Increasing biomass in Amazonian forest plots

A previous study by Phillips et al. of changes in the biomass of permanent sample plots in Amazonian forests was used to infer the presence of a regional carbon sink. However, these results generated a vigorous debate about sampling and methodological issues. Therefore we present a new analysis of biomass change in old-growth Amazonian forest plots using updated inventory data. We find that across 59 sites, the above-ground dry biomass in trees that are more than 10 cm in diameter (AGB) has increased since plot establishment by 1.22 ± 0.43 Mg per hectare per year (ha-1 yr-1), where 1 ha = 104 m2), or 0.98 ± 0.38 Mg ha-1 yr-1 if individual plot values are weighted by the number of hectare years of monitoring. This significant increase is neither confounded by spatial or temporal variation in wood specific gravity, nor dependent on the allometric equation used to estimate AGB. The conclusion is also robust to uncertainty about diameter measurements for problematic trees: for 34 plots in western Amazon forests a significant increase in AGB is found even with a conservative assumption of zero growth for all trees where diameter measurements were made using optical methods and/or growth rates needed to be estimated following fieldwork. Overall, our results suggest a slightly greater rate of net stand-level change than was reported by Phillips et al. Considering the spatial and temporal scale of sampling and associated studies showing increases in forest growth and stem turnover, the results presented here suggest that the total biomass of these plots has on average increased and that there has been a regional-scale carbon sink in old-growth Amazonian forests during the previous two decades

Soil pyrogenic carbon in southern Amazonia: Interaction between soil, climate, and above-ground biomass

The Amazon forest represents one of the world’s largest terrestrial carbon reservoirs. Here, we evaluated the role of soil texture, climate, vegetation, and distance to savanna on the distribution and stocks of soil pyrogenic carbon (PyC) in intact forests with no history of recent fire spanning the southern Amazonia forest-Cerrado Zone of Transition (ZOT). In 19 one hectare forest plots, including three Amazonian Dark Earth (ADE, terra preta) sites with high soil PyC, we measured all trees and lianas with diameter ≥ 10 cm and analyzed soil physicochemical properties, including texture and PyC stocks. We quantified PyC stocks as a proportion of total organic carbon using hydrogen pyrolysis. We used multiple linear regression and variance partitioning to determine which variables best explain soil PyC variation. For all forests combined, soil PyC stocks ranged between 0.9 and 6.8 Mg/ha to 30 cm depth (mean 2.3 ± 1.5 Mg/ha) and PyC, on average, represented 4.3% of the total soil organic carbon (SOC). The most parsimonious model (based on AICc) included soil clay content and above-ground biomass (AGB) as the main predictors, explaining 71% of soil PyC variation. After removal of the ADE plots, PyC stocks ranged between 0.9 and 3.8 Mg/ha (mean 1.9 ± 0.8 Mg/ha–1) and PyC continued to represent ∼4% of the total SOC. The most parsimonious models without ADE included AGB and sand as the best predictors, with sand and PyC having an inverse relationship, and sand explaining 65% of the soil PyC variation. Partial regression analysis did not identify any of the components (climatic, environmental, and edaphic), pure or shared, as important in explaining soil PyC variation with or without ADE plots. We observed a substantial amount of soil PyC, even excluding ADE forests; however, contrary to expectations, soil PyC stocks were not higher nearer to the fire-dependent Cerrado than more humid regions of Amazonia. Our findings that soil texture and AGB explain the distribution and amount of soil PyC in ZOT forests will help to improve model estimates of SOC change with further climatic warming

Increasing dominance of large lianas in Amazonian forests

Ecological orthodoxy suggests that old-growth forests should be close to dynamic equilibrium, but this view has been challenged by recent findings that neotropical forests are accumulating carbon and biomass, possibly in response to the increasing atmospheric concentrations of carbon dioxide. However, it is unclear whether the recent increase in tree biomass has been accompanied by a shift in community composition. Such changes could reduce or enhance the carbon storage potential of old-growth forests in the long term. Here we show that non-fragmented Amazon forests are experiencing a concerted increase in the density, basal area and mean size of woody climbing plants (lianas). Over the last two decades of the twentieth century the dominance of large lianas relative to trees has increased by 1.7–4.6% a year. Lianas enhance tree mortality and suppress tree growth, so their rapid increase implies that the tropical terrestrial carbon sink may shut down sooner than current models suggest. Predictions of future tropical carbon fluxes will need to account for the changing composition and dynamics of supposedly undisturbed forests

Rarity of monodominance in hyperdiverse Amazonian forests.

