings down to 1 cm in diameter were tagged, mapped and identified. Additional collections of the genus came from the 11 km trail leading to the plot. Classic herbarium techniques and field observations were used for the morphological identification and description of specimens at MO and YA and from our personal collections. Cola species richness and abundance was estimated from the plot data and compared to other African forest sites for which comparable data are available. The evaluation of the conservation status of the two new species described in this paper followed the IUCN Red List Categories and Criteria. Key results – Twenty-five species of Cola were identified in the southern part of the KNP, including four undescribed species, raising the total number of Cola species in Cameroon to 46. The abundance of the genus in KNP was three orders of magnitude higher than in the Rabi forest in southwestern Gabon or in the Ituri forest in eastern D.R. Congo. This high species richness and abundance suggests that KNP is part of the center of diversity of the genus. Two new species, Cola zemagoana Kenfack & D.W.Thomas and C. mamboana Kenfack & Sainge are described and illustrated. Both species are only known from the lowland rainforest of southwestern Cameroon. Cola zemagoana is narrow endemic of southern KNP and its conservation status is assessed as Endangered. Cola mamboana is confined to the lowland forests of southwestern Cameroon, is locally very abundant in protected areas and is also assigned the conservation status Endangered.

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

Analysis of patterns in the distribution of taxa can provide important insights into ecological and evolutionary processes. Microbial biogeographic patterns almost always appear to be weaker than those reported for plant and animal taxa. It is as yet unclear why this is the case. Some argue that microbial diversity scales differently over space because microbial taxa are fundamentally different in their abundance, longevity and dispersal abilities. Others have argued that differences in scaling are an artifact of how we assess microbial biogeography, driven, for example, by differences in taxonomic resolution, spatial scale, sampling effort or community activity/dormancy. We tested these alternative explanations by comparing bacterial biogeographic patterns in soil to those of trees found in a forest in Gabon. Altering taxonomic resolution, excluding inactive individuals, or adjusting for differences in spatial scale were insufficient to change the rate of microbial taxonomic turnover. In contrast, we account for the differences in spatial turnover between these groups by equalizing sampling extent. Our results suggest that spatial scaling differences between microbial and plant diversity are likely not due to fundamental differences in biology, and that sampling extent should be taken into account when comparing the biogeographic patterns of microorganisms and larger organisms.

International audience; In 2015 and 2016, the AfriSAR campaign was carried out as a collaborative effort among international space and National Park agencies (ESA, NASA, ONERA, DLR, ANPN and AGEOS) in support of the upcoming ESA BIOMASS, NASA-ISRO Synthetic Aperture Radar (NISAR) and NASA Global Ecosystem Dynamics Initiative (GEDI) missions. The NASA contribution to the campaign was conducted in 2016 with the NASA LVIS (Land Vegetation and Ice Sensor) Lidar, the NASA L-band UAVSAR (Uninhabited Aerial Vehicle Synthetic Aperture Radar). A central motivation for the AfriSAR deployment was the common AGBD estimation requirement for the three future spaceborne missions, the lack of sufficient airborne and ground calibration data covering the full range of ABGD in tropical forest systems, and the intercomparison and fusion of the technologies. During the campaign, over 7000 km2 of waveform Lidar data from LVIS and 30,000 km2 of UAVSAR data were collected over 10 key sites and transects. In addition, field measurements of forest structure and biomass were collected in sixteen 1-hectare sized plots. The campaign produced gridded Lidar canopy structure products, gridded aboveground biomass and associated uncertainties, Lidar based vegetation canopy cover profile products, Polarimetric Interferometric SAR and Tomographic SAR products and field measurements. Our results showcase the types of data products and scientific results expected from the spaceborne Lidar and SAR missions; we also expect that the AfriSAR campaign data will facilitate further analysis and use of waveform lidar and multiple baseline polarimetric SAR datasets for carbon cycle, biodiversity, water resources and more applications by the greater scientific community.

