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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.

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.

When Darwin visited the Galapagos archipelago, he observed that, in spite of the islands’ physical similarity, members of species that had dispersed to them recently were beginning to diverge from each other. He postulated that these divergences must have resulted primarily from interactions with sets of other species that had also diverged across these otherwise similar islands. By extrapolation, if Darwin is correct, such complex interactions must be driving species divergences across all ecosystems. However, many current general ecological theories that predict observed distributions of species in ecosystems do not take the details of between-species interactions into account. Here we quantify, in sixteen forest diversity plots (FDPs) worldwide, highly significant negative density-dependent (NDD) components of both conspecific and heterospecific between-tree interactions that affect the trees’ distributions, growth, recruitment, and mortality. These interactions decline smoothly in significance with increasing physical distance between trees. They also tend to decline in significance with increasing phylogenetic distance between the trees, but each FDP exhibits its own unique pattern of exceptions to this overall decline. Unique patterns of between-species interactions in ecosystems, of the general type that Darwin postulated, are likely to have contributed to the exceptions. We test the power of our null-model method by using a deliberately modified data set, and show that the method easily identifies the modifications. We examine how some of the exceptions, at the Wind River (USA) FDP, reveal new details of a known allelopathic effect of one of the Wind River gymnosperm species. Finally, we explore how similar analyses can be used to investigate details of many types of interactions in these complex ecosystems, and can provide clues to the evolution of these interactions., Author summary Worldwide, ecosystems are collapsing or in danger of collapse, but the precise causes of these collapses are often unknown. Observational and experimental evidence shows that all ecosystems are characterized by strong interactions between and among species, and that these webs of interactions can be important contributors to the preservation of ecosystem diversity. But many of the interactions–such as those involving pathogenic microorganisms and the chemical defenses that are mounted by their prey–are not easily identified and analyzed in ecosystems that may have hundreds or thousands of species. Here we use our equal-area-annulus analytical method to examine census data from over three million trees in forest plots from around the world. We show how the method can be used to flag pairs and groups of species that exhibit unusual levels of interaction and that are likely on further investigation to yield information about their causative mechanisms. We give a detailed example showing how some of these interactions can be traced to defense mechanisms that are possessed by one of the tree species. We explore how our method can be used to identify the between-species interactions that play the largest roles in the maintenance of ecosystems and their diversity.

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

© 2020 John Wiley & Sons Ltd Aim: Mapping tree species richness across the tropics is of great interest for effective conservation and biodiversity management. In this study, we evaluated the potential of full-waveform lidar data for mapping tree species richness across the tropics by relating measurements of vertical canopy structure, as a proxy for the occupation of vertical niche space, to tree species richness. Location: Tropics. Time period: Present. Major taxa studied: Trees. Methods: First, we evaluated the characteristics of vertical canopy structure across 15 study sites using (simulated) large-footprint full-waveform lidar data (22 m diameter) and related these findings to in-situ tree species information. Then, we developed structure–richness models at the local (within 25–50 ha plots), regional (biogeographical regions) and pan-tropical scale at three spatial resolutions (1.0, 0.25 and 0.0625 ha) using Poisson regression. Results: The results showed a weak structure–richness relationship at the local scale. At the regional scale (within a biogeographical region) a stronger relationship between canopy structure and tree species richness across different tropical forest types was found, for example across Central Africa and in South America [R2 ranging from.44–.56, root mean squared difference as a percentage of the mean (RMSD%) ranging between 23–61%]. Modelling the relationship pan-tropically, across four continents, 39% of the variation in tree species richness could be explained with canopy structure alone (R2 =.39 and RMSD% = 43%, 0.25-ha resolution). Main conclusions: Our results may serve as a basis for the future development of a set of structure–richness models to map high resolution tree species richness using vertical canopy structure information from the Global Ecosystem Dynamics Investigation (GEDI). The value of this effort would be enhanced by access to a larger set of field reference data for all tropical regions. Future research could also support the use of GEDI data in frameworks using environmental and spectral information for modelling tree species richness across the tropics.

© 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.

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.

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.

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.

