The puzzling ecology of African Marantaceae forests.

Marantaceae forests are tropical rainforests characterized by a continuous understory layer of perennial giant herbs and a near absence of tree regeneration. Although widespread in West-Central Africa, Marantaceae forests have rarely been considered in the international literature. Yet, they pose key challenges and opportunities for theoretical ecology that transcend the borders of the continent. Specifically, we ask in this review whether open Marantaceae forests and dense closed-canopy forests can be considered as one of the few documented examples of alternative stable states in tropical forests. First, we introduce the different ecological factors that have been posited to drive Marantaceae forests (climate, soil, historical and recent anthropogenic pressures, herbivores) and develop the different hypotheses that have been suggested to explain how Marantaceae forests establish in relation with other vegetation types (understory invasion, early succession after disturbance, and intermediate successional stage). Then, we review the underlying ecological mechanisms that can explain the stability of Marantaceae forests in the long term (tree recruitment inhibition, promotion of and resilience to fire, adaptive reproduction, maintenance by megaherbivores). Although some uncertainties remain and call for further empirical and theoretical research, we found converging evidence that Marantaceae forests are associated with an ecological succession that has been deflected or arrested. If verified, Marantaceae forests may provide a useful model to understand critical transitions in forest ecosystems, which is of particular relevance to achieve sustainable forest management and mitigate global climate change.

Tallo: A global tree allometry and crown architecture database.

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

Resistance of African tropical forests to an extreme climate anomaly.

The responses of tropical forests to environmental change are critical uncertainties in predicting the future impacts of climate change. The positive phase of the 2015-2016 El Niño Southern Oscillation resulted in unprecedented heat and low precipitation in the tropics with substantial impacts on the global carbon cycle. The role of African tropical forests is uncertain as their responses to short-term drought and temperature anomalies have yet to be determined using on-the-ground measurements. African tropical forests may be particularly sensitive because they exist in relatively dry conditions compared with Amazonian or Asian forests, or they may be more resistant because of an abundance of drought-adapted species. Here, we report responses of structurally intact old-growth lowland tropical forests inventoried within the African Tropical Rainforest Observatory Network (AfriTRON). We use 100 long-term inventory plots from six countries each measured at least twice prior to and once following the 2015-2016 El Niño event. These plots experienced the highest temperatures and driest conditions on record. The record temperature did not significantly reduce carbon gains from tree growth or significantly increase carbon losses from tree mortality, but the record drought did significantly decrease net carbon uptake. Overall, the long-term biomass increase of these forests was reduced due to the El Niño event, but these plots remained a live biomass carbon sink (0.51 ± 0.40 Mg C ha-1 y-1) despite extreme environmental conditions. Our analyses, while limited to African tropical forests, suggest they may be more resistant to climatic extremes than Amazonian and Asian forests.

Unveiling African rainforest composition and vulnerability to global change.

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

Daily Activity Patterns and Co-Occurrence of Duikers Revealed by an Intensive Camera Trap Survey across Central African Rainforests.

The duiker community in Central African rainforests includes a diversity of species that can coexist in the same area. The study of their activity patterns is needed to better understand habitat use or association between the species. Using camera traps, we studied the temporal activity patterns, and quantified for the first time the temporal overlap and spatial co-occurrence between species. Our results show that: (i) Two species are strongly diurnal: Cephalophus leucogaster, and Philantomba congica, (ii) two species are mostly diurnal: C.callipygus and C. nigrifrons, (iii) one species is strongly nocturnal: C.castaneus, (iv) and one species is mostly nocturnal: C.silvicultor. Analyses of temporal activities (for five species) identified four species pairs that highly overlapped (Δ^≥ 0.80), and six pairs that weakly overlapped (Δ^ between 0.06 and 0.35). Finally, co-occurrence tests reveal a truly random co-occurrence (plt > 0.05 and pgt > 0.05) for six species pairs, and a positive co-occurrence (pgt < 0.05) for four pairs. Positive co-occurrences are particularly noted for pairs formed by C.callipygus with the other species (except C. nigrifrons). These results are essential for a better understanding of the coexistence of duikers and the ecology of poorly known species (C. leucogaster and C. nigrifrons), and provide clarification on the activity patterns of C. silvicultor which was subject to controversy. Camera traps proved then to be a powerful tool for studying the activity patterns of free-ranging duiker populations.

