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.

Similar factors underlie tree abundance in forests in native and alien ranges.

AIM: Alien plant species can cause severe ecological and economic problems, and therefore attract a lot of research interest in biogeography and related fields. To identify potential future invasive species, we need to better understand the mechanisms underlying the abundances of invasive tree species in their new ranges, and whether these mechanisms differ between their native and alien ranges. Here, we test two hypotheses: that greater relative abundance is promoted by (a) functional difference from locally co-occurring trees, and (b) higher values than locally co-occurring trees for traits linked to competitive ability. LOCATION: Global. TIME PERIOD: Recent. MAJOR TAXA STUDIED: Trees. METHODS: We combined three global plant databases: sPlot vegetation-plot database, TRY plant trait database and Global Naturalized Alien Flora (GloNAF) database. We used a hierarchical Bayesian linear regression model to assess the factors associated with variation in local abundance, and how these relationships vary between native and alien ranges and depend on species' traits. RESULTS: In both ranges, species reach highest abundance if they are functionally similar to co-occurring species, yet are taller and have higher seed mass and wood density than co-occurring species. MAIN CONCLUSIONS: Our results suggest that light limitation leads to strong environmental and biotic filtering, and that it is advantageous to be taller and have denser wood. The striking similarities in abundance between native and alien ranges imply that information from tree species' native ranges can be used to predict in which habitats introduced species may become dominant.

An integrated pan-tropical biomass map using multiple reference datasets.

We combined two existing datasets of vegetation aboveground biomass (AGB) (Proceedings of the National Academy of Sciences of the United States of America, 108, 2011, 9899; Nature Climate Change, 2, 2012, 182) into a pan-tropical AGB map at 1-km resolution using an independent reference dataset of field observations and locally calibrated high-resolution biomass maps, harmonized and upscaled to 14 477 1-km AGB estimates. Our data fusion approach uses bias removal and weighted linear averaging that incorporates and spatializes the biomass patterns indicated by the reference data. The method was applied independently in areas (strata) with homogeneous error patterns of the input (Saatchi and Baccini) maps, which were estimated from the reference data and additional covariates. Based on the fused map, we estimated AGB stock for the tropics (23.4 N-23.4 S) of 375 Pg dry mass, 9-18% lower than the Saatchi and Baccini estimates. The fused map also showed differing spatial patterns of AGB over large areas, with higher AGB density in the dense forest areas in the Congo basin, Eastern Amazon and South-East Asia, and lower values in Central America and in most dry vegetation areas of Africa than either of the input maps. The validation exercise, based on 2118 estimates from the reference dataset not used in the fusion process, showed that the fused map had a RMSE 15-21% lower than that of the input maps and, most importantly, nearly unbiased estimates (mean bias 5 Mg dry mass ha(-1) vs. 21 and 28 Mg ha(-1) for the input maps). The fusion method can be applied at any scale including the policy-relevant national level, where it can provide improved biomass estimates by integrating existing regional biomass maps as input maps and additional, country-specific reference datasets.

Growth periodicity in semi-deciduous tropical tree species from the Congo Basin.

In the tropics, more precisely in equatorial dense rainforest, xylogenesis is driven by a little distinct climatological seasonality, and many tropical trees do not show clear growth rings. This makes retrospective analyses and modeling of future tree performance difficult. This research investigates the presence, the distinctness, and the periodicity of growth ring for dominant tree species in two semi-deciduous rainforests, which contrast in terms of precipitation dynamics. Eighteen tree species common to both forests were investigated. We used the cambial marking technique and then verified the presence and periodicity of growth-ring boundaries in the wood produced between pinning and collection by microscopic and macroscopic observation. The study showed that all eighteen species can form visible growth rings in both sites. However, the periodicity of ring formation varied significantly within and between species, and within sites. Trees from the site with clearly defined dry season had a higher likelihood to form periodical growth rings compared to those from the site where rainfall seasonality is less pronounced. The distinctness of the formed rings however did not show a site dependency. Periodical growth-ring formation was more likely in fast-growing trees. Furthermore, improvements can be made by a detailed study of the cambial activity through microcores taken at high temporal resolution, to get insight on the phenology of the lateral meristem.