The Acacia drepanolobium (also known as Vachellia drepanolobium) ant‐plant symbiosis is considered a classic case of species coexistence, in which four species of tree‐defending ants compete for nesting space in a single host tree species. Coexistence in this system has been explained by trade‐offs in the ability of the ant associates to compete with each other for occupied trees versus the ability to colonize unoccupied trees. We seek to understand the proximal reasons for how and why the ant species vary in competitive or colonizing abilities, which are largely unknown. In this study, we use RADseq‐derived SNPs to identify relatedness of workers in colonies to test the hypothesis that competitively dominant ants reach large colony sizes due to polygyny, that is, the presence of multiple egg‐laying queens in a single colony. We find that variation in polygyny is not associated with competitive ability; in fact, the most dominant species, unexpectedly, showed little evidence of polygyny. We also use these markers to investigate variation in mating behavior among the ant species and find that different species vary in the number of males fathering the offspring of each colony. Finally, we show that the nature of polygyny varies between the two commonly polygynous species, Crematogaster mimosae and Tetraponera penzigi: in C. mimosae, queens in the same colony are often related, while this is not the case for T. penzigi. These results shed light on factors influencing the evolution of species coexistence in an ant‐plant mutualism, as well as demonstrating the effectiveness of RADseq‐derived SNPs for parentage analysis.

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

Rhaptopetalumrabiense Kenfack & Nguema, sp. nov. from the Rabi forest in south-western Gabon is described, illustrated and assigned a provisional conservation status of “Critically Endangered”. An identification key to the five Gabonese species of Rhaptopetalum is also provided.

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.

AIM: The relationship between the proportion of sites occupied by a species and the area of a site [occupancy–area relationship (OAR)] offers key information for biodiversity management and has long fascinated ecologists. We quantified the variation in OAR for 3,157 woody species in 17 forest plots worldwide and tested the relative importance of environment and species traits for explaining this variation and evaluated overall model predictive ability. LOCATION: Global. TIME PERIOD: Early 21st century. MAJOR TAXA STUDIED: Woody plants. METHODS: We used mixed‐effect regression to examine the observed shape of the OAR (its “slope”) against species‐specific and plot‐wide predictors: coarse‐grain occupancy, tree size, plot species richness, energy availability and topographic complexity. RESULTS: We found large variation in OAR slopes, and the variation was strongest among species within plots. The OAR slopes showed a latitudinal trend and were steeper near the equator. As predicted, coarse‐grain occupancy and tree size negatively affected OAR slopes, whereas species richness had a positive effect and explained most of the variance between plots. Although hypothesized directionalities were broadly confirmed, traits and environment had relatively limited overall predictive power. MAIN CONCLUSIONS: These results document the variation of the OAR for 3,157 species at near‐global extent. We found a latitudinal gradient in OAR slopes and confirmed key hypothesized predictors. But at this global extent and over the large set of species analysed, the remaining unexplained variation in OAR slopes was substantial. Nevertheless, this large‐scale empirical analysis of the OAR offers an initial step towards a more general use of OARs for the fine‐scale prediction of species distributions and abundance.

Rhaptopetalumrabiense Kenfack & Nguema, sp. nov. from the Rabi forest in south-western Gabon is described, illustrated and assigned a provisional conservation status of “Critically Endangered”. An identification key to the five Gabonese species of Rhaptopetalum is also provided.

Gautier, L., O. Lachenaud, X. Van der Burgt & D. Kenfack (2016). Five new species of Englerophytum K. Krause (Sapotaceae) from central Africa. Candollea 71 ≥ : 287–305. In English, English and French abstracts. Five new species of Englerophytum K. Krause (Sapotaceae) are described : Englerophytum paludosum L. Gaut., Burgt & O. Lachenaud, Englerophytum gigantifolium O. Lachenaud & L. Gaut., Englerophytum libenii O. Lachenaud & L. Gaut., Englerophytum sylverianum Kenfack & L. Gaut., and Englerophytum ferrugineum L. Gaut. & O. Lachenaud. All five species are illustrated with line drawings and three of them with field photos. Distribution maps are also provided, and a preliminary extinction risk assessment according to IUCN Categories and Criteria is carried out : Englerophytum paludosum is assessed as “Least Concern”, Englerophytum sylverianum as “Vulnerable”, Englerophytum libenii and Englerophytum ferrugineum as “Endangered”, and Englerophytum gigantifolium as “Critically Endangered”.