Miocene Diversification in the Savannahs Precedes Tetraploid Rainforest Radiation in the African Tree Genus Afzelia (Detarioideae, Fabaceae).

The dating of diversification events, including transitions between biomes, is key to elucidate the processes that underlie the assembly and evolution of tropical biodiversity. Afzelia is a widespread genus of tropical trees, threatened by exploitation for its valuable timber, that presents an interesting system to investigate diversification events in Africa. Africa hosts diploid Afzelia species in the savannahs north and south of the Guineo-Congolian rainforest and autotetraploid species confined to the rainforest. Species delimitation and phylogenetic relationships among the diploid and tetraploid species remained unresolved in previous studies using small amounts of DNA sequence data. We used genotyping-by-sequencing in the five widespread Afzelia species in Africa, the savannah species A. africana and A. quanzensis and the rainforest species A. bipindensis, A. pachyloba, and A. bella. Maximum likelihood and coalescent approaches resolved all species as monophyletic and placed the savannah and rainforest taxa into two separate clades corresponding to contrasted ploidy levels. Our data are thus compatible with a single biome shift in Afzelia in Africa, although we were unable to conclude on its direction. SNAPP calibrated species trees show that the savannah diploids started to diversify early, at 12 (9.09-14.89) Ma, which contrasts with a recent and rapid diversification of the rainforest tetraploid clade, starting at 4.22 (3.12 - 5.36) Ma. This finding of older diversification in a tropical savannah clade vs. its sister rainforest clade is exceptional; it stands in opposition to the predominant observation of young ages for savannahs lineages in tropical regions during the relatively recent expansion of the savannah biome.

A map of African humid tropical forest aboveground biomass derived from management inventories.

Forest biomass is key in Earth carbon cycle and climate system, and thus under intense scrutiny in the context of international climate change mitigation initiatives (e.g. REDD+). In tropical forests, the spatial distribution of aboveground biomass (AGB) remains, however, highly uncertain. There is increasing recognition that progress is strongly limited by the lack of field observations over large and remote areas. Here, we introduce the Congo basin Forests AGB (CoFor-AGB) dataset that contains AGB estimations and associated uncertainty for 59,857 1-km pixels aggregated from nearly 100,000 ha of in situ forest management inventories for the 2000 - early 2010s period in five central African countries. A comprehensive error propagation scheme suggests that the uncertainty on AGB estimations derived from c. 0.5-ha inventory plots (8.6-15.0%) is only moderately higher than the error obtained from scientific sampling plots (8.3%). CoFor-AGB provides the first large scale view of forest AGB spatial variation from field data in central Africa, the second largest continuous tropical forest domain of the world.

Asynchronous carbon sink saturation in African and Amazonian tropical forests.

Structurally intact tropical forests sequestered about half of the global terrestrial carbon uptake over the 1990s and early 2000s, removing about 15 per cent of anthropogenic carbon dioxide emissions1-3. Climate-driven vegetation models typically predict that this tropical forest 'carbon sink' will continue for decades4,5. Here we assess trends in the carbon sink using 244 structurally intact African tropical forests spanning 11 countries, compare them with 321 published plots from Amazonia and investigate the underlying drivers of the trends. The carbon sink in live aboveground biomass in intact African tropical forests has been stable for the three decades to 2015, at 0.66 tonnes of carbon per hectare per year (95 per cent confidence interval 0.53-0.79), in contrast to the long-term decline in Amazonian forests6. Therefore the carbon sink responses of Earth's two largest expanses of tropical forest have diverged. The difference is largely driven by carbon losses from tree mortality, with no detectable multi-decadal trend in Africa and a long-term increase in Amazonia. Both continents show increasing tree growth, consistent with the expected net effect of rising atmospheric carbon dioxide and air temperature7-9. Despite the past stability of the African carbon sink, our most intensively monitored plots suggest a post-2010 increase in carbon losses, delayed compared to Amazonia, indicating asynchronous carbon sink saturation on the two continents. A statistical model including carbon dioxide, temperature, drought and forest dynamics accounts for the observed trends and indicates a long-term future decline in the African sink, whereas the Amazonian sink continues to weaken rapidly. Overall, the uptake of carbon into Earth's intact tropical forests peaked in the 1990s. Given that the global terrestrial carbon sink is increasing in size, independent observations indicating greater recent carbon uptake into the Northern Hemisphere landmass10 reinforce our conclusion that the intact tropical forest carbon sink has already peaked. This saturation and ongoing decline of the tropical forest carbon sink has consequences for policies intended to stabilize Earth's climate.

