Leaf thermotolerance of Hevea brasiliensis clones: intra- versus interclonal variation and relationships with other functional traits.

Land surface temperature is predicted to increase by 0.2 °C per decade due to climate change, although with considerable regional variability, and heatwaves are predicted to increase markedly in the future. These changes will affect where crops can be grown in the future. Understanding the thermal limits of plant physiological functioning and how flexible such limits are is thus important. Here, we report on the measurements of a core foliar thermotolerance trait, T50, defined as the temperature at which the maximum quantum yield (Fv/Fm) of photosystem II declines by 50%, across nine different Malaysian Hevea brasiliensis clones. We explore the relative importance of interclonal versus intraclonal variation in T50 as well as its association with leaf and hydraulic traits. We find very low variation in T50 within individual clones (mean intraclonal coefficient of variation (CoV) of 1.26%) and little variation across clones (interclonal CoV of 2.1%). The interclonal variation in T50 was lower than for all other functional traits considered. The T50 was negatively related to leaf mass per area and leaf dry matter content, but it was not related to hydraulic traits such as embolism resistance (P50) or hydraulic safety margins (HSM50). The range of T50 observed (42.9-46.2 °C) is well above the current maximum air temperatures Tmax,obs (T50 - Tmax,obs >5.8 °C), suggesting that H. brasiliensis is likely thermally safe in this south-east Asian region of Malaysia.

Tropical forests are approaching critical temperature thresholds.

The critical temperature beyond which photosynthetic machinery in tropical trees begins to fail averages approximately 46.7 °C (Tcrit)1. However, it remains unclear whether leaf temperatures experienced by tropical vegetation approach this threshold or soon will under climate change. Here we found that pantropical canopy temperatures independently triangulated from individual leaf thermocouples, pyrgeometers and remote sensing (ECOSTRESS) have midday peak temperatures of approximately 34 °C during dry periods, with a long high-temperature tail that can exceed 40 °C. Leaf thermocouple data from multiple sites across the tropics suggest that even within pixels of moderate temperatures, upper canopy leaves exceed Tcrit 0.01% of the time. Furthermore, upper canopy leaf warming experiments (+2, 3 and 4 °C in Brazil, Puerto Rico and Australia, respectively) increased leaf temperatures non-linearly, with peak leaf temperatures exceeding Tcrit 1.3% of the time (11% for more than 43.5 °C, and 0.3% for more than 49.9 °C). Using an empirical model incorporating these dynamics (validated with warming experiment data), we found that tropical forests can withstand up to a 3.9 ± 0.5 °C increase in air temperatures before a potential tipping point in metabolic function, but remaining uncertainty in the plasticity and range of Tcrit in tropical trees and the effect of leaf death on tree death could drastically change this prediction. The 4.0 °C estimate is within the 'worst-case scenario' (representative concentration pathway (RCP) 8.5) of climate change predictions2 for tropical forests and therefore it is still within our power to decide (for example, by not taking the RCP 6.0 or 8.5 route) the fate of these critical realms of carbon, water and biodiversity3,4.

Anatomical functional traits and hydraulic vulnerability of trees in different water conditions in southern Amazonia.

PREMISE: Understanding tree species' responses to drought is critical for predicting the future of tropical forests, especially in regions where the climate is changing rapidly. METHODS: We compared anatomical and functional traits of the dominant tree species of two tropical forests in southern Amazonia, one on deep, well-drained soils (cerradão [CD]) and one in a riparian environment (gallery forest [GF]), to examine potential anatomical indicators of resistance or vulnerability to drought. RESULTS: Leaves of CD species generally had a thicker cuticle, upper epidermis, and mesophyll than those of GF species, traits that are indicative of adaptation to water deficit. In the GF, the theoretical hydraulic conductivity of the stems was significantly higher, indicating lower investment in drought resistance. The anatomical functional traits of CD species indicate a greater potential for surviving water restriction compared to the GF. Even so, it is possible that CD species could also be affected by extreme climate changes due to the more water-limited environment. CONCLUSIONS: In addition to the marked anatomical and functional differences between these phytophysiognomies, tree diversity within each is associated with a large range of hydraulic morphofunctional niches. Our results suggest the strong potential for floristic and functional compositional shifts under continued climate change, especially in the GF.

Long-term drought effects on the thermal sensitivity of Amazon forest trees.

The continued functioning of tropical forests under climate change depends on their resilience to drought and heat. However, there is little understanding of how tropical forests will respond to combinations of these stresses, and no field studies to date have explicitly evaluated whether sustained drought alters sensitivity to temperature. We measured the temperature response of net photosynthesis, foliar respiration and the maximum quantum efficiency of photosystem II (Fv /Fm ) of eight hyper-dominant Amazonian tree species at the world's longest-running tropical forest drought experiment, to investigate the effect of drought on forest thermal sensitivity. Despite a 0.6°C-2°C increase in canopy air temperatures following long-term drought, no change in overall thermal sensitivity of net photosynthesis or respiration was observed. However, photosystem II tolerance to extreme-heat damage (T50 ) was reduced from 50.0 ± 0.3°C to 48.5 ± 0.3°C under drought. Our results suggest that long-term reductions in precipitation, as projected across much of Amazonia by climate models, are unlikely to greatly alter the response of tropical forests to rising mean temperatures but may increase the risk of leaf thermal damage during heatwaves.

