Tree mode of death and mortality risk factors across Amazon forests.

The carbon sink capacity of tropical forests is substantially affected by tree mortality. However, the main drivers of tropical tree death remain largely unknown. Here we present a pan-Amazonian assessment of how and why trees die, analysing over 120,000 trees representing > 3800 species from 189 long-term RAINFOR forest plots. While tree mortality rates vary greatly Amazon-wide, on average trees are as likely to die standing as they are broken or uprooted-modes of death with different ecological consequences. Species-level growth rate is the single most important predictor of tree death in Amazonia, with faster-growing species being at higher risk. Within species, however, the slowest-growing trees are at greatest risk while the effect of tree size varies across the basin. In the driest Amazonian region species-level bioclimatic distributional patterns also predict the risk of death, suggesting that these forests are experiencing climatic conditions beyond their adaptative limits. These results provide not only a holistic pan-Amazonian picture of tree death but large-scale evidence for the overarching importance of the growth-survival trade-off in driving tropical tree mortality.

Long-term thermal sensitivity of Earth's tropical forests.

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.

The Forest Observation System, building a global reference dataset for remote sensing of forest biomass.

Forest biomass is an essential indicator for monitoring the Earth's ecosystems and climate. It is a critical input to greenhouse gas accounting, estimation of carbon losses and forest degradation, assessment of renewable energy potential, and for developing climate change mitigation policies such as REDD+, among others. Wall-to-wall mapping of aboveground biomass (AGB) is now possible with satellite remote sensing (RS). However, RS methods require extant, up-to-date, reliable, representative and comparable in situ data for calibration and validation. Here, we present the Forest Observation System (FOS) initiative, an international cooperation to establish and maintain a global in situ forest biomass database. AGB and canopy height estimates with their associated uncertainties are derived at a 0.25 ha scale from field measurements made in permanent research plots across the world's forests. All plot estimates are geolocated and have a size that allows for direct comparison with many RS measurements. The FOS offers the potential to improve the accuracy of RS-based biomass products while developing new synergies between the RS and ground-based ecosystem research communities.

Nutrient-cycling mechanisms other than the direct absorption from soil may control forest structure and dynamics in poor Amazonian soils.

Tropical forests store large amounts of biomass despite they generally grow in nutrient-poor soils, suggesting that the role of soil characteristics in the structure and dynamics of tropical forests is complex. We used data for >34 000 trees from several permanent plots in French Guiana to investigate if soil characteristics could predict the structure (tree diameter, density and aboveground biomass), and dynamics (growth, mortality, aboveground wood productivity) of nutrient-poor tropical forests. Most variables did not covary with site-level changes in soil nutrient content, indicating that nutrient-cycling mechanisms other than the direct absorption from soil (e.g. the nutrient uptake from litter, the resorption, or the storage of nutrients in the biomass), may strongly control forest structure and dynamics. Ecosystem-level adaptations to low soil nutrient availability and long-term low levels of disturbance may help to account for the lower productivity and higher accumulation of biomass in nutrient-poor forests compared to nutrient-richer forests.

Continuous soil carbon storage of old permanent pastures in Amazonia.

Amazonian forests continuously accumulate carbon (C) in biomass and in soil, representing a carbon sink of 0.42-0.65 GtC yr-1 . In recent decades, more than 15% of Amazonian forests have been converted into pastures, resulting in net C emissions (~200 tC ha-1 ) due to biomass burning and litter mineralization in the first years after deforestation. However, little is known about the capacity of tropical pastures to restore a C sink. Our study shows in French Amazonia that the C storage observed in native forest can be partly restored in old (≥24 year) tropical pastures managed with a low stocking rate (±1 LSU ha-1 ) and without the use of fire since their establishment. A unique combination of a large chronosequence study and eddy covariance measurements showed that pastures stored between -1.27 ± 0.37 and -5.31 ± 2.08 tC ha-1  yr-1 while the nearby native forest stored -3.31 ± 0.44 tC ha-1  yr-1 . This carbon is mainly sequestered in the humus of deep soil layers (20-100 cm), whereas no C storage was observed in the 0- to 20-cm layer. C storage in C4 tropical pasture is associated with the installation and development of C3 species, which increase either the input of N to the ecosystem or the C:N ratio of soil organic matter. Efforts to curb deforestation remain an obvious priority to preserve forest C stocks and biodiversity. However, our results show that if sustainable management is applied in tropical pastures coming from deforestation (avoiding fires and overgrazing, using a grazing rotation plan and a mixture of C3 and C4 species), they can ensure a continuous C storage, thereby adding to the current C sink of Amazonian forests.