Historical tree phenology data reveal the seasonal rhythms of the Congo Basin rainforest.

Tropical forest phenology directly affects regional carbon cycles, but the relation between species-specific and whole-canopy phenology remains largely uncharacterized. We present a unique analysis of historical tropical tree phenology collected in the central Congo Basin, before large-scale impacts of human-induced climate change. Ground-based long-term (1937-1956) phenological observations of 140 tropical tree species are recovered, species-specific phenological patterns analyzed and related to historical meteorological records, and scaled to characterize stand-level canopy dynamics. High phenological variability within and across species and in climate-phenology relationships is observed. The onset of leaf phenophases in deciduous species was triggered by drought and light availability for a subset of species and showed a species-specific decoupling in time along a bi-modal seasonality. The majority of the species remain evergreen, although central African forests experience relatively low rainfall. Annually a maximum of 1.5% of the canopy is in leaf senescence or leaf turnover, with overall phenological variability dominated by a few deciduous species, while substantial variability is attributed to asynchronous events of large and/or abundant trees. Our results underscore the importance of accounting for constituent signals in canopy-wide scaling and the interpretation of remotely sensed phenology signals.

Geographical Distribution and Genetic Diversity of the Banana Fusarium Wilt Fungus in Laos and Vietnam.

Fusarium wilt, caused by the fungus Fusarium oxysporum f. sp. cubense (Foc), poses a major threat to global banana production. The tropical race 4 (TR4) variant of Foc is a highly virulent form with a large host range, and severely affects Cavendish bananas. Foc TR4 was recently observed within the Greater Mekong Subregion, after Chinese private companies expanded Cavendish production to the region. In this study, extensive surveys conducted across Laos and Vietnam show that Foc TR4 is still mainly constricted to the northern regions of these countries and is limited to Cavendish cultivation settings. In Laos, Foc TR4 is associated with large-scale Cavendish plantations owned by or involved with Chinese companies through which infected planting material could have been imported. In Vietnam, mostly small-holder Cavendish farmers and backyard gardens were affected by Foc TR4. In Vietnam, no direct link is found with Chinese growers, and it is expected the pathogen mainly spreads through local and regional movement of infected planting materials. Foc TR4 was not recorded on banana cultivars other than Cavendish. The extensively cultivated 'Pisang Awak' cultivar was solely infected by VCGs belonging to Foc race 1 and 2, with a high occurrence of VCG 0123 across Laos, and of VCG 0124/5 in Vietnam. Substantial diversity of Foc VCGs was recorded (VCGs 0123, 0124/5, 01218 and 01221) from northern to southern regions in both countries, suggesting that Fusarium wilt is well established in the region. Interviews with farmers indicated that the local knowledge of Fusarium wilt epidemiology and options for disease management was limited. Clear communication efforts on disease epidemiology and management with emphasis on biosecurity practices need to be improved in order to prevent further spread of Foc TR4 to mixed variety smallholder settings.

Large-sized rare tree species contribute disproportionately to functional diversity in resource acquisition in African tropical forest.

Increasing evidence is available for a positive effect of biodiversity on ecosystem productivity and standing biomass, also in highly diverse systems as tropical forests. Biodiversity conservation could therefore be a critical aspect of climate mitigation policies. There is, however, limited understanding of the role of individual species for this relationship, which could aid in focusing conservation efforts and forest management planning. This study characterizes the functional specialization and redundancy for 95% of all tree species (basal area weighted percentage) in a diverse tropical forest in the central Congo Basin and relates this to species' abundance, contribution to aboveground carbon, and maximum size. Functional characterization is based on a set of traits related to resource acquisition (wood density, specific leaf area, leaf carbon, nitrogen and phosphorus content, and leaf stable carbon isotope composition). We show that within both mixed and monodominant tropical forest ecosystems, the highest functional specialization and lowest functional redundancy are solely found in rare tree species and significantly more in rare species holding large-sized individuals. Rare species cover the entire range of low and high functional redundancy, contributing both unique and redundant functions. Loss of species supporting functional redundancy could be buffered by other species in the community, including more abundant species. This is not the case for species supporting high functional specialization and low functional redundancy, which would need specific conservation attention. In terms of tropical forest management planning, we argue that specific conservation of large-sized trees is imperative for long-term maintenance of ecosystem functioning.