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

Climate is widely recognised as an important determinant of the latitudinal diversity gradient. However, most existing studies make no distinction between direct and indirect effects of climate, which substantially hinders our understanding of how climate constrains biodiversity globally. Using data from 35 large forest plots, we test hypothesised relationships amongst climate, topography, forest structural attributes (stem abundance, tree size variation and stand basal area) and tree species richness to better understand drivers of latitudinal tree diversity patterns. Climate influences tree richness both directly, with more species in warm, moist, aseasonal climates and indirectly, with more species at higher stem abundance. These results imply direct limitation of species diversity by climatic stress and more rapid (co-)evolution and narrower niche partitioning in warm climates. They also support the idea that increased numbers of individuals associated with high primary productivity are partitioned to support a greater number of species.

The African genus Crotonogynopsis Pax (Euphorbiaceae) is revised to include four species, including two novelties, C. korupensis Kenfack & D. W. Thomas from the Korup National Park, Cameroon, and the Reserva Natural de Rio Campo in Equatorial Guinea; and C. australis Kenfack & Gereau from the southern part of the Eastern Arc Mountains of Tanzania, with a distant outlier in Mozambique. Three of the four species are assigned the IUCN Red List category of Least Concern (LC) on the basis of their occurrence in protected areas with no known threats.

Background and aims – There is increasing recognition that plant traits mediate environmental influence on species distribution, justifying non-random community assembly. We studied the influence of local scale edaphic factors on the distribution of functional traits in a tropical rainforest of Cameroon with the aim to find correlations between the main edaphic gradient and community functional trait metrics (weighted mean trait, functional divergence and intraspecific variation). Methods – Within the Korup Forest Dynamics Plot (50 ha), we randomly selected 44 quadrats of 0.04 ha each, collected soils and analysed 11 topography and soil variables. Leaves were harvested from all 98 tree species found in the quadrats to calculate community trait metrics [quadrat-level weighted mean (qk) and functional divergence (FDivk)] for leaf area (LA), specific leaf area (SLA), leaf phosphorus (LPC), leaf nitrogen concentration (LNC) and nitrogen to phosphorus ratio (N:P ratio). We examined relationships between the main edaphic gradient with qk, with FDivk and with intraspecific variation and interpreted correlations as the effects of abiotic filtering and competitive interaction. Key results – Soil fertility was the main edaphic gradient and was significantly correlated with qk for LPC, LNC and LA and with FDivk for LPC, N:P ratio, LA and SLA, confirming the influence of abiotic filtering and competitive interaction by the soil fertility gradient, respectively. For a given trait, quadrats were either over-dispersed or under-dispersed, accounting for 7–33 % of non-random trait distribution along the soil fertility gradient. Trends in intraspecific traits variation were consistently lower than quadrat-level mean traits along the soil fertility gradient. Conclusions – This study demonstrates the influence of soil fertility gradient on local scale community trait distribution and its contribution to non-random community assembly., SCOPUS: ar.j, info:eu-repo/semantics/published

With the emergence of a new field, biodiversity informatics, an important task has been to evaluate completeness of biodiversity information that is existing and available for various countries and regions. This paper offers a first and very basic assessment of sampling gaps and inventory completeness across the Cameroon Mountains. Because digital accessible knowledge is severely limited for the region, we relied on qualitative evaluations of inventory completeness, supplemented by large amounts of data from the National Herbarium of Cameroon (YA) database. Detailed botanical inventories have been developed for Mt Cameroon, the Kupe-Mwanenguba Mountains, Mt Oku, and the Mambila Plateau, leaving substantial geographic and environmental coverage gaps corresponding to Rumpi Hills, Mt Nlonako, Kimbi Fungom National Park, Bali and Bafut Ngemba, Mt Bamboutos, Kagwene, and Tchabal Mbabo. This paper provides a roadmap for a comprehensive botanical survey for this region. Completing this survey plan, the resulting data will allow researchers to track changes in biodiversity and identify priority areas for conservation on the various mountain ranges that make up this important biodiversity hotspot.