Population genomics of the widespread African savannah trees Afzelia africana and Afzelia quanzensis reveals no significant past fragmentation of their distribution ranges.

PREMISE: Few studies have addressed the evolutionary history of tree species from African savannahs. Afzelia contains economically important timber species, including two species widely distributed in African savannahs: A. africana in the Sudanian region and A. quanzensis in the Zambezian region. We aimed to infer whether these species underwent range fragmentation and/or demographic changes, possibly reflecting how savannahs responded to Quaternary climate changes. METHODS: We characterized the genetic diversity and structure of these species across their distribution ranges using nuclear microsatellites (SSRs) and genotyping-by-sequencing (GBS) markers. Six SSR loci were genotyped in 241 A. africana and 113 A. quanzensis individuals, while 2800 high-quality single nucleotide polymorphisms (SNPs) were identified in 30 A. africana individuals. RESULTS: Both species appeared to be mainly outcrossing. The kinship between individuals decayed with the logarithm of the distance at similar rates across species and markers, leading to relatively small Sp statistics (0.0056 for SSR and 0.0054 for SNP in A. africana, 0.0075 for SSR in A. quanzensis). The patterns were consistent with isolation by distance expectations in the absence of large-scale geographic gradients. Bayesian clustering of SSR genotypes did not detect genetic clusters within species. In contrast, SNP data resolved intraspecific genetic clusters in A. africana, illustrating the higher resolving power of GBS. However, these clusters revealed low levels of differentiation and no clear geographical entities, so that they were interpreted as resulting from the isolation by distance pattern rather than from past population fragmentation. CONCLUSIONS: These results suggest that populations have remained connected throughout the large, continuous savannah landscapes. The absence of clear phylogeographic discontinuities, also found in a few other African savannah trees, indicates that their distribution ranges have not been significantly fragmented during the climatic oscillations of the Pleistocene, in contrast to patterns commonly found in African rainforest trees.

Highlighting convergent evolution in morphological traits in response to climatic gradient in African tropical tree species: The case of genus Guibourtia Benn.

Adaptive evolution is a major driver of organism diversification, but the links between phenotypic traits and environmental niche remain little documented in tropical trees. Moreover, trait-niche relationships are complex because a correlation between the traits and environmental niches displayed by a sample of species may result from (a) convergent evolution if different environmental conditions have selected different sets of traits, and/or (b) phylogenetic inertia if niche and morphological differences between species are simply function of their phylogenetic divergence, in which case the trait-niche correlation does not imply any direct causal link. Here, we aim to assess the respective roles of phylogenetic inertia and convergent evolution in shaping the differences of botanical traits and environmental niches among congeneric African tree species that evolved in different biomes.This issue was addressed with the tree genus Guibourtia Benn. (Leguminosae and Detarioideae), which contains 13 African species occupying various forest habitat types, from rain forest to dry woodlands, with different climate and soil conditions. To this end, we combined morphological data with ecological niche modelling and used a highly resolved plastid phylogeny of the 13 African Guibourtia species.First, we demonstrated phylogenetic signals in both morphological traits (Mantel test between phylogenetic and morphological distances between species: r = .24, p = .031) and environmental niches (Mantel test between phylogenetic and niche distances between species: r = .23, p = .025). Second, we found a significant correlation between morphology and niche, at least between some of their respective dimensions (Mantel's r = .32, p = .013), even after accounting for phylogenetic inertia (Phylogenetic Independent Contrast: r = .69, p = .018). This correlation occurred between some leaflet and flower traits and solar radiation, relative humidity, precipitations, and temperature range.Our results demonstrate the convergent evolution of some morphological traits in response to climatic factors in congeneric tree species and highlight the action of selective forces, along with neutral ones, in shaping the divergence between congeneric tropical plants.