Disentangling how management affects biomass stock and productivity of tropical secondary forests fallows.

A better understanding of biomass production in secondary forests after cultivation is essential for assessing the resilience of slash and burn systems and their capacity to deliver ecosystem services. Biomass production is influenced by management legacies, landscape configuration and soil, but these drivers are rarely studied simultaneously, nor is the role of changes in vegetation properties in linking them to biomass production. We assessed how management legacies affect biomass in secondary forests created by slash and burn in the central Congo Basin, and tested whether changes in productivity could be attributed to changes in stem density, functional diversity, functional identity or soil. Using data from 6452 trees in 96 fallow plots nested in 3 study sites, we looked for the main determinants of aboveground biomass (AGB) of woody vegetation in fallow systems. Next, using a subset of 58 plots in fallow fields aged 5 to 10 years, we used confirmatory path analysis to explore the relations between management history, soil, vegetation properties and biomass productivity. The sampled fallow fields had, on average, 58.4 (±46.2) Mg ha-1 AGB. AGB was positively related to both fallow age and to the proportion of remnant trees in AGB and negatively related to the number of previous cultivation cycles. Biomass productivity varied with the number of previous slash-and-burn cycles, with notable declines in the fourth cycle. The effect of management history was mainly through a reduction in the dominance of fast growing tree species and in the number of regenerating stems, which were also indirectly affected by an increase in C. odorata cover. Soil fertility status and the biomass of remnant trees also modified biomass productivity. Our findings suggest that under the current management intensity the capacity of the slash and burn system to provide important ecosystem functions, such as carbon sequestration, is declining.

Reconciling biodiversity and carbon stock conservation in an Afrotropical forest landscape.

Protecting aboveground carbon stocks in tropical forests is essential for mitigating global climate change and is assumed to simultaneously conserve biodiversity. Although the relationship between tree diversity and carbon stocks is generally positive, the relationship remains unclear for consumers or decomposers. We assessed this relationship for multiple trophic levels across the tree of life (10 organismal groups, 3 kingdoms) in lowland rainforests of the Congo Basin. Comparisons across regrowth and old-growth forests evinced the expected positive relationship for trees, but not for other organismal groups. Moreover, differences in species composition between forests increased with difference in carbon stock. These variable associations across the tree of life contradict the implicit assumption that maximum co-benefits to biodiversity are associated with conservation of forests with the highest carbon storage. Initiatives targeting climate change mitigation and biodiversity conservation should include both old-growth and regenerating forests to optimally benefit biodiversity and carbon storage.

Isotopic evidence for oligotrophication of terrestrial ecosystems.

Human societies depend on an Earth system that operates within a constrained range of nutrient availability, yet the recent trajectory of terrestrial nitrogen (N) availability is uncertain. Examining patterns of foliar N concentrations and isotope ratios (δ15N) from more than 43,000 samples acquired over 37 years, here we show that foliar N concentration declined by 9% and foliar δ15N declined by 0.6-1.6‰. Examining patterns across different climate spaces, foliar δ15N declined across the entire range of mean annual temperature and mean annual precipitation tested. These results suggest declines in N supply relative to plant demand at the global scale. In all, there are now multiple lines of evidence of declining N availability in many unfertilized terrestrial ecosystems, including declines in δ15N of tree rings and leaves from herbarium samples over the past 75-150 years. These patterns are consistent with the proposed consequences of elevated atmospheric carbon dioxide and longer growing seasons. These declines will limit future terrestrial carbon uptake and increase nutritional stress for herbivores.

Functional community structure of African monodominant Gilbertiodendron dewevrei forest influenced by local environmental filtering.

