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One of the current threats to the bonobo (Pan paniscus), a highly endangered ape species only found in the Democratic Republic of the Congo, are anthropozoonoses caused by human respiratory viruses. To date, epidemiological information regarding respiratory viral infections in bonobos is limited. In this study, we examined fecal immunoglobulin A antibodies against human respiratory viruses in bonobos, which may help estimating the viral prevalence. A substantial proportion of bonobos were positive for the antiviral antibodies, including those against parainfluenza virus, respiratory syncytial virus, influenza virus, rhinovirus, and mumps virus. The prevalence of the antibodies was found to depend on the viral species and bonobo populations, suggesting that the bonobos had been exposed to these respiratory viruses. These results may indicate the need for an epidemiological evidence-based action plan for the protection of bonobos from anthropozoonoses.

OBJECTIVES The little known guenon Cercopithecus dryas has a controversial taxonomic history with some recognizing two taxa (C. dryas and C. salongo) instead of one. New adult specimens from the TL2 region of the central Congo Basin allow further assessment of C. dryas morphology and, along with CT scans of the juvenile holotype, provide ontogenetically stable comparisons across all C. dryas and "C. salongo" specimens for the first time. MATERIALS AND METHODS The skins and skulls of two newly acquired C. dryas specimens, male YPM MAM 16890 and female YPM MAM 17066, were compared to previously described C. dryas and "C. salongo" specimens, along with a broader guenon comparative sample (cranial sample n = 146, dental sample n = 102). Qualitative and quantitative assessments were made on the basis of commonly noted pelage features as well as craniodental characters in the form of shape ratios and multivariate discriminant analyses. RESULTS All C. dryas specimens, including the TL2 adults, are comparatively small in overall cranial size, have relatively small I1 s, and display tall molar cusps; these osteological characters, along with pelage features, are shared with known "C. salongo" specimens. Discriminant analyses of dental features separate C. dryas/salongo specimens from all other guenons. DISCUSSION In addition to pelage-based evidence, direct osteological evidence suggests "C. salongo" is a junior synonym of C. dryas. Combined with molecular analyses suggesting C. dryas is most closely related to Chlorocebus spp., we emend the species diagnosis and support its transfer to Chlorocebus or possibly a new genus to reflect its distinctiveness.

The Endangered dryas monkey Cercopithecus dryas, endemic to the Democratic Republic of the Congo, is one of Africa's most enigmatic primates. The discovery of a dryas monkey killed by a hunter in the buffer zone of Lomami National Park in 2014 prompted field research on the species’ distribution, habitat use and stratum preference. We used local knowledge to determine the distribution of this species and to select sites for camera-trap surveys in Lomami National Park and its buffer zone. We employed a multi-strata (0–29 m) camera-trap placement technique to determine habitat use at Camp Bartho in Lomami National Park and Bafundo Forest in the Park's buffer zone. We confirmed the occurrence of the dryas monkey at seven locations over a total area of 3,453 km2, in both the Park and its buffer zone. Dryas monkeys were detected most frequently (2.22 events/100 trap-days) in disturbed areas of Bafundo Forest and less in mature forest in Camp Bartho (0.82 events/100 trap-days). Dryas monkeys appear to prefer structurally complex understories and forest edges. We found that camera traps at 2–10 m above ground over at least 365 trap-days are required to determine if the species is present. We recommend utilizing local knowledge and using this species-specific camera-trap method in other areas of the central Congo basin to determine the wider distribution of the dryas monkey.

The natural history and taxonomic status of two central African primates, Cercopithecus dryas and Cercopithecus salongo have long been in question. Recent studies confirmed that C. dryas is a basal member of the savanna monkey clade, and that it prefers dense undergrowth in lowland rainforest. While these studies advanced our knowledge of this enigmatic species, key aspects of its natural history remain poorly documented. Furthermore, the lack of a field study that documents pelage patterns of both sexes and different age classes of C. dryas has led to a disagreement over the validity of C. salongo as a sister taxon to C. dryas. Using the results of two multi-strata camera trap surveys in Lomami National Park (Democratic Republic of the Congo) and its buffer zone, we conducted a third survey in the understory of degraded forest to accumulate videos of C. dryas/salongo. We used these videos to test the hypothesis that C. dryas and C. salongo are synonymous, and to assess the species' group composition, density, behavior and vocalizations. Camera traps revealed an ontogenetic change in pelage pattern that supports the view that C. salongo is the adult of C. dryas. Videos revealed that adult males develop a blue perineum and scrotum, and a red subcaudal patch, similar to other savanna monkeys. We provide a preliminary assessment of C. dryas' group composition, density, behavior, and vocalizations. This long-overlooked monkey is an exceptional member of the Chlorocebus clade, and all aspects of its biology require further investigation.