Seed and pollen dispersal distances in two African legume timber trees and their reproductive potential under selective logging.

The natural regeneration of tree species depends on seed and pollen dispersal. To assess whether limited dispersal could be critical for the sustainability of selective logging practices, we performed parentage analyses in two Central African legume canopy species displaying contrasted floral and fruit traits: Distemonanthus benthamianus and Erythrophleum suaveolens. We also developed new tools linking forward dispersal kernels with backward migration rates to better characterize long-distance dispersal. Much longer pollen dispersal in D. benthamianus (mean distance dp  = 700 m, mp  = 52% immigration rate in 6 km2 plot, s = 7% selfing rate) than in E. suaveolens (dp  = 294 m, mp  = 22% in 2 km2 plot, s = 20%) might reflect different insect pollinators. At a local scale, secondary seed dispersal by vertebrates led to larger seed dispersal distances in the barochorous E. suaveolens (ds  = 175 m) than in the wind-dispersed D. benthamianus (ds  = 71 m). Yet, seed dispersal appeared much more fat-tailed in the latter species (15%-25% seeds dispersing >500 m), putatively due to storm winds (papery pods). The reproductive success was correlated to trunk diameter in E. suaveolens and crown dominance in D. benthamianus. Contrary to D. benthamianus, E. suaveolens underwent significant assortative mating, increasing further the already high inbreeding of its juveniles due to selfing, which seems offset by strong inbreeding depression. To achieve sustainable exploitation, seed and pollen dispersal distances did not appear limiting, but the natural regeneration of E. suaveolens might become insufficient if all trees above the minimum legal cutting diameter were exploited. This highlights the importance of assessing the diameter structure of reproductive trees for logged species.

Do topography and fruit presence influence occurrence and intensity of crop-raiding by forest elephants (Loxodonta africana cyclotis)?

Crop damage by forest elephants (Loxodonta africana cyclotis) and the resulting human-elephant conflict are issues of great concern for both the conservation of the species and the protection of rural livelihoods in Central Africa. Addressing these problems requires identifying the factors that facilitate or impede crop-raiding by forest elephants. Yet to date, the environmental or anthropogenic factors that influence the occurrence and intensity of crop-raiding by forest elephants are largely unknown. We used a multivariate approach to investigate conditions under which forest elephants raid some fields and not others in the buffer zone of Monts de Cristal National Park (MCNP), Gabon. We first interviewed 121 farmers from 11 villages situated within 10 km of MCNP regarding the occurrence of elephant crop-raiding of their fields. We then collected data on 39 explanatory variables to characterize the agricultural fields. Of these, the most important predictors of elephant raid occurrence of crop damage were presence of fruit trees, elephant deterrents (scarecrows, fire, wire string fences and empty barrels), and field topography. We secondly assessed the effect of stage of crop growth, presence of fruit trees, field topography and presence of elephant deterrents on crop-raiding occurrence and intensity by counting raids and measuring areas of crop damage every week in 17 plantations over 19 weeks in the most elephant-impacted zone of the study area. We found that fruit presence and stage of crop growth led to more intense damage to crops, whereas local deterrents did not inhibit raiding events and crop damage by elephants. We report a tradeoff between non-timber forest products (NTFP) services and crop-raiding by elephants. We show for the first time that steep topography impedes elephant damage to crops with no raids recorded in fields with surrounding slopes greater than 25%. We discuss whether farming on steep fields could be used as a strategy for mitigating crop-raiding to favor human-elephant coexistence and enhance elephant conservation.

The persistence of carbon in the African forest understory.