Tropical forests are known for their high diversity. Yet, forest patches do occur in the tropics where a single tree species is dominant. Such "monodominant" forests are known from all of the main tropical regions. For Amazonia, we sampled the occurrence of monodominance in a massive, basin-wide database of forest-inventory plots from the Amazon Tree Diversity Network (ATDN). Utilizing a simple defining metric of at least half of the trees ≥ 10 cm diameter belonging to one species, we found only a few occurrences of monodominance in Amazonia, and the phenomenon was not significantly linked to previously hypothesized life history traits such wood density, seed mass, ectomycorrhizal associations, or Rhizobium nodulation. In our analysis, coppicing (the formation of sprouts at the base of the tree or on roots) was the only trait significantly linked to monodominance. While at specific locales coppicing or ectomycorrhizal associations may confer a considerable advantage to a tree species and lead to its monodominance, very few species have these traits. Mining of the ATDN dataset suggests that monodominance is quite rare in Amazonia, and may be linked primarily to edaphic factors

MODIS Vegetation Continuous Fields tree cover needs calibrating in tropical savannas

The Moderate Resolution Imaging Spectroradiometer Vegetation Continuous Fields (MODIS VCF) Earth observation product is widely used to estimate forest cover changes and to parameterize vegetation and Earth system models and as a reference for validation or calibration where field data are limited. However, although limited independent validations of MODIS VCF have shown that MODIS VCF's accuracy decreases when estimating tree cover in sparsely vegetated areas such as tropical savannas, no study has yet assessed the impact this may have on the VCF-based tree cover data used by many in their research. Using tropical forest and savanna inventory data collected by the Tropical Biomes in Transition (TROBIT) project, we produce a series of calibration scenarios that take into account (i) the spatial disparity between the in situ plot size and the MODIS VCF pixel and (ii) the trees' spatial distribution within in situ plots. To identify if a disparity also exists in products trained using VCF, we used a similar approach to evaluate the finer-scale Landsat Tree Canopy Cover (TCC) product. For MODIS VCF, we then applied our calibrations to areas identified as forest or savanna in the International Geosphere-Biosphere Programme (IGBP) land cover mapping product. All IGBP classes identified as “savanna” show substantial increases in cover after calibration, indicating that the most recent version of MODIS VCF consistently underestimates woody cover in tropical savannas. We also found that these biases are propagated in the finer-scale Landsat TCC. Our scenarios suggest that MODIS VCF accuracy can vary substantially, with tree cover underestimation ranging from 0 % to 29 %. Models that use MODIS VCF as their benchmark could therefore be underestimating the carbon uptake in forest–savanna areas and misrepresenting forest–savanna dynamics. Because of the limited in situ plot number, our results are designed to be used as an indicator of where the product is potentially more or less reliable. Until more in situ data are available to produce more accurate calibrations, we recommend caution when using uncalibrated MODIS VCF data in tropical savannas

Evenness mediates the global relationship between forest productivity and richness

1. Biodiversity is an important component of natural ecosystems, with higher species richness often correlating with an increase in ecosystem productivity. Yet, this relationship varies substantially across environments, typically becoming less pronounced at high levels of species richness. However, species richness alone cannot reflect all important properties of a community, including community evenness, which may mediate the relationship between biodiversity and productivity. If the evenness of a community correlates negatively with richness across forests globally, then a greater number of species may not always increase overall diversity and productivity of the system. Theoretical work and local empirical studies have shown that the effect of evenness on ecosystem functioning may be especially strong at high richness levels, yet the consistency of this remains untested at a global scale. 2. Here, we used a dataset of forests from across the globe, which includes composition, biomass accumulation and net primary productivity, to explore whether productivity correlates with community evenness and richness in a way that evenness appears to buffer the effect of richness. Specifically, we evaluated whether low levels of evenness in speciose communities correlate with the attenuation of the richness–productivity relationship. 3. We found that tree species richness and evenness are negatively correlated across forests globally, with highly speciose forests typically comprising a few dominant and many rare species. Furthermore, we found that the correlation between diversity and productivity changes with evenness: at low richness, uneven communities are more productive, while at high richness, even communities are more productive. 4. Synthesis. Collectively, these results demonstrate that evenness is an integral component of the relationship between biodiversity and productivity, and that the attenuating effect of richness on forest productivity might be partly explained by low evenness in speciose communities. Productivity generally increases with species richness, until reduced evenness limits the overall increases in community diversity. Our research suggests that evenness is a fundamental component of biodiversity– ecosystem function relationships, and is of critical importance for guiding conservation and sustainable ecosystem management decisions