Significance The CD4 protein of primates has undergone rapid diversification, but the reasons for this remain unknown. Here we show that within-species diversity of the HIV/simian immunodeficiency virus (SIV) envelope (Env) binding (D1) domain is common among African primate species, and that these polymorphisms can inhibit SIV Env-mediated cell entry. Amino acid replacements in the D1 domain changed putative Env contact residues as well as potential N-linked glycosylation sites in many species, with evidence for parallel evolution and trans-specific polymorphism. These data suggest that the primate CD4 receptor is under long-term balancing selection and that this diversification has been the result of a coevolutionary arms race between primate lentiviruses and their hosts., Infection with human and simian immunodeficiency viruses (HIV/SIV) requires binding of the viral envelope glycoprotein (Env) to the host protein CD4 on the surface of immune cells. Although invariant in humans, the Env binding domain of the chimpanzee CD4 is highly polymorphic, with nine coding variants circulating in wild populations. Here, we show that within-species CD4 diversity is not unique to chimpanzees but found in many African primate species. Characterizing the outermost (D1) domain of the CD4 protein in over 500 monkeys and apes, we found polymorphic residues in 24 of 29 primate species, with as many as 11 different coding variants identified within a single species. D1 domain amino acid replacements affected SIV Env-mediated cell entry in a single-round infection assay, restricting infection in a strain- and allele-specific fashion. Several identical CD4 polymorphisms, including the addition of N-linked glycosylation sites, were found in primate species from different genera, providing striking examples of parallel evolution. Moreover, seven different guenons (Cercopithecus spp.) shared multiple distinct D1 domain variants, pointing to long-term trans-specific polymorphism. These data indicate that the HIV/SIV Env binding region of the primate CD4 protein is highly variable, both within and between species, and suggest that this diversity has been maintained by balancing selection for millions of years, at least in part to confer protection against primate lentiviruses. Although long-term SIV-infected species have evolved specific mechanisms to avoid disease progression, primate lentiviruses are intrinsically pathogenic and have left their mark on the host genome.

The sensitivity of tropical forest carbon to climate is a key uncertainty in predicting global climate change. Although short-term drying and warming are known to affect forests, it is unknown if such effects translate into long-term responses. Here, we analyze 590 permanent plots measured across the tropics to derive the equilibrium climate controls on forest carbon. Maximum temperature is the most important predictor of aboveground biomass (-9.1 megagrams of carbon per hectare per degree Celsius), primarily by reducing woody productivity, and has a greater impact per °C in the hottest forests (>32.2°C). Our results nevertheless reveal greater thermal resilience than observations of short-term variation imply. To realize the long-term climate adaptation potential of tropical forests requires both protecting them and stabilizing Earth's climate., Our plot monitoring networks havebeen supported by multiple grants from a large number of funding bodies: European Research Council; Natural Environment Research Council; European Union’s Fifth, Sixth, and Seventh Framework Programme; Royal Society, Leverhulme Trust; Gordon and Betty Moore Foundation; David and Lucile Packard Foundation; State of São Paulo Research Foundation (FAPESP); National Council for Science and Technology Development of Brazil (CNPq); Agence Nationale de la Recherche; Conservation International; Missouri Botanical Garden; Smithsonian Institution; Wildlife Conservation Society; National Geographic Society; Centre for International Forestry; and Gabon’s National Park Agency.

The giant pangolin (Smutsia gigantea) is the largest of all living pangolins, approaching a total length of 180 cm and a weight of more than 30 kg. The species is widely distributed in West and Central Africa in a variety of habitats, including primary and secondary forest, swamp forest and wooded savanna, ranging from Senegal to western Tanzania south to central Democratic Republic of the Congo. Primarily solitary and nocturnal, giant pangolins use their large muscular forelimbs to open subterranean termitaries in contrast to Temminck’s pangolin (S. temminckii) which do little digging. Available data suggest the species is uncommon and declining in many parts of its range, with no recent data for several countries from which it has been previously recorded. Despite their presence in many protected areas and international trade controls, hunting and poaching for bushmeat and international trade remains the primary threat to the species.

Genomic data can be a powerful tool for inferring ecology, behaviour and conservation needs of highly elusive species, particularly when other sources of information are hard to come by. Here we focus on the dryas monkey, an endangered primate endemic to the Congo Basin with cryptic behaviour and possibly less than 250 remaining individuals. Using whole genome data we show that the dryas monkey represents a sister lineage to the vervet monkeys and has diverged from them at least 1 million years ago with additional bi-directional gene flow 590,000 – 360,000 years ago. After bonobo-chimpanzee admixture, this is the second reported case of gene flow that most likely involved crossing the Congo River, a strong dispersal barrier. As the demographic history of bonobos and dryas monkey shows similar patterns of population increase during this time period, we hypothesise that the fluvial topology of the Congo River might have been more dynamic than previously recognised. As a result of dryas monkey - vervet admixture, genes involved in resistance to the simian immunodeficiency virus (SIV) have been exchanged, possibly indicating adaptive introgression. Despite the presence of several homozygous loss-of-function mutations in genes associated with reduced sperm mobility and immunity, we find high genetic diversity and low levels of inbreeding and genetic load in the studied dryas monkey individual. This suggests that the current population carries sufficient genetic variability for the long-term survival of this species. We thus provide an example of how genomic data can directly improve our understanding of elusive species.

