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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.

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.

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.

© 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.

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.

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.

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”.

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

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

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

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.

Defaunation is causing declines of large-seeded animal-dispersed trees in tropical forests worldwide, but whether and how these declines will affect carbon storage across this biome is unclear. Here we show, using a pan-tropical data set, that simulated declines of large-seeded animal-dispersed trees have contrasting effects on aboveground carbon stocks across Earth's tropical forests. In our simulations, African, American and South Asian forests, which have high proportions of animal-dispersed species, consistently show carbon losses (2–12%), but Southeast Asian and Australian forests, where there are more abiotically dispersed species, show little to no carbon losses or marginal gains (±1%). These patterns result primarily from changes in wood volume, and are underlain by consistent relationships in our empirical data (∼2,100 species), wherein, large-seeded animal-dispersed species are larger as adults than small-seeded animal-dispersed species, but are smaller than abiotically dispersed species. Thus, floristic differences and distinct dispersal mode–seed size–adult size combinations can drive contrasting regional responses to defaunation., Defaunation is linked to the decline of tree species that depend on large animals for seed dispersal, but it is unclear if this affects carbon storage. Here the authors show that defaunation effects on carbon storage vary across continents, driven by relationships between seed dispersal strategies and adult tree size.

Tropical forests have long been recognized for their biodiversity and ecosystem services. Despite their importance, tropical forests, and particularly those of central Africa, remain understudied. Until recently, most forest inventories in Central Africa have focused on trees ≥10 cm in diameter, even though several studies have shown that small-diameter tree population may be important to demographic rates and nutrient cycling. To determine the ecological importance of small-diameter trees in central African forests, we used data from a 25-ha permanent plot that we established in the rainforest of Gabon to study the diversity and dynamics of these forests. Within the plot, we censused 175,830 trees ≥1 cm dbh from 54 families, 192 genera, and 345 species. Average tree density was 7,026 trees/ha, basal area 31.64 m2/ha, and above-ground biomass 369.40 Mg/ha. Fabaceae, Ebenaceae and Euphorbiaceae were the most important families by basal area, density and above-ground biomass. Small-diameter trees (1 cm ≥ dbh

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.

Carapa comprises trees confined to tropical forests in Africa and America. Previous revisions of the genus based primarily on herbarium specimens recognized few variable species, but also stressed the need for further research to clarify and understand the patterns of morphological variation in these species complexes. In the present revision, 27 species are recognized in Carapa of which 16 occur in Africa and 11 in America. Nine new species are described and illustrated, and a new combination is proposed. The remaining 17 species correspond to either recently described species or previously described species that had been placed in synonymy. A key to the 27 species and their distribution maps are provided.

Niche differentiation with respect to habitat has been hypothesized to shape patterns of diversity and species distributions in plant communities. African forests have been reported to be relatively less diverse compared to highly diversed regions of the Amazonian or Southeast Asian forests, and might be expected to have less niche differentiation. We examined patterns of structural and floristic differences among five topographically defined habitats for 494 species with stems ≥1 cm dbh in a 50-ha plot in Korup National Park, Cameroon. In addition, we tested for species–habitat associations for 272 species (with more than 50 individuals in the plot) using Torus translation randomization tests. Tree density and basal area were lowest in areas with negative convexity, which contained streams or were inundated during rainy periods and highest in moist well-drained habitats. Species composition and diversity varied along the topographical gradient from low flat to ridge top habitats. The low depression and low flat habitats were characterized by high diversity and similar species composition, relative to slopes, high gullies and ridge tops. Sixty-three percent of the species evaluated showed significant positive associations with at least one of the five habitat types. The majority of associations were with low depressions (75 species) and the fewest with ridge tops (8 species). The large number of species–habitat associations and the pronounced contrast between low (valley) and elevated (ridgetop) habitats in the Korup plot shows that niche differentiation with respect to edaphic variables (e.g., soil moisture, nutrients) contributes to local scale tree species distributions and to the maintenance of diversity in African forests.

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.

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.

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.

Accurately monitoring tropical forest carbon stocks is an outstanding challenge. Allometric models that consider tree diameter, height and wood density as predictors are currently used in most tropical forest carbon studies. In particular, a pantropical biomass model has been widely used for approximately a decade, and its most recent version will certainly constitute a reference in the coming years. However, this reference model shows a systematic bias for the largest trees. Because large trees are key drivers of forest carbon stocks and dynamics, understanding the origin and the consequences of this bias is of utmost concern. In this study, we compiled a unique tree mass dataset on 673 trees measured in five tropical countries (101 trees > 100 cm in diameter) and an original dataset of 130 forest plots (1 ha) from central Africa to quantify the error of biomass allometric models at the individual and plot levels when explicitly accounting or not accounting for crown mass variations. We first showed that the proportion of crown to total tree aboveground biomass is highly variable among trees, ranging from 3 to 88 %. This proportion was constant on average for trees < 10 Mg (mean of 34 %) but, above this threshold, increased sharply with tree mass and exceeded 50 % on average for trees ≥ 45 Mg. This increase coincided with a progressive deviation between the pantropical biomass model estimations and actual tree mass. Accounting for a crown mass proxy in a newly developed model consistently removed the bias observed for large trees (> 1 Mg) and reduced the range of plot-level error from −23–16 to 0–10 %. The disproportionally higher allocation of large trees to crown mass may thus explain the bias observed recently in the reference pantropical model. This bias leads to far-from-negligible, but often overlooked, systematic errors at the plot level and may be easily corrected by accounting for a crown mass proxy for the largest trees in a stand, thus suggesting that the accuracy of forest carbon estimates can be significantly improved at a minimal cost.

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.

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

The high species richness of tropical forests has long been recognized, yet there remains substantial uncertainty regarding the actual number of tropical tree species. Using a pantropical tree inventory database from closed canopy forests, consisting of 657,630 trees belonging to 11,371 species, we use a fitted value of Fisher's alpha and an approximate pantropical stem total to estimate the minimum number of tropical forest tree species to fall between ∼40,000 and ∼53,000, i.e. at the high end of previous estimates. Contrary to common assumption, the Indo-Pacific region was found to be as species-rich as the Neotropics, with both regions having a minimum of ∼19,000-25,000 tree species. Continental Africa is relatively depauperate with a minimum of ∼4,500-6,000 tree species. Very few species are shared among the African, American, and the Indo-Pacific regions. We provide a methodological framework for estimating species richness in trees that may help refine species richness estimates of tree-dependent taxa., 0, SCOPUS: ar.j, info:eu-repo/semantics/published

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.

Structural changes in secondary forests are less known in West Africa, and this precludes their management. This study aims at providing quantitative information on floristic composition and structure of the Lama secondary forests (Benin), so as to contribute to their restoration, and fill part of knowledge gaps on West African secondary forests. Data of 77 permanent plots each of 0.5 ha were used to analyze the floristic composition, the trajectory of the recovery and the recovery of stocking in these forests, compared to nearby old-growth forests. The results showed that the forests were less diversified with few species very common in the forest stands; the most dominant were Lonchocarpus sericeus and Anogeissus leiocarpa in the secondary forests, and Dialium guineense , Diospyros mespiliformis and Afzelia africana in the old-growth forests. The secondary forests hold more species than the mature ones. Their compositions will recover that of the original forest because species of the original forest were actively regenerating in the secondary forests. About 28 years after recovery, large trees were insufficient and basal area was about 60% of those of the mature forests. Further studies are needed to elucidate barriers to tree regeneration and dynamics of tree population. © 2016 International Formulae Group. All rights reserved. Keywords: Secondary forest, recovery, diversity, species composition, timber stock, Benin