Background: The Rumpi Hills Forest Reserve (RHFR) is a montane forest area in south-western Cameroon. Although RHFR is presumed to be rich in biodiversity and vegetation types, little information exists regarding its floristic composition and vegetation patterns. Objectives: Our goal was to characterise vegetation patterns in the reserve and to understand how elevation influences distributions and diversity of species. We aimed to provide a first detailed plant species inventory for this important forest area, as well as basic information on forest structure. Method: We characterised floristic composition and vegetation patterns of the reserve in 25 1-ha plots along an elevational gradient from 50 m to 1778 m. In each plot, trees and lianas of diameter at breast height (dbh) ≥ 10 cm were measured; shrubs < 10 cm were measured in nested plots of 0.01 ha. Results: In all, 16 761 trees, shrubs and lianas with dbh ≥ 1 cm were censused, representing 71 families, 279 genera and 617 morphospecies. Floristic composition ranged from 94 to 132 species, with a mean of 117.5 species per hectare in lowland forest (50 m – 200 m) and 36–41 species, with a mean of 38.5 species per hectare in montane cloud forest (1600 m – 1778 m) near the summit of Mount Rata. Two-way indicator species analysis classified the 25 plots into six vegetation types corresponding to lowland evergreen rainforest, lowland evergreen rainforest on basalt rocks, middle-elevation evergreen forest, submontane forest, transitional submontane forest and montane cloud forest. In all, 0.006% of the reserve was included in our sample plots. Detrended correspondence analysis highlighted the importance of elevation in shaping vegetation patterns. Conclusion: The RHFR is composed of different vegetation types, which show impressive variation in terms of structure, species composition and diversity. The detailed, fine-scale inventory data obtained in this study could be useful in planning efficient management of this and other montane tropical forests.

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”.

© 2018 John Wiley & Sons Ltd 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.

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

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.

The Neotropics, Afrotropics and Madagascar have different histories which have influenced their respective patterns of diversity. Based on current knowledge of these histories, we developed the following predictions about the phylogenetic structure and composition of rainforest tree communities: (Hypothesis 1) isolation of Gondwanan biotas generated differences in phylogenetic composition among biogeographical regions; (H2) major Cenozoic extinction events led to lack of phylogenetic structure in Afrotropical and Malagasy communities; (H3) greater angiosperm diversification in the Neotropics led to greater phylogenetic clustering there than elsewhere; (H4) phylogenetic overdispersion is expected near the Andes due to the co-occurrence of magnoliids tracking conserved habitat preferences and recently diversified eudicot lineages. Using abundance data of tropical rainforest tree species from 94 communities in the Neotropics, Afrotropics and Madagascar, we computed net relatedness index (NRI) to assess local phylogenetic structure, i.e. phylogenetic clustering vs. overdispersion relative to regional species pools, and principal coordinates of phylogenetic structure (PCPS) to assess variation in phylogenetic composition across communities. We observed significant differences in phylogenetic composition among biogeographical regions (agreement with H1). Overall phylogenetic structure did not differ among biogeographical regions, but results indicated variation from Andes to Amazon. We found widespread phylogenetic randomness in most Afrotropical and all Malagasy communities (agreement with H2). Most of central Amazonian communities were phylogenetically random, although some communities presented phylogenetic clustering (partial agreement with H3). We observed phylogenetic overdispersion near the Andes (agreement with H4). We were able to identify how differences in lineage composition are related to local phylogenetic co-occurrences across biogeographical regions that have been undergoing different climatic and orographic histories during the past 100 Myr. We observed imprints of the history following Gondwana breakup on phylobetadiversity and local phylogenetic structure of rainforest tree communities in the Neotropics, Afrotropics and Madagascar. © 2016 The Authors

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.

Carapa vasquezii from western Amazonia is described and illustrated. The new species differs from the closely related C. guianensis in having leaflets with an obtuse to rounded apex and a farinose midrib, farinose inflorescence and flower pedicels, 6-ovulate locules, and smaller seeds with a very much reduced hilum.

A taxonomic revision of Carapa (Meliaceae) in Ecuador is provided with the recognition of four species, C. megistocarpa which has cauliflorous inflorescences, C. nicaraguensis previously described and currently placed as synonym of C. guianensis and two new species (C. alticola and C. longipetala). The new species are close to C. guianensis based on their 4-merous flowers borne at the end of the branches. However, C. alticola differs from C. guianensis in having larger leaflets with prominent secondary veins, seeds with rounded edges and short poorly ramified inflorescences, while C. longipetala can be distinguised from C. guianensis in having distinctly pedicellate flowers and 6-ovulate ovary loculi. The new species are described, illustrated, and a key to the four species recognized in Ecuador is provided.