The Laverania clade comprises the human malaria parasite Plasmodium falciparum as well as at least seven additional parasite species that infect wild African apes. A recent analysis of Laverania genome sequences (Otto TD, et al. 2018. Genomes of all known members of a Plasmodium subgenus reveal paths to virulent human malaria. Nat Microbiol. 3:687-697) reported three instances of inter-species gene transfer, one of which had previously been described. Generating gene sequences from additional ape parasites and re-examining sequencing reads generated in the Otto et al. study, we identified one of the newly described gene transfers as an assembly artefact of sequences derived from a sample co-infected by two parasite species. The second gene transfer between ancestors of two divergent chimpanzee parasite lineages was confirmed, but involved a much larger number of genes than originally described, many of which encode exported proteins that remodel, or bind to, erythrocytes. Because successful hybridisation between Laverania species is very rare, it will be important to determine to what extent these gene transfers have shaped their host interactions.

Structurally intact tropical forests sequestered about half of the global terrestrial carbon uptake over the 1990s and early 2000s, removing about 15 per cent of anthropogenic carbon dioxide emissions1,2,3. Climate-driven vegetation models typically predict that this tropical forest ‘carbon sink’ will continue for decades4,5. Here we assess trends in the carbon sink using 244 structurally intact African tropical forests spanning 11 countries, compare them with 321 published plots from Amazonia and investigate the underlying drivers of the trends. The carbon sink in live aboveground biomass in intact African tropical forests has been stable for the three decades to 2015, at 0.66 tonnes of carbon per hectare per year (95 per cent confidence interval 0.53–0.79), in contrast to the long-term decline in Amazonian forests6. Therefore the carbon sink responses of Earth’s two largest expanses of tropical forest have diverged. The difference is largely driven by carbon losses from tree mortality, with no detectable multi-decadal trend in Africa and a long-term increase in Amazonia. Both continents show increasing tree growth, consistent with the expected net effect of rising atmospheric carbon dioxide and air temperature7,8,9. Despite the past stability of the African carbon sink, our most intensively monitored plots suggest a post-2010 increase in carbon losses, delayed compared to Amazonia, indicating asynchronous carbon sink saturation on the two continents. A statistical model including carbon dioxide, temperature, drought and forest dynamics accounts for the observed trends and indicates a long-term future decline in the African sink, whereas the Amazonian sink continues to weaken rapidly. Overall, the uptake of carbon into Earth’s intact tropical forests peaked in the 1990s. Given that the global terrestrial carbon sink is increasing in size, independent observations indicating greater recent carbon uptake into the Northern Hemisphere landmass10 reinforce our conclusion that the intact tropical forest carbon sink has already peaked. This saturation and ongoing decline of the tropical forest carbon sink has consequences for policies intended to stabilize Earth’s climate.

OBJECTIVES The guenons (tribe Cercopithecini) are a diverse and primarily arboreal radiation of Old World monkeys from Africa. However, preliminary behavioral observations of the lesula (Cercopithecus lomamiensis), a little-known guenon species described in 2012, report it spending substantial amounts of time on the ground. New specimens allow us to present the first description of lesula postcranial morphology and apply a comparative functional morphology approach to supplement our knowledge of its locomotor behavior. MATERIALS AND METHODS To infer the substrate use preferences of the lesula, 22 postcranial variables correlated with locomotion were assessed in a sample of 151 adult guenon specimens, including two C. lomamiensis. Using multivariate statistical analyses, we predict the amount of time the lesula spends on the ground relative to the comparative sample. RESULTS Results suggest that the lesula spends nearly half its time on the ground, and the two available individuals were classified as semiterrestrial and terrestrial with strong support. Comparisons with two outgroup cercopithecid taxa (Colobus guereza and Macaca mulatta) demonstrate that, as a group, guenons retain signals of a generalized, semiterrestrially adapted postcranium compared to specialized arboreal cercopithecids. DISCUSSION These results corroborate preliminary behavioral observations of the lesula as a semiterrestrial to terrestrial primate and imply multiple evolutionary transitions in substrate use among the guenon radiation. A broader view of cercopithecoid evolution suggests that a semiterrestrial ancestor for extant guenons is more parsimonious than an arboreal one, indicating that the arboreal members of the group are probably recently derived from a more semiterrestrial ancestor.