Aboveground forest biomass varies across continents, ecological zones and successional stages: refined IPCC default values for tropical and subtropical forests

For monitoring and reporting forest carbon stocks and fluxes, many countries in the tropics and subtropics rely on default values of forest aboveground biomass (AGB) from the Intergovernmental Panel on Climate Change (IPCC) guidelines for National Greenhouse Gas (GHG) Inventories. Default IPCC forest AGB values originated from 2006, and are relatively crude estimates of average values per continent and ecological zone. The 2006 default values were based on limited plot data available at the time, methods for their derivation were not fully clear, and no distinction between successional stages was made. As part of the 2019 Refinement to the 2006 IPCC Guidelines for GHG Inventories, we updated the default AGB values for tropical and subtropical forests based on AGB data from >25 000 plots in natural forests and a global AGB map where no plot data were available. We calculated refined AGB default values per continent, ecological zone, and successional stage, and provided a measure of uncertainty. AGB in tropical and subtropical forests varies by an order of magnitude across continents, ecological zones, and successional stage. Our refined default values generally reflect the climatic gradients in the tropics, with more AGB in wetter areas. AGB is generally higher in old-growth than in secondary forests, and higher in older secondary (regrowth >20 years old and degraded/logged forests) than in young secondary forests (20 years old). While refined default values for tropical old-growth forest are largely similar to the previous 2006 default values, the new default values are 4.0-7.7-fold lower for young secondary forests. Thus, the refined values will strongly alter estimated carbon stocks and fluxes, and emphasize the critical importance of old-growth forest conservation. We provide a reproducible approach to facilitate future refinements and encourage targeted efforts to establish permanent plots in areas with data gaps

Expanding tropical forest monitoring into Dry Forests: The DRYFLOR protocol for permanent plots

This is the final version. Available on open access from Wiley via the DOI in this recordSocietal Impact Statement Understanding of tropical forests has been revolutionized by monitoring in permanent plots. Data from global plot networks have transformed our knowledge of forests’ diversity, function, contribution to global biogeochemical cycles, and sensitivity to climate change. Monitoring has thus far been concentrated in rain forests. Despite increasing appreciation of their threatened status, biodiversity, and importance to the global carbon cycle, monitoring in tropical dry forests is still in its infancy. We provide a protocol for permanent monitoring plots in tropical dry forests. Expanding monitoring into dry biomes is critical for overcoming the linked challenges of climate change, land use change, and the biodiversity crisis.Newton FundNatural Environment Research Council (NERC)Fundação de Amparo à Pesquisa do Estado de São PauloCYTE

Variation in stem mortality rates determines patterns of above-ground biomass in Amazonian forests: implications for dynamic global vegetation models

Understanding the processes that determine above-ground biomass (AGB) in Amazonian forests is important for predicting the sensitivity of these ecosystems to environmental change and for designing and evaluating dynamic global vegetation models (DGVMs). AGB is determined by inputs from woody productivity [woody net primary productivity (NPP)] and the rate at which carbon is lost through tree mortality. Here, we test whether two direct metrics of tree mortality (the absolute rate of woody biomass loss and the rate of stem mortality) and/or woody NPP, control variation in AGB among 167 plots in intact forest across Amazonia. We then compare these relationships and the observed variation in AGB and woody NPP with the predictions of four DGVMs. The observations show that stem mortality rates, rather than absolute rates of woody biomass loss, are the most important predictor of AGB, which is consistent with the importance of stand size structure for determining spatial variation in AGB. The relationship between stem mortality rates and AGB varies among different regions of Amazonia, indicating that variation in wood density and height/diameter relationships also influences AGB. In contrast to previous findings, we find that woody NPP is not correlated with stem mortality rates and is weakly positively correlated with AGB. Across the four models, basin-wide average AGB is similar to the mean of the observations. However, the models consistently overestimate woody NPP and poorly represent the spatial patterns of both AGB and woody NPP estimated using plot data. In marked contrast to the observations, DGVMs typically show strong positive relationships between woody NPP and AGB. Resolving these differences will require incorporating forest size structure, mechanistic models of stem mortality and variation in functional composition in DGVMs