Our aim was to evaluate the extent to which we can predict and map tree alpha diversity across broad spatial scales either by using climate and remote sensing data or by exploiting spatial autocorrelation patterns in tropical rain forest, West Africa and Atlantic Central Africa.

The taxonomy of the amphi-Atlantic tree genus Carapa (Meliaceae) has long been controversial. Of the three species currently recognized in the genus, two are known to present substantial morphological variation that has been used in the past to distinguish several taxa, most of which are currently placed in synonymy. Here, a combination of field observations, univariate analyses of leaf, floral and seed characters and principal coordinate analyses of floral characters in the context of a molecular phylogenetic analysis was used to investigate the patterns of variation and delimit morphological species anew in the genus. These results support the recognition of 27 species in Carapa, of which 16 are previously described and 11 are new. In general, phylogenetically related species occurred in the same geographical area, but were morphologically distinct. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165, 186–221.

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

Species spatial turnover, or β-diversity, induces a decay of community similarity with geographic distance known as the distance–decay relationship. Although this relationship is central to biodiversity and biogeography, its theoretical underpinnings remain poorly understood. Here, we develop a general framework to describe how the distance–decay relationship is influenced by population aggregation and the landscape-scale species-abundance distribution. We utilize this general framework and data from three tropical forests to show that rare species have a weak influence on distance–decay curves, and that overall similarity and rates of decay are primarily influenced by species abundances and population aggregation respectively. We illustrate the utility of the framework by deriving an exact analytical expression of the distance–decay relationship when population aggregation is characterized by the Poisson Cluster Process. Our study provides a foundation for understanding the distance–decay relationship, and for predicting and testing patterns of beta-diversity under competing theories in ecology. Ecology Letters (2008) 11: 904–917

A distinctive new species of Rubiaceae from Rio Muni, Equatorial Guinea, is described and illustrated. Leptactina rheophytica is the only rheophyte known in the genus. Its diagnostic characters are elucidated, its taxonomic affinities are discussed, and notes on its conservation status are provided. © 2007 The Linnean Society of London.

A new species of Triuridaceae, Kihansia jengiensis Sainge & Kenfack is described from the semi-deciduous forest of Southeastern Cameroon. The new species is illustrated and a key to the identification of the two species in the genus provided. The species constitutes the first record of the genus in central Africa and its conservation status is assessed as Critically Endangered.

We censused all trees ‡1 cm dbh in 50 ha of forest in Korup National Park, southwest Cameroon, in the central African coastal forest known for high diversity and endemism. The plot included 329,519 individuals and 493 species, but 128 of those taxa remain partially identified. Abundance varied over four orders of magnitude, from 1 individual per 50 ha (34 species) to Phyllobotryon spathulatum, with 26,741 trees; basal area varied over six orders of magnitude. Abundance patterns, both the percentage of rare species and the dominance of abundant species were similar to those from 50-ha plots censused the same way in Asia and Latin America. Rare species in the Korup plot were much less likely to be identified than common species: 42% of taxa with

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

The new species Cassipourea korupensis Kenfack & Sainge (Rhizophoraceae) from western Cameroon is described and illustrated, and its conservation status discussed. Unlike other species of subgenus Lasiosepalum Alston, in which it is provisionally placed, the new species has a distinctive glabrous ovary and a cupuliform calyx.

A recent increase in published studies of lianas has been paralleled by a proliferation of protocols for censusing lianas. This article seeks to increase uniformity in liana inventories by providing specific recommendations for the determination of which taxa to include, the location of diameter measurement points on individual stems, the setting of minimum stem diameter cutoffs, the treatment of multiple-stemmed and rooted clonal groups, and the measurement of noncylindrical stems. Use of more uniform liana censusing protocols may facilitate comparison of independently collected data sets and further our understanding of global patterns in liana abundance, diversity, biomass, and dynamics.