Monodominant patches of forest dominated by Gilbertiodendron dewevrei are commonly found in central African tropical forests, alongside forests with high species diversity. Although these forests are generally found sparsely distributed along rivers, their occurrence is not thought to be (clearly) driven by edaphic conditions but rather by trait combinations of G. dewevrei that aid in achieving monodominance. Functional community structure between these monodominant and mixed forests has, however, not yet been compared. Additionally, little is known about nondominant species in the monodominant forest community. These two topics are addressed in this study. We investigate the functional community structure of 10 one-hectare plots of monodominant and mixed forests in a central region of the Congo basin, in DR Congo. Thirteen leaf and wood traits are measured, covering 95% (basal area weighted) of all species present in the plots, including leaf nutrient contents, leaf isotopic compositions, specific leaf area, wood density, and vessel anatomy. The trait-based assessment of G. dewevrei shows an ensemble of traits related to water use and transport that could be favorable for its location near forest rivers. Moreover, indications have been found for N and P limitations in the monodominant forest, possibly related to ectomycorrhizal associations formed with G. dewevrei. Reduced leaf N and P contents are found at the community level for the monodominant forest and for different nondominant groups, as compared to those in the mixed forest. In summary, this work shows that environmental filtering does prevail in the monodominant G. dewevrei forest, leading to lower functional diversity in this forest type, with the dominant species showing beneficial traits related to its common riverine locations and with reduced soil N and P availability found in this environment, both coregulating the tree community assembly.

Correction: Aboveground vs. Belowground Carbon Stocks in African Tropical Lowland Rainforest: Drivers and Implications.

[This corrects the article DOI: 10.1371/journal.pone.0143209.].

Aboveground vs. Belowground Carbon Stocks in African Tropical Lowland Rainforest: Drivers and Implications.

BACKGROUND: African tropical rainforests are one of the most important hotspots to look for changes in the upcoming decades when it comes to C storage and release. The focus of studying C dynamics in these systems lies traditionally on living aboveground biomass. Belowground soil organic carbon stocks have received little attention and estimates of the size, controls and distribution of soil organic carbon stocks are highly uncertain. In our study on lowland rainforest in the central Congo basin, we combine both an assessment of the aboveground C stock with an assessment of the belowground C stock and analyze the latter in terms of functional pools and controlling factors. PRINCIPAL FINDINGS: Our study shows that despite similar vegetation, soil and climatic conditions, soil organic carbon stocks in an area with greater tree height (= larger aboveground carbon stock) were only half compared to an area with lower tree height (= smaller aboveground carbon stock). This suggests that substantial variability in the aboveground vs. belowground C allocation strategy and/or C turnover in two similar tropical forest systems can lead to significant differences in total soil organic C content and C fractions with important consequences for the assessment of the total C stock of the system. CONCLUSIONS/SIGNIFICANCE: We suggest nutrient limitation, especially potassium, as the driver for aboveground versus belowground C allocation. However, other drivers such as C turnover, tree functional traits or demographic considerations cannot be excluded. We argue that large and unaccounted variability in C stocks is to be expected in African tropical rain-forests. Currently, these differences in aboveground and belowground C stocks are not adequately verified and implemented mechanistically into Earth System Models. This will, hence, introduce additional uncertainty to models and predictions of the response of C storage of the Congo basin forest to climate change and its contribution to the terrestrial C budget.

Conventional tree height-diameter relationships significantly overestimate aboveground carbon stocks in the Central Congo Basin.

Policies to reduce emissions from deforestation and forest degradation largely depend on accurate estimates of tropical forest carbon stocks. Here we present the first field-based carbon stock data for the Central Congo Basin in Yangambi, Democratic Republic of Congo. We find an average aboveground carbon stock of 162 ± 20 Mg C ha(-1) for intact old-growth forest, which is significantly lower than stocks recorded in the outer regions of the Congo Basin. The best available tree height-diameter relationships derived for Central Africa do not render accurate canopy height estimates for our study area. Aboveground carbon stocks would be overestimated by 24% if these inaccurate relationships were used. The studied forests have a lower stature compared with forests in the outer regions of the basin, which confirms remotely sensed patterns. Additionally, we find an average soil carbon stock of 111 ± 24 Mg C ha(-1), slightly influenced by the current land-use change.

Above-ground biomass and structure of 260 African tropical forests.

We report above-ground biomass (AGB), basal area, stem density 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- compared with neotropical 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 relationships with C : N ratio (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.