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

Classical ecology provides principles for construction and function of biological communities, but to what extent these apply to the animal-associated microbiota is just beginning to be assessed. Here, we investigated the influence of several well-known ecological principles on animal-associated microbiota by characterizing gut microbial specimens from bilaterally symmetrical animals (Bilateria) ranging from flies to whales. A rigorously vetted sample set containing 265 specimens from 64 species was assembled. Bacterial lineages were characterized by 16S rRNA gene sequencing. Previously published samples were also compared, allowing analysis of over 1,098 samples in total. A restricted number of bacterial phyla was found to account for the great majority of gut colonists. Gut microbial composition was associated with host phylogeny and diet. We identified numerous gut bacterial 16S rRNA gene sequences that diverged deeply from previously studied taxa, identifying opportunities to discover new bacterial types. The number of bacterial lineages per gut sample was positively associated with animal mass, paralleling known species-area relationships from island biogeography and implicating body size as a determinant of community stability and niche complexity. Samples from larger animals harbored greater numbers of anaerobic communities, specifying a mechanism for generating more-complex microbial environments. Predictions for species/abundance relationships from models of neutral colonization did not match the data set, pointing to alternative mechanisms such as selection of specific colonists by environmental niche. Taken together, the data suggest that niche complexity increases with gut size and that niche selection forces dominate gut community construction., IMPORTANCE The intestinal microbiome of animals is essential for health, contributing to digestion of foods, proper immune development, inhibition of pathogen colonization, and catabolism of xenobiotic compounds. How these communities assemble and persist is just beginning to be investigated. Here we interrogated a set of gut samples from a wide range of animals to investigate the roles of selection and random processes in microbial community construction. We show that the numbers of bacterial species increased with the weight of host organisms, paralleling findings from studies of island biogeography. Communities in larger organisms tended to be more anaerobic, suggesting one mechanism for niche diversification. Nonselective processes enable specific predictions for community structure, but our samples did not match the predictions of the neutral model. Thus, these findings highlight the importance of niche selection in community construction and suggest mechanisms of niche diversification.

Malaria parasites, though widespread among wild chimpanzees and gorillas, have not been detected in bonobos. Here, we show that wild-living bonobos are endemically Plasmodium infected in the eastern-most part of their range. Testing 1556 faecal samples from 11 field sites, we identify high prevalence Laverania infections in the Tshuapa-Lomami-Lualaba (TL2) area, but not at other locations across the Congo. TL2 bonobos harbour P. gaboni, formerly only found in chimpanzees, as well as a potential new species, Plasmodium lomamiensis sp. nov. Rare co-infections with non-Laverania parasites were also observed. Phylogenetic relationships among Laverania species are consistent with co-divergence with their gorilla, chimpanzee and bonobo hosts, suggesting a timescale for their evolution. The absence of Plasmodium from most field sites could not be explained by parasite seasonality, nor by bonobo population structure, diet or gut microbiota. Thus, the geographic restriction of bonobo Plasmodium reflects still unidentified factors that likely influence parasite transmission., Unlike chimpanzees and gorillas, bonobos have not been found infected by malaria parasites in the wild. Here, Liu et al. report more thorough survey and sequencing results showing that bonobos host malaria parasites, including a yet-unknown species, but only in the eastern-most part of their range.

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., 0, SCOPUS: ar.j, info:eu-repo/semantics/published

Global change is impacting forests worldwide, threatening biodiversity and ecosystem services including climate regulation. Understanding how forests respond is critical to forest conservation and climate protection. This review describes an international network of 59 long-term forest dynamics research sites (CTFS-ForestGEO) useful for characterizing forest responses to global change. Within very large plots (median size 25ha), all stems 1cm diameter are identified to species, mapped, and regularly recensused according to standardized protocols. CTFS-ForestGEO spans 25 degrees S-61 degrees N latitude, is generally representative of the range of bioclimatic, edaphic, and topographic conditions experienced by forests worldwide, and is the only forest monitoring network that applies a standardized protocol to each of the world's major forest biomes. Supplementary standardized measurements at subsets of the sites provide additional information on plants, animals, and ecosystem and environmental variables. CTFS-ForestGEO sites are experiencing multifaceted anthropogenic global change pressures including warming (average 0.61 degrees C), changes in precipitation (up to +/- 30% change), atmospheric deposition of nitrogen and sulfur compounds (up to 3.8g Nm(-2)yr(-1) and 3.1g Sm(-2)yr(-1)), and forest fragmentation in the surrounding landscape (up to 88% reduced tree cover within 5km). The broad suite of measurements made at CTFS-ForestGEO sites makes it possible to investigate the complex ways in which global change is impacting forest dynamics. Ongoing research across the CTFS-ForestGEO network is yielding insights into how and why the forests are changing, and continued monitoring will provide vital contributions to understanding worldwide forest diversity and dynamics in an era of global change.

