Maxime Réjou-Méchain

Improved allometric models to estimate the aboveground biomass of tropical trees

Terrestrial carbon stock mapping is important for the successful implementation of climate change mitigation policies. Its accuracy depends on the availability of reliable allometric models to infer oven-dry aboveground biomass of trees from census data. The degree of uncertainty associated with previously published pantropical aboveground biomass allometries is large. We analyzed a global database of directly harvested trees at 58 sites, spanning a wide range of climatic conditions and vegetation types (4004 trees ≥ 5 cm trunk diameter). When trunk diameter, total tree height, and wood specific gravity were included in the aboveground biomass model as covariates, a single model was found to hold across tropical vegetation types, with no detectable effect of region or environmental factors. The mean percent bias and variance of this model was only slightly higher than that of locally fitted models. Wood specific gravity was an important predictor of aboveground biomass, especially when including a much broader range of vegetation types than previous studies. The generic tree diameter-height relationship depended linearly on a bioclimatic stress variable E, which compounds indices of temperature variability, precipitation variability, and drought intensity. For cases in which total tree height is unavailable for aboveground biomass estimation, a pantropical model incorporating wood density, trunk diameter, and the variable E outperformed previously published models without height. However, to minimize bias, the development of locally derived diameter-height relationships is advised whenever possible. Both new allometric models should contribute to improve the accuracy of biomass assessment protocols in tropical vegetation types, and to advancing our understanding of architectural and evolutionary constraints on woody plant development.

Seeing Central African forests through their largest trees

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.

Amazon forest response to repeated droughts

The Amazon Basin has experienced more variable climate over the last decade, with a severe and widespread drought in 2005 causing large basin-wide losses of biomass. A drought of similar climatological magnitude occurred again in 2010; however, there has been no basin-wide ground-based evaluation of effects on vegetation. We examine to what extent the 2010 drought affected forest dynamics using ground-based observations of mortality and growth from an extensive forest plot network. We find that during the 2010 drought interval, forests did not gain biomass (net change: −0.43 Mg ha−1, confidence interval (CI): −1.11, 0.19, n = 97), regardless of whether forests experienced precipitation deficit anomalies. This contrasted with a long-term biomass sink during the baseline pre-2010 drought period (1998 to pre-2010) of 1.33 Mg ha−1 yr−1 (CI: 0.90, 1.74, p < 0.01). The resulting net impact of the 2010 drought (i.e., reversal of the baseline net sink) was −1.95 Mg ha−1 yr−1 (CI:−2.77, −1.18; p < 0.001). This net biomass impact was driven by an increase in biomass mortality (1.45 Mg ha−1 yr−1 CI: 0.66, 2.25, p < 0.001) and a decline in biomass productivity (−0.50 Mg ha−1 yr−1, CI:−0.78, −0.31; p < 0.001). Surprisingly, the magnitude of the losses through tree mortality was unrelated to estimated local precipitation anomalies and was independent of estimated local pre-2010 drought history. Thus, there was no evidence that pre-2010 droughts compounded the effects of the 2010 drought. We detected a systematic basin-wide impact of the 2010 drought on tree growth rates across Amazonia, which was related to the strength of the moisture deficit. This impact differed from the drought event in 2005 which did not affect productivity. Based on these ground data, live biomass in trees and corresponding estimates of live biomass in lianas and roots, we estimate that intact forests in Amazonia were carbon neutral in 2010 (−0.07 Pg C yr−1 CI:−0.42, 0.23), consistent with results from an independent analysis of airborne estimates of land-atmospheric fluxes during 2010. Relative to the long-term mean, the 2010 drought resulted in a reduction in biomass carbon uptake of 1.1 Pg C, compared to 1.6 Pg C for the 2005 event.

Geological substrates shape tree species and trait distributions in African moist forests.