Reassessment of carbon emissions from fires and a new estimate of net carbon uptake in Russian forests in 2001-2021.

Russia has the largest forest area on earth. Its boreal forests officially store about 97 Pg C, which significantly affect the global carbon cycle. In recent years, forest fires have been intensifying on the planet, leading to increased carbon emissions. Here we review how differences in fire control management of Russian forests affect fire related emissions. Carbon emissions due to fire were estimated using satellite data and compared to official reports for 2001-2021. We found that the relative areas affected by fire did differ between different fire protection zones, and 89 % of the area burnt was in forests controlled by fire-fighting aircraft or areas without protection. As a result, 417.7 Mha of poor or unprotected Russian forests (42 % of total) account about a half of total carbon emissions. According to our estimates, the average area of burnt forests in Russia was about 8.3 Mha per year between 2016 and 2021, resulting in annual carbon emission of 193 million metric tons (Mt) C emissions, and 53 % of them were from unprotected forest. These estimated carbon emissions are significantly higher than official national reports (79 Mt C yr-1). We estimated that net carbon uptake for Russia for 2015-2021 was about 333 ± 37 Mt C, which is roughly double the official estimates. Our results highlight large spatial differences in fire protection and prevention strategies in fire related emissions. The so-called control zone which stretches across large parts of Eastern Russia has no fire control and is the region of major recent fires. Our study shows that to estimate the Russian forest carbon balance it is critical to include this area. Implementation of some forest management in the remote areas (i.e., control zone) would help to decrease forest loss and resulting carbon emissions.

A novel in situ passive heating method for evaluating whole-tree responses to daytime warming in remote environments.

BACKGROUND: Many significant ecosystems, including important non-forest woody ecosystems such as the Cerrado (Brazilian savannah), are under threat from climate change, yet our understanding of how increasing temperatures will impact native vegetation remains limited. Temperature manipulation experiments are important tools for investigating such impacts, but are often constrained by access to power supply and limited to low-stature species, juvenile individuals, or heating of target organs, perhaps not fully revealing how entire or mature individuals and ecosystems will react to higher temperatures. RESULTS: We present a novel, modified open top chamber design for in situ passive heating of whole individuals up to 2.5 m tall (but easily expandable) in remote field environments with strong solar irradiance. We built multiple whole-tree heating structures (WTHSs) in an area of Cerrado around native woody species Davilla elliptica and Erythroxylum suberosum to test the design and its effects on air temperature and humidity, while also studying the physiological responses of E. suberosum to short-term heating. The WTHSs raised internal air temperature by approximately 2.5 °C above ambient during the daytime. This increased to 3.4 °C between 09:00 and 17:00 local time when thermal impact was greatest, and during which time mean internal temperatures corresponded closely with maximum ambient temperatures. Heating was consistent over time and across WTHSs of variable size and shape, and they had minimal effect on humidity. E. suberosum showed no detectable response of photosynthesis or respiration to short-term experimental heating, but some indication of acclimation to natural temperature changes. CONCLUSIONS: Our WTHSs produced a consistent and reproducible level of daytime heating in line with mid-range climate predictions for the Cerrado biome by the end of the century. The whole-tree in situ passive heating design is flexible, low-cost, simple to build using commonly available materials, and minimises negative impacts associated with passive chambers. It could be employed to investigate the high temperature responses of many understudied species in a range of complex non-forest environments with sufficient solar irradiance, providing new and important insights into the possible impacts of our changing climate.

Non-structural carbohydrates mediate seasonal water stress across Amazon forests.

Non-structural carbohydrates (NSC) are major substrates for plant metabolism and have been implicated in mediating drought-induced tree mortality. Despite their significance, NSC dynamics in tropical forests remain little studied. We present leaf and branch NSC data for 82 Amazon canopy tree species in six sites spanning a broad precipitation gradient. During the wet season, total NSC (NSCT) concentrations in both organs were remarkably similar across communities. However, NSCT and its soluble sugar (SS) and starch components varied much more across sites during the dry season. Notably, the proportion of leaf NSCT in the form of SS (SS:NSCT) increased greatly in the dry season in almost all species in the driest sites, implying an important role of SS in mediating water stress in these sites. This adjustment of leaf NSC balance was not observed in tree species less-adapted to water deficit, even under exceptionally dry conditions. Thus, leaf carbon metabolism may help to explain floristic sorting across water availability gradients in Amazonia and enable better prediction of forest responses to future climate change.

Large-scale variations in the dynamics of Amazon forest canopy gaps from airborne lidar data and opportunities for tree mortality estimates.