Mbau forest covers much of the Congo, and shifts in its composition could have a large impact on the African tropics. The Ituri forest in east Congo is near a boundary between the monodominant mbau type and non-mbau mixed forest, and two 20-ha censuses of trees ≥ 1 cm diameter were carried out over 12 y to monitor forest change. Based on published diameter allometry, mbau forest had 535 Mg ha−1 biomass above ground and gained 1.1 Mg ha−1 y−1. Mixed forest had 399 Mg ha−1 and gained 3 Mg ha−1 y−1. The mbau tree (Gilbertiodendron dewevrei) increased its share of biomass from 4.1% to 4.4% in mixed forest; other common species also increased. Sapling density declined at both sites, likely because increased biomass meant shadier understorey, but the mbau tree increased in sapling density, suggesting it will become more important in the future. Tree mortality and growth rates were low relative to other tropical forests, especially in the mbau plots. Shifting toward G. dewevrei would represent a large gain in carbon in the mixed forest, but mbau is presently more important as a high-carbon stock: biomass lost during forest harvest could not recuperate for centuries due to slow community dynamics.

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.

Large tropical trees and a few dominant species were recently identified as the main structuring elements of tropical forests. However, such result did not translate yet into quantitative approaches which are essential to understand, predict and monitor forest functions and composition over large, often poorly accessible territories. Here we show that the above-ground biomass (AGB) of the whole forest can be predicted from a few large trees and that the relationship is proved strikingly stable in 175 1-ha plots investigated across 8 sites spanning Central Africa. We designed a generic model predicting AGB with an error of 14% when based on only 5% of the stems, which points to universality in forest structural properties. For the first time in Africa, we identified some dominant species that disproportionally contribute to forest AGB with 1.5% of recorded species accounting for over 50% of the stock of AGB. Consequently, focusing on large trees and dominant species provides precise information on the whole forest stand. This offers new perspectives for understanding the functioning of tropical forests and opens new doors for the development of innovative monitoring strategies., SCOPUS: ar.j, info:eu-repo/semantics/published

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.

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 Ituri Forest, Democratic Republic of Congo (formerly Zaire) is an example of a closed canopy forest showing extreme depletion in (13)C. delta(13)C values for plants from the canopy top, from gaps in the canopy, and from the subcanopy average -29.0+/-1.7 per thousand, -30.4+/-0.9 per thousand, and -34.0+/-1.5 per thousand, respectively. The delta(13)C of forest mammals show these differences, with the subcanopy browsers (okapi, dwarf antelope) having delta(13)C values for tooth enamel much more negative than subcanopy frugivores who derive their food from the canopy top, and from folivores and omnivores living in gap or clearing areas. Nitrogen isotopes in plants from this ecosystem have an average delta(15)N value of 5.4+/-1.8 per thousand and do not show significant differences at the 95% confidence interval between plants from the canopy top, from gaps in the canopy, and from the subcanopy. The delta(18)O(SMOW) values of surface waters in the study area are between -2.0 and -2.7. The delta(18)O(PDB) for tooth enamel ranged from -3 to +7 per thousand.

We report above-ground biomass (AGB), basal area, stemdensity and wood mass density estimates from 260 sample plots (mean size: 1.2 ha) in intact closed-canopy tropical forests across 12 African countries. Mean AGB is 395.7 Mg dry mass ha-1 (95% CI: 14.3), substantially higher than Amazonian values, with the Congo Basin and contiguous forest region attaining AGB values (429 Mg ha-1) similar to those of Bornean forests, and significantly greater than East or West African forests. AGB therefore appears generally higher in palaeo- comparedwithneotropical forests.However, mean stem density is low(426±11 stems ha-1 greater than or equal to 100 mm diameter) compared with both Amazonian and Bornean forests (cf. approx. 600) and is the signature structural feature of African tropical forests. While spatial autocorrelation complicates analyses, AGB shows a positive relationship with rainfall in the driest nine months of the year, and an opposite association with the wettest three months of the year; a negative relationship with temperature; positive relationship with clay-rich soils; and negative relationshipswith C:Nratio (suggesting a positive soil phosphorus- AGB relationship), and soil fertility computed as the sum of base cations. The results indicate that AGB is mediated by both climate and soils, and suggest that the AGB of African closed-canopy tropical forests may be particularly sensitive to future precipitation and temperature changes. © 2013 The Authors., 0, SCOPUS: ar.j, info:eu-repo/semantics/published

The central Ituri Forest of Zaïre is characterized by monodominant stands of Gilbert-iodendron dewevrei (mbau forest) within a matrix of mixed forest where Julbernardia seretii is subdominant. Both species produce supra-annual, synchronous (mast) seed crops. The relationship of mast seed production to monodominance for the two tree species was evaluated by investigating: (1) the relationship between seed density and seed survival, (2) the relative impact of mammal versus insect seed predators in high and low seed density areas, and (3) post-germination survival in the understorey. For the latter, tagged seedlings were monitored over 10 years and both species were censused in different understorey size classes. Percentage seed survival was significantly greater for Julbernardia (but not for Gilbertiodendron) in areas of high conspecific seed density. Generalist fruit- and seed-eating mammals were the principal predators on seeds of both species when removed to areas without seed-fall. In mast areas high seed densities appeared to satiate mammalian seed predators but not the specialized beetle seed predators that were the major cause of seed destruction. Gilbertiodendron seedlings survived longer in the forest understorey than did seedlings of Julbernardia. Furthermore, Gilbertiodendron made up a greater proportion of the larger subcanopy size classes in monodominant forest than did Julbernardia in mixed forests. These results suggest that, irrespective of mortality to seeds, the ability to persist in the understorey may confer on Gilbertiodendron a greater probability of reaching and dominating the canopy.