Quantifying carbon dynamics in forests is critical for understanding their role in long-term climate regulation1-4. Yet little is known about tree longevity in tropical forests3,5-8, a factor that is vital for estimating carbon persistence3,4. Here we calculate mean carbon age (the period that carbon is fixed in trees7) in different strata of African tropical forests using (1) growth-ring records with a unique timestamp accurately demarcating 66 years of growth in one site and (2) measurements of diameter increments from the African Tropical Rainforest Observation Network (23 sites). We find that in spite of their much smaller size, in understory trees mean carbon age (74 years) is greater than in sub-canopy (54 years) and canopy (57 years) trees and similar to carbon age in emergent trees (66 years). The remarkable carbon longevity in the understory results from slow and aperiodic growth as an adaptation to limited resource availability9-11. Our analysis also reveals that while the understory represents a small share (11%) of the carbon stock12,13, it contributes disproportionally to the forest carbon sink (20%). We conclude that accounting for the diversity of carbon age and carbon sequestration among different forest strata is critical for effective conservation management14-16 and for accurate modelling of carbon cycling4.

The limited contribution of large trees to annual biomass production in an old-growth tropical forest.

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

Evolution in the Amphi-Atlantic tropical genus Guibourtia (Fabaceae, Detarioideae), combining NGS phylogeny and morphology.

Tropical rain forests support a remarkable diversity of tree species, questioning how and when this diversity arose. The genus Guibourtia (Fabaceae, Detarioideae), characterized by two South American and 13 African tree species growing in various tropical biomes, is an interesting model to address the role of biogeographic processes and adaptation to contrasted environments on species diversification. Combining whole plastid genome sequencing and morphological characters analysis, we studied the timing of speciation and diversification processes in Guibourtia through molecular dating and ancestral habitats reconstruction. All species except G. demeusei and G. copallifera appear monophyletic. Dispersal from Africa to America across the Atlantic Ocean is the most plausible hypothesis to explain the occurrence of Neotropical Guibourtia species, which diverged ca. 11.8 Ma from their closest African relatives. The diversification of the three main clades of African Guibourtia is concomitant to Miocene global climate changes, highlighting pre-Quaternary speciation events. These clades differ by their reproductive characters, which validates the three subgenera previously described: Pseudocopaiva, Guibourtia and Gorskia. Within most monophyletic species, plastid lineages start diverging from each other during the Pliocene or early Pleistocene, suggesting that these species already arose during this period. The multiple transitions between rain forests and dry forests/savannahs inferred here through the plastid phylogeny in each Guibourtia subgenus address thus new questions about the role of phylogenetic relationships in shaping ecological niche and morphological similarity among taxa.

History of the fragmentation of the African rain forest in the Dahomey Gap: insight from the demographic history of Terminalia superba.

Paleo-environmental reconstructions show that the distribution of tropical African rain forests was affected by Quaternary climate changes. They suggest that the Dahomey Gap (DG)-the savanna corridor that currently separates Upper Guinean (UG, West Africa) and Lower Guinean (LG, western Central Africa) rain forest blocks-was forested during the African Humid Holocene period (from at least 9 ka till 4.5 ka), and possibly during other interglacial periods, while an open vegetation developed in the DG under drier conditions, notably during glacial maxima. Nowadays, relics of semi-deciduous forests containing UG and LG forest species are still present within the DG. We used one of these species, the pioneer tree Terminalia superba (Combretaceae), to study past forest fragmentation in the DG and its impact on infraspecific biodiversity. A Bayesian clustering analysis of 299 individuals genotyped at 14 nuclear microsatellites revealed five parapatric genetic clusters (UG, DG, and three in LG) with low to moderate genetic differentiation (Fst from 0.02 to 0.24). Approximate Bayesian Computation analyses inferred a demographic bottleneck around the penultimate glacial period in all populations. They also supported an origin of the DG population by admixture of UG and LG populations around 54,000 (27,600-161,000) years BP, thus before the Last Glacial Maximum. These results contrast with those obtained on Distemonanthus benthamianus where the DG population seems to originate from the Humid Holocene period. We discuss these differences in light of the ecology of each species. Our results challenge the simplistic view linking population fragmentation/expansion with glacial/interglacial periods in African forest species.

Characterization of microsatellite markers in the African tropical tree species Guibourtia ehie (Fabaceae, Detarioideae).