Background Understanding the factors that shape the distribution of tropical tree species at large scales is a central issue in ecology, conservation and forest management. The aims of this study were to (i) assess the importance of environmental factors relative to historical factors for tree species distributions in the semi-evergreen forests of the northern Congo basin; and to (ii) identify potential mechanisms explaining distribution patterns through a trait-based approach. Methodology/Principal Findings We analyzed the distribution patterns of 31 common tree species in an area of more than 700,000 km2 spanning the borders of Cameroon, the Central African Republic, and the Republic of Congo using forest inventory data from 56,445 0.5-ha plots. Spatial variation of environmental (climate, topography and geology) and historical factors (human disturbance) were quantified from maps and satellite records. Four key functional traits (leaf phenology, shade tolerance, wood density, and maximum growth rate) were extracted from the literature. The geological substrate was of major importance for the distribution of the focal species, while climate and past human disturbances had a significant but lesser impact. Species distribution patterns were significantly related to functional traits. Species associated with sandy soils typical of sandstone and alluvium were characterized by slow growth rates, shade tolerance, evergreen leaves, and high wood density, traits allowing persistence on resource-poor soils. In contrast, fast-growing pioneer species rarely occurred on sandy soils, except for Lophira alata. Conclusions/Significance The results indicate strong environmental filtering due to differential soil resource availability across geological substrates. Additionally, long-term human disturbances in resource-rich areas may have accentuated the observed patterns of species and trait distributions. Trait differences across geological substrates imply pronounced differences in population and ecosystem processes, and call for different conservation and management strategies.

Regional variation in tropical forest tree species composition in the Central African Republic: an assessment based on inventories by forest companies

Understanding how species assemblages are structured in relation to environmental variation is a central issue in community ecology. However, factors that create regional variation in relative species abundances have been little studied due to the rarity of large-scale datasets. Here, we investigated a large dataset (30 180 0.5-ha plots spread over 1 600 000 ha) gathered from forest planning inventories in the semi-deciduous forest of the south western Central African Republic. We used Correspondence Analysis and Non-Symmetric Correspondence Analysis on Instrumental Variables to analyse variation in the abundance of 73 common tree species in relation to soil type, rainfall and proximity to villages. Together, environmental variables explained 10.3% of multi-species floristic variation among plots, and the regional spatial structure almost disappeared when the effects of these variables were removed. A Trend Surface Analysis using a third order polynomial function of the geographical coordinates of the plots explained 14.5% of the floristic variation and more than 75% of this variation was explained by environmental variables. Sandy soil was the most influential factor affecting floristic composition. Residual spatial variation not explained by the environmental variables probably reflects the natural and anthropogenic history of the vegetation.

Prevalence of phylogenetic clustering at multiple scales in an African rain forest tree community

Summary 1. In highly diverse ecosystems, such as tropical forests, the relative importance of mechanisms underlying species coexistence (e.g. habitat filtering, competitive exclusion, neutral dynamics) is still poorly known and probably varies depending on spatial and phylogenetic scales. 2. Here, we develop new approaches for dissecting simultaneously the phylogenetic structure of communities at different phylogenetic depths and spatial scales. We tested with simulations that our method is able to disentangle overdispersion and clustering effects occurring at contrasted phylogenetic depths. 3. We applied our approaches to a 50 ha Forest Dynamic Plot located in Korup National Park (Cameroon) where 329,000 tree stems ≥ 1 cm in diameter were identified and mapped, and using a newly generated dated molecular phylogenetic tree based on 2 plastid loci (rbcL and matK), including 272 species from Korup (97% of the individuals). 4. Significant patterns of phylogenetic turnover were detected across 20 9 20 m 2 quadrats at most spatial scales, with higher turnover between topographic habitats than within habitats, indicating the prevalence of habitat filtering processes. Spatial phylogenetic clustering was detected over the entire range of phylogenetic depths indicating that competitive exclusion does not generate a pattern of phylogenetic overdispersion at this scale, even at a shallow phylogenetic depth. 5. Using an individual-based approach, we also show that closely related species tended to aggregate spatially until a scale of 1 m. However, the signal vanishes at smaller distance, suggesting that competitive exclusion can balance the impact of environmental filtering at a very fine spatial scale. 6. Synthesis. Using new methods to characterize the structure of communities across spatial and phylogenetic scales, we inferred the relative importance of the mechanisms underlying species coexistence in tropical forests. Our analysis confirms that environmental filtering processes are key in the structuring of natural communities at most spatial scales. Although negative-density tends to limit coexistence of closely related species at very short distance (<1 m), its influence is largely veiled by environmental filtering at larger distances.