In June 2007, a previously undescribed monkey known locally as “lesula” was found in the forests of the middle Lomami Basin in central Democratic Republic of Congo (DRC). We describe this new species as Cercopithecus lomamiensis sp. nov., and provide data on its distribution, morphology, genetics, ecology and behavior. C. lomamiensis is restricted to the lowland rain forests of central DRC between the middle Lomami and the upper Tshuapa Rivers. Morphological and molecular data confirm that C. lomamiensis is distinct from its nearest congener, C. hamlyni, from which it is separated geographically by both the Congo (Lualaba) and the Lomami Rivers. C. lomamiensis, like C. hamlyni, is semi-terrestrial with a diet containing terrestrial herbaceous vegetation. The discovery of C. lomamiensis highlights the biogeographic significance and importance for conservation of central Congo’s interfluvial TL2 region, defined from the upper Tshuapa River through the Lomami Basin to the Congo (Lualaba) River. The TL2 region has been found to contain a high diversity of anthropoid primates including three forms, in addition to C. lomamiensis, that are endemic to the area. We recommend the common name, lesula, for this new species, as it is the vernacular name used over most of its known range.

The response of terrestrial vegetation to a globally changing environment is central to predictions of future levels of atmospheric carbon dioxide1, 2. The role of tropical forests is critical because they are carbon-dense and highly productive3, 4. Inventory plots across Amazonia show that old-growth forests have increased in carbon storage over recent decades5, 6, 7, but the response of one-third of the world's tropical forests in Africa8 is largely unknown owing to an absence of spatially extensive observation networks9, 10. Here we report data from a ten-country network of long-term monitoring plots in African tropical forests. We find that across 79 plots (163 ha) above-ground carbon storage in live trees increased by 0.63 Mg C ha-1 yr-1 between 1968 and 2007 (95% confidence interval (CI), 0.22¿0.94; mean interval, 1987¿96). Extrapolation to unmeasured forest components (live roots, small trees, necromass) and scaling to the continent implies a total increase in carbon storage in African tropical forest trees of 0.34 Pg C yr-1 (CI, 0.15¿0.43). These reported changes in carbon storage are similar to those reported for Amazonian forests per unit area6, 7, providing evidence that increasing carbon storage in old-growth forests is a pan-tropical phenomenon. Indeed, combining all standardized inventory data from this study and from tropical America and Asia5, 6, 11 together yields a comparable figure of 0.49 Mg C ha-1 yr-1 (n = 156; 562 ha; CI, 0.29¿0.66; mean interval, 1987¿97). This indicates a carbon sink of 1.3 Pg C yr-1 (CI, 0.8¿1.6) across all tropical forests during recent decades. Taxon-specific analyses of African inventory and other data12 suggest that widespread changes in resource availability, such as increasing atmospheric carbon dioxide concentrations, may be the cause of the increase in carbon stocks13, as some theory14 and models2, 10, 15 predict.

In Amazonian tropical forests, recent studies have reported increases in aboveground biomass and in primary productivity, as well as shifts in plant species composition favouring fast-growing species over slow-growing ones. This pervasive alteration of mature tropical forests was attributed to global environmental change, such as an increase in atmospheric CO2 concentration, nutrient deposition, temperature, drought frequency, and/or irradiance. We used standardized, repeated measurements of over 2 million trees in ten large (16–52 ha each) forest plots on three continents to evaluate the generality of these findings across tropical forests. Aboveground biomass increased at seven of our ten plots, significantly so at four plots, and showed a large decrease at a single plot. Carbon accumulation pooled across sites was significant (+0.24 MgC ha−1 y−1, 95% confidence intervals [0.07, 0.39] MgC ha−1 y−1), but lower than reported previously for Amazonia. At three sites for which we had data for multiple census intervals, we found no concerted increase in biomass gain, in conflict with the increased productivity hypothesis. Over all ten plots, the fastest-growing quartile of species gained biomass (+0.33 [0.09, 0.55] % y−1) compared with the tree community as a whole (+0.15 % y−1); however, this significant trend was due to a single plot. Biomass of slow-growing species increased significantly when calculated over all plots (+0.21 [0.02, 0.37] % y−1), and in half of our plots when calculated individually. Our results do not support the hypothesis that fast-growing species are consistently increasing in dominance in tropical tree communities. Instead, they suggest that our plots may be simultaneously recovering from past disturbances and affected by changes in resource availability. More long-term studies are necessary to clarify the contribution of global change to the functioning of tropical forests., Author Summary Recent studies have reported major changes in mature tropical forests, with increases in both forest biomass and net primary productivity, as well as shifts in plant species composition that favour fast-growing species over slow-growing ones. These pervasive alterations were attributed to global environmental change, and may result in dramatic shifts in the functioning of tropical forest ecosystems. We reassessed these findings using a dataset of large permanent forest plots on three continents. We found that tree biomass increased at seven of our ten plots, and showed a large decrease at a single plot. Overall, this increase was significant, albeit lower than reported previously for Amazonian forests. At three sites for which we had data for multiple census intervals, we found no concerted increase in biomass gain, in conflict with the increased productivity hypothesis. With the exception of one plot, slow-growing species gained more biomass than either fast-growing species or the tree community as a whole. Hence, our results do not support the hypothesis that fast-growing species are consistently increasing in dominance in tropical tree communities. Overall, our results suggest that our plots may be simultaneously recovering from past disturbances and affected by changes in resource availability., An analysis of changes in biomass and primary productivity across ten large tropical forest plots revealed that aboveground biomass increased significantly, fast-growing species increased significantly in dominance at a single site, and slow-growing species increased significantly at half of our sites.