PREMISE OF THE STUDY: Microsatellite primers (simple sequence repeats [SSRs]) were developed in Guibourtia ehie (Fabaceae, Detarioideae) to study population genetic structure and the history of African vegetation. METHODS AND RESULTS: We isolated 18 polymorphic SSRs from a nonenriched genomic library. This set of primer pairs was tested on four populations, and the results showed two to 16 alleles per locus with mean observed and expected heterozygosities of 0.27 ± 0.05 and 0.57 ± 0.05, respectively. Cross-amplification tests in 13 congeneric species were successful for the four taxa belonging to the subgenus Gorskia. CONCLUSIONS: This set of microsatellite markers will be useful to investigate the phylogeography and population genetics of G. ehie, a key representative of African semideciduous moist forests.

Extensive seed and pollen dispersal and assortative mating in the rain forest tree Entandrophragma cylindricum (Meliaceae) inferred from indirect and direct analyses.

Pollen and seed dispersal are key processes affecting the demographic and evolutionary dynamics of plant species and are also important considerations for the sustainable management of timber trees. Through direct and indirect genetic analyses, we studied the mating system and the extent of pollen and seed dispersal in an economically important timber species, Entandrophragma cylindricum (Meliaceae). We genotyped adult trees, seeds and saplings from a 400-ha study plot in a natural forest from East Cameroon using eight nuclear microsatellite markers. The species is mainly outcrossed (t = 0.92), but seeds from the same fruit are often pollinated by the same father (correlated paternity, rp  = 0.77). An average of 4.76 effective pollen donors (Nep ) per seed tree contributes to the pollination. Seed dispersal was as extensive as pollen dispersal, with a mean dispersal distance in the study plot approaching 600 m, and immigration rates from outside the plot to the central part of the plot reaching 40% for both pollen and seeds. Extensive pollen- and seed-mediated gene flow is further supported by the weak, fine-scale spatial genetic structure (Sp statistic = 0.0058), corresponding to historical gene dispersal distances (σg ) reaching approximately 1,500 m. Using an original approach, we showed that the relatedness between mating individuals (Fij  = 0.06) was higher than expected by chance, given the extent of pollen dispersal distances (expected Fij  = 0.02 according to simulations). This remarkable pattern of assortative mating could be a phenomenon of potentially consequential evolutionary and management significance that deserves to be studied in other plant populations.

Diversity and carbon storage across the tropical forest biome.

Tropical forests are global centres of biodiversity and carbon storage. Many tropical countries aspire to protect forest to fulfil biodiversity and climate mitigation policy targets, but the conservation strategies needed to achieve these two functions depend critically on the tropical forest tree diversity-carbon storage relationship. Assessing this relationship is challenging due to the scarcity of inventories where carbon stocks in aboveground biomass and species identifications have been simultaneously and robustly quantified. Here, we compile a unique pan-tropical dataset of 360 plots located in structurally intact old-growth closed-canopy forest, surveyed using standardised methods, allowing a multi-scale evaluation of diversity-carbon relationships in tropical forests. Diversity-carbon relationships among all plots at 1 ha scale across the tropics are absent, and within continents are either weak (Asia) or absent (Amazonia, Africa). A weak positive relationship is detectable within 1 ha plots, indicating that diversity effects in tropical forests may be scale dependent. The absence of clear diversity-carbon relationships at scales relevant to conservation planning means that carbon-centred conservation strategies will inevitably miss many high diversity ecosystems. As tropical forests can have any combination of tree diversity and carbon stocks both require explicit consideration when optimising policies to manage tropical carbon and biodiversity.

Present-day central African forest is a legacy of the 19th century human history.

The populations of light-demanding trees that dominate the canopy of central African forests are now aging. Here, we show that the lack of regeneration of these populations began ca. 165 ya (around 1850) after major anthropogenic disturbances ceased. Since 1885, less itinerancy and disturbance in the forest has occurred because the colonial administrations concentrated people and villages along the primary communication axes. Local populations formerly gardened the forest by creating scattered openings, which were sufficiently large for the establishment of light-demanding trees. Currently, common logging operations do not create suitable openings for the regeneration of these species, whereas deforestation degrades landscapes. Using an interdisciplinary approach, which included paleoecological, archaeological, historical, and dendrological data, we highlight the long-term history of human activities across central African forests and assess the contribution of these activities to present-day forest structure and composition. The conclusions of this sobering analysis present challenges to current silvicultural practices and to those of the future.