Evidence for arrested succession in a liana-infested Amazonian forest

1. Empirical evidence and modelling both suggest that global changes may lead to an increased dominance of lianas and thus to an increased prevalence of liana-infested forest formations in tropical forests. The implications for tropical forest structure and the carbon cycle remain poorly understood. 2. We studied the ecological processes underpinning the structure and dynamics of a liana-infested forest in French Guiana, using a combination of long-term surveys (tree, liana, seedling and litterfall), soil chemical analyses and remote-sensing approaches (LiDAR and Landsat). 3. At stand scale and for adult trees, the liana-infested forest had higher growth, recruitment and mortality rates than the neighbouring high-canopy forest. Both total seedling density and tree seedling recruitment were lower in the liana-infested forest. Stand scale above-ground biomass of the liana-infested forest was 58% lower than in the high-canopy forest. 4. Above-ground net primary productivity (ANPP) was comparable in the liana-infested and highcanopy forests. However, due to more abundant leaf production, the relative contribution of fast turnover carbon pools to ANPP was larger in the liana-infested forest and the carbon residence time was half that of the high-canopy forest. 5. Although soils of the liana-infested forest were richer in nutrients, soil elemental ratios suggest that liana-infested forest and high-canopy forest soils both derive from the same geological substrate. The higher nutrient concentration in the liana-infested forest may therefore be the result of a release of nutrients from vegetation after a forest blowdown. 6. Using small-footprint LiDAR campaigns, we show that the overall extent of the liana-infested forest has remained stable from 2007 to 2012 but about 10% of the forest area changed in forest cover type. Landsat optical imagery confirms the liana-infested forest presence in the landscape for at least 25 years. 7. Synthesis. Because persistently high rates of liana infestation are maintained by the fast dynamics of the liana-infested forest, liana-infested forests here appear to be the result of an arrested tropical forest succession. If the prevalence of such arrested succession forests were to increase in the future, this would have important implications for the carbon sink potential of Amazonian forests.

Guidelines for documenting and reporting tree allometric equations

1 IntroductionGiven the pressing need to quantify carbon fluxes associatedwith terrestrial vegetation dynamics, an increasing number ofresearchers have sought to improve estimates of tree volume,biomass, and carbon stocks. Tree allometric equations arecritical tools for such purpose and have the potential toimprove our understanding about carbon sequestration inwoody vegetation, to support the implementation of policiesand mechanisms designed to mitigate climate change (e.g.CDM and REDD+; Agrawal et al. 2011), to calculate costsand benefits associated with forest carbon projects, and toimprove bioenergy systems and sustainable forest manage-ment (Henry et al. 2013).

Properties of Similarity Indices under Niche-Based and Dispersal-Based Processes in Communities

Little is known about the combined impact of habitat filtering and dispersal limitation on species turnover patterns. To gain new insights, we constructed a spatially explicit community model wherein we controlled dispersal distances, the strength of habitat filtering, and the grain of habitat heterogeneity to study the distance decay of several (dis)similarity indices. The impact of habitat filtering is dependent on the ratio between the grain of habitats and the mean dispersal distance. The behavior of (dis)similarity indices varies. First, incidence-based measures of species overlap are less affected by habitat filtering than are abundance-based indices. Second, species identity–based indices, derived from population genetics’ FST, show interesting capacities to infer dispersal processes under neutrality but are also highly sensitive to habitat filtering. All indices except FST-related indices under neutrality are very sensitive to overall species richness. Hence, community patterns showing contrasted diversity levels should be compared with caution. Partitioning similarity indices within and between habitats appears to be an efficient approach to assess the strength of habitat filtering, and we show that a torus-translation test is powerful for this purpose. We finally highlight the need for further analytical studies to achieve theoretical expectations of similarity decay under dispersal and niche processes.

High-resolution forest mapping for behavioural studies in the Nature Reserve ‘Les Nouragues’, French Guiana

For animals with spatially complex behaviours at relatively small scales, the resolution of a global positioning system (GPS) receiver location is often below the resolution needed to correctly map animals’ spatial behaviour. Natural conditions such as canopy cover, canyons or clouds can further degrade GPS receiver reception. Here we present a detailed, high-resolution map of a 4.6 ha Neotropical river island and a 8.3 ha mainland plot with the location of every tree >5 cm DBH and all structures on the forest floor, which are relevant to our study species, the territorial frog Allobates femoralis (Dendrobatidae). The map was derived using distance- and compass-based survey techniques, rooted on dGPS reference points, and incorporates altitudinal information based on a LiDAR survey of the area.