Old-growth rain forests that are dominated by a single canopy species occur throughout the tropics, though they account for a limited proportion of the total rain forest area. These forests have been considered anomalies in which development of a more diverse community is deflected by harsh conditions. Very poor soils or an otherwise extreme environment may promote monodominance by excluding potentially competing species, but it is now apparent that monodominant tropical forests also develop under more benign conditions. Field studies have shown that a single species may dominate on undisturbed sites where the soils are similar to those of adjacent old-growth, mixed forests. In these situations the dominant is a superior competitor and/or is particularly tolerant to stresses such as shade. Assertion of dominance by a single species in an old-growth forest appears most likely in areas where the species pool contains few late-succession species with similar life history traits.

Most ecological hypotheses about species coexistence hinge on species differences, but quantifying trait differences across species in diverse communities is often unfeasible. We examined the variation of demographic traits using a global tropical forest data set covering 4500 species in 10 large-scale tree inventories. With a hierarchical Bayesian approach, we quantified the distribution of mortality and growth rates of all tree species at each site. This allowed us to test the prediction that demographic differences facilitate species richness, as suggested by the theory that a tradeoff between high growth and high survival allows species to coexist. Contrary to the prediction, the most diverse forests had the least demographic variation. Although demographic differences may foster coexistence, they do not explain any of the 16-fold variation in tree species richness observed across the tropics.

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

Bonobos (Pan paniscus) inhabit regions south of the Congo River including all areas between its southerly tributaries. To investigate the genetic diversity and evolutionary relationship among bonobo populations, we sequenced mitochondrial DNA from 376 fecal samples collected in seven study populations located within the eastern and western limits of the species' range. In 136 effective samples from different individuals (range: 7-37 per population), we distinguished 54 haplotypes in six clades (A1, A2, B1, B2, C, D), which included a newly identified clade (D). MtDNA haplotypes were regionally clustered; 83 percent of haplotypes were locality-specific. The distribution of haplotypes across populations and the genetic diversity within populations thus showed highly geographical patterns. Using population distance measures, seven populations were categorized in three clusters: the east, central, and west cohorts. Although further elucidation of historical changes in the geological setting is required, the geographical patterns of genetic diversity seem to be shaped by paleoenvironmental changes during the Pleistocene. The present day riverine barriers appeared to have a weak effect on gene flow among populations, except for the Lomami River, which separates the TL2 population from the others. The central cohort preserves a high genetic diversity, and two unique clades of haplotypes were found in the Wamba/Iyondji populations in the central cohort and in the TL2 population in the eastern cohort respectively. This knowledge may contribute to the planning of bonobo conservation.

Gretchen Vogel (“Conflict in Congo threatens bonobos and rare gorillas,” News of the Week, 31 Mar., p. [2386][1]) describes the current plight of apes in the Democratic Republic of Congo (DRC) whose habitat is on the front lines of a civil war that began about 18 months ago. The apes, including bonobos and the eastern lowland gorilla, are being poached for food by troops and refugees. The protection from poaching that national parks would normally provide for wildlife has been thwarted by the disarming of park guards by military leaders. The very presence of researchers and local dedicated conservationists, Vogel notes, has helped to alleviate the situation; however, the presence of dedicated Congolese researchers with outside support can be much more effective than efforts of foreign researchers. The Wildlife Conservation Society (WCS) has been operating throughout the current civil war in collaboration with the Congolese national parks in the Okapi Wildlife Reserve. In Okapi, it has been elephants and ungulates that have been heavily hunted to feed armed forces, which have passed through from Bunia to Kisangani. WCS has been able to work with Gilman International Conservation to support the rehabilitation of infrastructure as well as the salaries of Congolese conservationists. The presence of committed Congolese conservationists has meant that each successive incoming administration or army command has been approached and educated about the importance of the reserve for conservation. This has not stopped all poaching, but it has led to the rearming of the reserve guards and has led to reduced hunting by armed forces. The effectiveness of committed Congolese, in Okapi and elsewhere, has been recognized. UNESCO, with support from the United Nations Foundation (supported through a $1 billion gift from media magnate Ted Turner), has promised nearly $3 million over 4 years to five World Heritage Sites in DRC (Okapi and Garamba, Virunga, Kahuzi-Biega, and Salonga national parks). This support will be used for the salaries of park staff, the only presence on the grounds in some parks, as well as to provide equipment and monitoring. This initiative, which came from several conservation organizations with UNESCO, might become a model for United Nations support to conservation in areas of conflict. [1]: /lookup/doi/10.1126/science.287.5462.2386