Rapid tree carbon stock recovery in managed Amazonian forests.

While around 20% of the Amazonian forest has been cleared for pastures and agriculture, one fourth of the remaining forest is dedicated to wood production. Most of these production forests have been or will be selectively harvested for commercial timber, but recent studies show that even soon after logging, harvested stands retain much of their tree-biomass carbon and biodiversity. Comparing species richness of various animal taxa among logged and unlogged forests across the tropics, Burivalova et al. found that despite some variability among taxa, biodiversity loss was generally explained by logging intensity (the number of trees extracted). Here, we use a network of 79 permanent sample plots (376 ha total) located at 10 sites across the Amazon Basin to assess the main drivers of time-to-recovery of post-logging tree carbon (Table S1). Recovery time is of direct relevance to policies governing management practices (i.e., allowable volumes cut and cutting cycle lengths), and indirectly to forest-based climate change mitigation interventions.

An estimate of the number of tropical tree species.

The high species richness of tropical forests has long been recognized, yet there remains substantial uncertainty regarding the actual number of tropical tree species. Using a pantropical tree inventory database from closed canopy forests, consisting of 657,630 trees belonging to 11,371 species, we use a fitted value of Fisher's alpha and an approximate pantropical stem total to estimate the minimum number of tropical forest tree species to fall between ∼ 40,000 and ∼ 53,000, i.e., at the high end of previous estimates. Contrary to common assumption, the Indo-Pacific region was found to be as species-rich as the Neotropics, with both regions having a minimum of ∼ 19,000-25,000 tree species. Continental Africa is relatively depauperate with a minimum of ∼ 4,500-6,000 tree species. Very few species are shared among the African, American, and the Indo-Pacific regions. We provide a methodological framework for estimating species richness in trees that may help refine species richness estimates of tree-dependent taxa.

Drought-mortality relationships for tropical forests.

*The rich ecology of tropical forests is intimately tied to their moisture status. Multi-site syntheses can provide a macro-scale view of these linkages and their susceptibility to changing climates. Here, we report pan-tropical and regional-scale analyses of tree vulnerability to drought. *We assembled available data on tropical forest tree stem mortality before, during, and after recent drought events, from 119 monitoring plots in 10 countries concentrated in Amazonia and Borneo. *In most sites, larger trees are disproportionately at risk. At least within Amazonia, low wood density trees are also at greater risk of drought-associated mortality, independent of size. For comparable drought intensities, trees in Borneo are more vulnerable than trees in the Amazon. There is some evidence for lagged impacts of drought, with mortality rates remaining elevated 2 yr after the meteorological event is over. *These findings indicate that repeated droughts would shift the functional composition of tropical forests toward smaller, denser-wooded trees. At very high drought intensities, the linear relationship between tree mortality and moisture stress apparently breaks down, suggesting the existence of moisture stress thresholds beyond which some tropical forests would suffer catastrophic tree mortality.

Modeling decay rates of dead wood in a neotropical forest.

Variation of dead wood decay rates among tropical trees remains one source of uncertainty in global models of the carbon cycle. Taking advantage of a broad forest plot network surveyed for tree mortality over a 23-year period, we measured the remaining fraction of boles from 367 dead trees from 26 neotropical species widely varying in wood density (0.23-1.24 g cm(-3)) and tree circumference at death time (31.5-272.0 cm). We modeled decay rates within a Bayesian framework assuming a first order differential equation to model the decomposition process and tested for the effects of forest management (selective logging vs. unexploited), of mode of death (standing vs. downed) and of topographical levels (bottomlands vs. hillsides vs. hilltops) on wood decay rates. The general decay model predicts the observed remaining fraction of dead wood (R2 = 60%) with only two biological predictors: tree circumference at death time and wood specific density. Neither selective logging nor local topography had a differential effect on wood decay rates. Including the mode of death into the model revealed that standing dead trees decomposed faster than downed dead trees, but the gain of model accuracy remains rather marginal. Overall, these results suggest that the release of carbon from tropical dead trees to the atmosphere can be simply estimated using tree circumference at death time and wood density.