Ant-plant interactions mediated by special structures provided by plants such as domatia, extrafloral nectaries (EFNs) and food bodies, are very frequent in tropical ecosystems. To understand why ants are frequently encountered on most species of Carapa Aubl. (Meliaceae), we investigated the presence of extranuptial nectaries (ENNs) in all 27 species of the genus, spanning its entire distributional range in tropical Africa and America. We report for the first time in the genus the occurrence of extrafloral nectaries (at the base of the petiole, along the rachis of the pinnately compound leaf, on bracts) petaline nectaries (on the outer surface of petals), and pericarpial nectaries (on the surface of fruits), and confirm the presence of nectaries on leaflets in Carapa. Petiolar nectaries are the most common, occurring in 85% of the species. Nectaries were mainly active in young developing plant organs. Ants were observed foraging on exudates from these nectaries. The secretions from these glands help to explain the abundance of ants on Carapa trees. Although similar nectaries were also found in other members of the subfamily Cedreloideae, their position and frequency provide new characters for the identification of Carapa species in the field and the herbarium. As in other myrmecophilous plants, ENNs probably confer adaptive advantages to Carapa trees.

Morphometric analysis of Phyllobotryon Muell.Arg. in the Korup Forest Area of Cameroon recognizes three distinct morphospecies (1-3), which show significant variation in several leaf and fruit characters. In order to clarify the taxonomy of Phyllobotryon, we conducted univariate and multivariate analyses on sixteen quantitative and four qualitative characters scored from 111 fresh samples. Analysis of Variance revealed nine significant quantitative characters from which the first three Principal Components accounted for 74.6 % of the total variation. Results from Discriminant Analysis strongly support the existence of two groups (96.2 % and 100 %) representing Morphospecies 2 and 3, but morphospecies 2 is only weakly supported (88.9 %) as distinct from morphospecies 1. Characters such as petiole length, fruit surface ornamentation, style & calyx persistence, flower and fruit orientation and leaf shape are of

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.

Summary 1. In highly diverse ecosystems, such as tropical forests, the relative importance of mechanisms underlying species coexistence (e.g. habitat filtering, competitive exclusion, neutral dynamics) is still poorly known and probably varies depending on spatial and phylogenetic scales. 2. Here, we develop new approaches for dissecting simultaneously the phylogenetic structure of communities at different phylogenetic depths and spatial scales. We tested with simulations that our method is able to disentangle overdispersion and clustering effects occurring at contrasted phylogenetic depths. 3. We applied our approaches to a 50 ha Forest Dynamic Plot located in Korup National Park (Cameroon) where 329,000 tree stems ≥ 1 cm in diameter were identified and mapped, and using a newly generated dated molecular phylogenetic tree based on 2 plastid loci (rbcL and matK), including 272 species from Korup (97% of the individuals). 4. Significant patterns of phylogenetic turnover were detected across 20 9 20 m 2 quadrats at most spatial scales, with higher turnover between topographic habitats than within habitats, indicating the prevalence of habitat filtering processes. Spatial phylogenetic clustering was detected over the entire range of phylogenetic depths indicating that competitive exclusion does not generate a pattern of phylogenetic overdispersion at this scale, even at a shallow phylogenetic depth. 5. Using an individual-based approach, we also show that closely related species tended to aggregate spatially until a scale of 1 m. However, the signal vanishes at smaller distance, suggesting that competitive exclusion can balance the impact of environmental filtering at a very fine spatial scale. 6. Synthesis. Using new methods to characterize the structure of communities across spatial and phylogenetic scales, we inferred the relative importance of the mechanisms underlying species coexistence in tropical forests. Our analysis confirms that environmental filtering processes are key in the structuring of natural communities at most spatial scales. Although negative-density tends to limit coexistence of closely related species at very short distance (

Summary. Two new species of Afrothismia are described within the framework of an intended revision of the genus in Cameroon, A. fungiformis from the submontane forest of Mt Kupe and A. pusilla from Mt Kala. The new species are illustrated and the conservation status for both is assessed as Critically Endangered.