A study of the structure and floristics at a transition zone from a monodominant to a more diverse forest in the African humid tropics was conducted to elucidate the mechanisms maintaining floristic diversity and the discontinuity between mixed forests and forests dominated by a single tree species (monodominant). The mixed forest's greater diversity could not be explained by substrate differences, greater maturity, or greater predation on seeds or juveniles. The dominant species of the monodominant forest was shade-tolerant and had poorly dispersed seeds. Tree species associated with the dominant were also found in the mixed forest. Monodominant and mixed forests occur side by side in the Asian and American tropics as well. As in the African example, many of these monodominant forests share most species with neighboring mixed forests. Characteristically, the dominant species have large seeds and shade-tolerant seedlings. Monodominant tropical forests are widespread and may indicate areas that have not ex...

While it is well established that the Mbuti pygmies, hunter-gatherers of the Ituri Forest of Zaire, trade forest products and labor for agricultural foods, it has been assumed that this is a recent development, following contact with shifting cultivators. From field investigations over a five-year period, Hart and Hart found that, while the Mbuti can identify a large number of edible plant species, they actually gather very few of them. Instead, they trade meat, which is seasonally abundant, as well as their own labor for agricultural products rich in starch and carbohydrates. While game is available throughout the year, they cannot profitably utilize protein as a main caloric intake; lean meat requires more calories to digest than it supplies, a point illustrated by what the explorer Henry Stanley called in 1890 the “starvation camps” in the middle of the Ituri: People were dying while surrounded by game. Hart and Hart found that almost all of the oil-rich plant foods collected came from secondary forests rather than from that part of the Ituri hitherto unpenetrated by cultivators. Using historical data from early travellers, archaeological evidence, and linguistic connections linking various Mbuti groups to different cultivators, Hart and Hart make the case that the great evergreen primary forests of the Ituri were probably not capable of sustaining hunter-gatherers in significant numbers prior to the coming of cultivators, about 4000 years ago. This a good illustration of complex and interdependent ties that link what might appear to be distinct adaptations. Here agriculture has made the forest a more hospitable environment for specialized hunter-gatherers by providing a reliable source of energy-rich foods as well as by producing patches of secondary forest in which to forage.

International audience; Plasmodium vivax is the leading cause of human malaria in Asia and Latin America but is absent from most of central Africa due to the near fixation of a mutation that inhibits the expression of its receptor, the Duffy antigen, on human erythrocytes. The emergence of this protective allele is not understood because P. vivax is believed to have originated in Asia. Here we show, using a noninvasive approach, that wild chimpanzees and gorillas throughout central Africa are endemically infected with parasites that are closely related to human P. vivax. Sequence analyses reveal that ape parasites lack host specificity and are much more diverse than human parasites, which form a monophyletic lineage within the ape parasite radiation. These findings indicate that human P. vivax is of African origin and likely selected for the Duffy-negative mutation. All extant human P. vivax parasites are derived from a single ancestor that escaped out of Africa.

Tropical forests store 40–50 per cent of terrestrial vegetation carbon 1. However, spatial variations in aboveground live tree biomass carbon (AGC) stocks remain poorly understood, in particular in tropical montane forests 2. Owing to climatic and soil changes with increasing elevation 3, AGC stocks are lower in tropical montane forests compared with lowland forests 2. Here we assemble and analyse a dataset of structurally intact old-growth forests (AfriMont) spanning 44 montane sites in 12 African countries. We find that montane sites in the AfriMont plot network have a mean AGC stock of 149.4 megagrams of carbon per hectare (95% confidence interval 137.1–164.2), which is comparable to lowland forests in the African Tropical Rainforest Observation Network 4 and about 70 per cent and 32 per cent higher than averages from plot networks in montane 2,5,6 and lowland 7 forests in the Neotropics, respectively. Notably, our results are two-thirds higher than the Intergovernmental Panel on Climate Change default values for these forests in Africa 8. We find that the low stem density and high abundance of large trees of African lowland forests 4 is mirrored in the montane forests sampled. This carbon store is endangered: we estimate that 0.8 million hectares of old-growth African montane forest have been lost since 2000. We provide country-specific montane forest AGC stock estimates modelled from our plot network to help to guide forest conservation and reforestation interventions. Our findings highlight the need for conserving these biodiverse 9,10 and carbon-rich ecosystems.