Dynamics of aboveground carbon stocks in a selectively logged tropical forest.

The expansion of selective logging in tropical forests may be an important source of global carbon emissions. However, the effects of logging practices on the carbon cycle have never been quantified over long periods of time. We followed the fate of more than 60 000 tropical trees over 23 years to assess changes in aboveground carbon stocks in 48 1.56-ha plots in French Guiana that represent a gradient of timber harvest intensities, with and without intensive timber stand improvement (TSI) treatments to stimulate timber tree growth. Conventional selective logging led to emissions equivalent to more than a third of aboveground carbon stocks in plots without TSI (85 Mg C/ha), while plots with TSI lost more than one-half of aboveground carbon stocks (142 Mg C/ha). Within 20 years of logging, plots without TSI sequestered aboveground carbon equivalent to more than 80% of aboveground carbon lost to logging (-70.7 Mg C/ha), and our simulations predicted an equilibrium aboveground carbon balance within 45 years of logging. In contrast, plots with intensive TSI are predicted to require more than 100 years to sequester aboveground carbon lost to emissions. These results indicate that in some tropical forests aboveground carbon storage can be recovered within half a century after conventional logging at moderate harvest intensities.

Drought sensitivity of the Amazon rainforest.

Amazon forests are a key but poorly understood component of the global carbon cycle. If, as anticipated, they dry this century, they might accelerate climate change through carbon losses and changed surface energy balances. We used records from multiple long-term monitoring plots across Amazonia to assess forest responses to the intense 2005 drought, a possible analog of future events. Affected forest lost biomass, reversing a large long-term carbon sink, with the greatest impacts observed where the dry season was unusually intense. Relative to pre-2005 conditions, forest subjected to a 100-millimeter increase in water deficit lost 5.3 megagrams of aboveground biomass of carbon per hectare. The drought had a total biomass carbon impact of 1.2 to 1.6 petagrams (1.2 x 10(15) to 1.6 x 10(15) grams). Amazon forests therefore appear vulnerable to increasing moisture stress, with the potential for large carbon losses to exert feedback on climate change.

Seasonal water stress tolerance and habitat associations within four neotropical tree genera.

We investigated the relationship between habitat association and physiological performance in four congeneric species pairs exhibiting contrasting distributions between seasonally flooded and terra firme habitats in lowland tropical rain forests of French Guiana, including Virola and Iryanthera (Myristicaceae), Symphonia (Clusiaceae), and Eperua (Caesalpiniaceae). We analyzed 10-year data sets of mapped and measured saplings (stems >150 cm in height and <10 cm diameter at breast height [dbh]) and trees (stems > or =10 cm dbh) across 37.5 ha of permanent plots covering a 300-ha zone, within which seasonally flooded areas (where the water table never descends below 1 m) have been mapped. Additionally, we tested the response of growth, survival, and leaf functional traits of these species to drought and flood stress in a controlled experiment. We tested for habitat preference using a modification of the torus translation method. Strong contrasting associations of the species pairs of Iryanthera, Virola, and Symphonia were observed at the sapling stage, and these associations strengthened for the tree stage. Neither species of Eperua was significantly associated with flooded habitats at the sapling stage, but E. falcata was significantly and positively associated with flooded forests at the tree stage, and trees of E. grandiflora were found almost exclusively in nonflooded habitats. Differential performance provided limited explanatory support for the observed habitat associations, with only congeners of Iryanthera exhibiting divergent sapling survival and tree growth. Seedlings of species associated with flooded forest tended to have higher photosynthetic capacity than their congeners at field capacity. In addition, they tended to have the largest reductions in leaf gas exchange and growth rate in response to experimental drought stress and the least reductions in response to experimental inundation. The corroboration of habitat association with differences in functional traits and, to a lesser extent, measures of performance provides an explanation for the regional coexistence of these species pairs. We suggest that specialization to seasonally flooded habitats may explain patterns of adaptive radiation in many tropical tree genera and thereby provide a substantial contribution to regional tree diversity.