Background: Many tree species in tropical forests have distributions tracking local ridge-slope-valley topography. Previous work in a 50-ha plot in Korup National Park, Cameroon, demonstrated that 272 species, or 63% of those tested, were significantly associated with topography. Methods: We used two censuses of 329,000 trees ≥1 cm dbh to examine demographic variation at this site that would account for those observed habitat preferences. We tested two predictions. First, within a given topographic habitat, species specializing on that habitat (‘residents’) should outperform species that are specialists of other habitats (‘foreigners’). Second, across different topographic habitats, species should perform best in the habitat on which they specialize (‘home’) compared to other habitats (‘away’). Species’ performance was estimated using growth and mortality rates. Results: In hierarchical models with species identity as a random effect, we found no evidence of a demographic advantage to resident species. Indeed, growth rates were most often higher for foreign species. Similarly, comparisons of species on their home vs. away habitats revealed no sign of a performance advantage on the home habitat. Conclusions: We reject the hypothesis that species distributions along a ridge-valley catena at Korup are caused by species differences in trees ≥1 cm dbh. Since there must be a demographic cause for habitat specialization, we offer three alternatives. First, the demographic advantage specialists have at home occurs at the reproductive or seedling stage, in sizes smaller than we census in the forest plot. Second, species may have higher performance on their preferred habitat when density is low, but when population builds up, there are negative density-dependent feedbacks that reduce performance. Third, demographic filtering may be produced by extreme environmental conditions that we did not observe during the census interval.

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

Tetagastris panamensis is a tropical forest canopy tree that is broadly distributed across Central and South America. We isolated 15 microsatellite loci and characterized them in populations from Panama and Ecuador. The number of alleles per locus in 61 individuals ranged from 4 to 22, and expected heterozygosity ranged from 0.044 to 0.909. The high overall single-parent exclusion probability (P = 1.00) highlights the utility of these loci for parentage-based analyses of gene flow.

Carapa akuri, a new species endemic to central Guyana, is described and illustrated. It is compared to the two other species (C. guianensis and C. surinamensis) occurring in the Guianas.

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

Between 1990 and 2000 a number of vegetation surveys were undertaken on the Mount Cameroon area using mainly three methods: 0.25 ha sample plots, 20 m wide transects and rapid botanical surveys. A comparison of these methods in terms of data collected, operational characteristics and output relevance, revealed their strengths and weaknesses. The methods vary markedly in their ecological value (0.25 ha sample plots superior), taxonomic scope and rigour (rapid botanical survey superior) and logistic efficiency (transects superior). The combined outcome of all these surveys is discussed and recommendations made on the use of the rapid botanical survey in order to rapidly fill the remaining gaps on plant distribution.

During the past decade, two long-term studies of tropical forest trees and lianas were initiated in the Korup National Park, Cameroon and in the Reserve de Faune 'a Okapis, Democratic Republic of Congo. These two large forest dynamics programs are part of a global network of research sites coordinated by the Smithsonian Tropical Research Institute, and are leading to the discovery of many new plant species (e.g., Gereau & Kenfack 2000; Ewango & Breteler 2001; Sonk6 et al. 2002). During the study of the Korup plot vouchers at BR, MO, K, SCA and YA, some specimens belonging to the genus Manilkara Adans. could not be matched, and did not fit any of the species described from western and central Africa (Aubreville 1961, 1964; Gautier 1997). Critical examination of the Korup specimens revealed sufficient differences to justify the description of a new species. Further surveys in the Korup area to collect additional specimens of this taxon have so far been unsuccessful. This description is therefore based on a very small number of individuals, and we have little data on intra-specific variation. However, since we have both flowering and fruiting material showing a suite of distinctive characters, there is no reason to delay publication. The new species clearly belongs in the genus Manilkara because of the calyx, which has two whorls of 3 sepals each. Although Manilkara species are often difficult to separate, ours does not resemble any other described species very closely. M. pellegriniana Tisser. & Sillans is the only other species from the region with persistent stipules and glabrous, laciniate staminodes, and the new species can be distinguished from it by a number of well-defined characters. Manilkara lososiana Kenfack & Ewango sp. nov. Manilkarae pellegrinianae Tisser. & Sillans similis, sed stipulis triangularibus non filiformibus, petiolo breviore 5 15 non 20 40 mm longo, pedicello longiore, foliorum indumento brunneo (nec griseoargenteo) distinguitur. Typus: Cameroon, Southwest Province, Korup National Park, Kenfack 625 (holotypus YA; isotypi SCA, MO, K).