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

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

Gambeya korupensis Ewango & Kenfack (Sapotaceae: Chrysophylloideae), a new rain forest tree species from the Southwest Region in Cameroon, is described and illustrated. A distribution map is provided. G. korupensis has the leaf blade below pubescent on the midribs and secondary nerves, flowers with a pedicel 0.5 – 1 mm long, and a fruit which is ovoid, attenuate at the apex, 5-ridged, verrucose between the ridges, and bright red at maturity. The conservation status of G. korupensis is assessed as Vulnerable according to IUCN criteria.

Aims With the aim of understanding why some of the world's forests exhibit higher tree beta diversity values than others, we asked: (1) what is the contribution of environmentally related variation versus pure spatial and local stochastic variation to tree beta diversity assessed at the forest plot scale; (2) at what resolution are these beta-diversity components more apparent; and (3) what determines the variation in tree beta diversity observed across regions/continents? Location World-wide. Methods We compiled an unprecedented data set of 10 large-scale stem-mapping forest plots differing in latitude, tree species richness and topographic variability. We assessed the tree beta diversity found within each forest plot separately. The non-directional variation in tree species composition among cells of the plot was our measure of beta diversity. We compared the beta diversity of each plot with the value expected under a null model. We also apportioned the beta diversity into four components: pure topographic, spatially structured topographic, pure spatial and unexplained. We used linear mixed models to interpret the variation of beta diversity values across the plots. Results Total tree beta diversity within a forest plot decreased with increasing cell size, and increased with tree species richness and the amount of topographic variability of the plot. The topography-related component of beta diversity was correlated with the amount of topographic variability but was unrelated to its species richness. The unexplained variation was correlated with the beta diversity expected under the null model and with species richness. Main conclusions Because different components of beta diversity have different determinants, comparisons of tree beta diversity across regions should quantify not only overall variation in species composition but also its components. Global-scale patterns in tree beta diversity are largely coupled with changes in gamma richness due to the relationship between the latter and the variation generated by local stochastic assembly processes.

Plant species delimitation within tropical ecosystems is often difficult because of the lack of diagnostic morphological characters that are clearly visible. The development of an integrated approach, which utilizes several different types of markers (both morphological and molecular), would be extremely useful in this context. Here we have addressed species delimitation of sympatric tropical tree species that belong to Carapa spp. (Meliaceae) in Central Africa. We adopted a population genetics approach, sampling numerous individuals from three locations where sympatric Carapa species are known to exist. Comparisons between morphological markers (the presence or absence of characters, leaf-shape traits) and molecular markers (chloroplast sequences, ribosomal internal transcribed spacer region (ITS) sequences, and nuclear microsatellites) demonstrated the following: (i) a strong correlation between morphological and nuclear markers; (ii) despite substantial polymorphism, the inability of chloroplast DNA to discriminate between species, suggesting that cytoplasmic markers represent ineffective DNA barcodes; (iii) lineage sorting effects when using ITS sequences; and (iv) a complex evolutionary history within the genus Carapa, which includes frequent inter-specific gene flow. Our results support the use of a population genetics approach, based on ultra-polymorphic markers, to address species delimitation within complex taxonomic groups.

A new species of Cassipourea is described from the banks of the Mana River, in the southeastern boundary of the Korup National Park in Cameroon and placed provisionally within the subgenus Cassipourea. The leaves of the new species are close to those of C. afzelii, but its flowers differ from those of the latter and the rest of the species of the subgenus Cassipourea in having a glabrous ovary and a diplostemonous androceum, with filaments distinctly of two lengths. The illustration of the new species is provided and based on its narrow distribution, the provisional conservation status Vulnerable VU D1 is assigned.