We report large-scale estimates of Amazonian gap dynamics using a novel approach with large datasets of airborne light detection and ranging (lidar), including five multi-temporal and 610 single-date lidar datasets. Specifically, we (1) compared the fixed height and relative height methods for gap delineation and established a relationship between static and dynamic gaps (newly created gaps); (2) explored potential environmental/climate drivers explaining gap occurrence using generalized linear models; and (3) cross-related our findings to mortality estimates from 181 field plots. Our findings suggest that static gaps are significantly correlated to dynamic gaps and can inform about structural changes in the forest canopy. Moreover, the relative height outperformed the fixed height method for gap delineation. Well-defined and consistent spatial patterns of dynamic gaps were found over the Amazon, while also revealing the dynamics of areas never sampled in the field. The predominant pattern indicates 20-35% higher gap dynamics at the west and southeast than at the central-east and north. These estimates were notably consistent with field mortality patterns, but they showed 60% lower magnitude likely due to the predominant detection of the broken/uprooted mode of death. While topographic predictors did not explain gap occurrence, the water deficit, soil fertility, forest flooding and degradation were key drivers of gap variability at the regional scale. These findings highlight the importance of lidar in providing opportunities for large-scale gap dynamics and tree mortality monitoring over the Amazon.

Photosynthetic quantum efficiency in south-eastern Amazonian trees may be already affected by climate change.

Tropical forests are experiencing unprecedented high-temperature conditions due to climate change that could limit their photosynthetic functions. We studied the high-temperature sensitivity of photosynthesis in a rainforest site in southern Amazonia, where some of the highest temperatures and most rapid warming in the Tropics have been recorded. The quantum yield (Fv /Fm ) of photosystem II was measured in seven dominant tree species using leaf discs exposed to varying levels of heat stress. T50 was calculated as the temperature at which Fv /Fm was half the maximum value. T5 is defined as the breakpoint temperature, at which Fv /Fm decline was initiated. Leaf thermotolerance in the rapidly warming southern Amazonia was the highest recorded for forest tree species globally. T50 and T5 varied between species, with one mid-storey species, Amaioua guianensis, exhibiting particularly high T50 and T5 values. While the T50 values of the species sampled were several degrees above the maximum air temperatures experienced in southern Amazonia, the T5 values of several species are now exceeded under present-day maximum air temperatures.

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.

Intra-annual oxygen isotopes in the tree rings record precipitation extremes and water reservoir levels in the Metropolitan Area of São Paulo, Brazil.

The impacts of climate change on precipitation and the growing demand for water have increased the water risks worldwide. Water scarcity is one of the main challenges of the 21st century, and the assessment of water risks is only possible from spatially distributed records of historical climate and levels of water reservoirs. One potential method to assess water supply is the reconstruction of oxygen isotopes in rainfall. We here investigated the use of tree-ring stable isotopes in urban trees to assess spatial/temporal variation in precipitation and level of water reservoirs. We analyzed the intra-annual variation of δ13C and δ18O in the tree rings of Tipuana tipu trees from northern and southern Metropolitan Area of São Paulo (MASP), Brazil. While variation in δ13C indicates low leaf-level enrichments from evapotranspiration, δ18O variation clearly reflects precipitation extremes. Tree-ring δ18O was highest during the 2014 drought, associated with the lowest historical reservoir levels in the city. The δ18O values from the middle of the tree rings have a strong association with the mid-summer precipitation (r = -0.71), similar to the association between the volume of precipitation and its δ18O signature (r = -0.76). These consistent results allowed us to test the association between tree-ring δ18O and water-level of the main reservoirs that supply the MASP. We observed a strong association between intra-annual tree-ring δ18O and the water-level of reservoirs in the northern and southern MASP (r = -0.94, r = -0.90, respectively). These results point to the potential use of high-resolution tree-ring stable isotopes to put precipitation extremes, and water supply, in a historical perspective assisting public policies related to water risks and climate change. The ability to record precipitation extremes, and previously reported capacity to record air pollution, place Tipuana tipu in a prominent position as a reliable environmental monitor for urban locations.

Asynchronous carbon sink saturation in African and Amazonian tropical forests.

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 emissions1-3. 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 temperature7-9. 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.

Mapping Atlantic rainforest degradation and regeneration history with indicator species using convolutional network.

The Atlantic rainforest of Brazil is one of the global terrestrial hotspots of biodiversity. Despite having undergone large scale deforestation, forest cover has shown signs of increases in the last decades. Here, to understand the degradation and regeneration history of Atlantic rainforest remnants near São Paulo, we combine a unique dataset of very high resolution images from Worldview-2 and Worldview-3 (0.5 and 0.3m spatial resolution, respectively), georeferenced aerial photographs from 1962 and use a deep learning method called U-net to map (i) the forest cover and changes and (ii) two pioneer tree species, Cecropia hololeuca and Tibouchina pulchra. For Tibouchina pulchra, all the individuals were mapped in February, when the trees undergo mass-flowering with purple and pink blossoms. Additionally, elevation data at 30m spatial resolution from NASA Shuttle Radar Topography Mission (SRTM) and annual mean climate variables (Terraclimate datasets at ∼ 4km of spatial resolution) were used to analyse the forest and species distributions. We found that natural forests are currently more frequently found on south-facing slopes, likely because of geomorphology and past land use, and that Tibouchina is restricted to the wetter part of the region (southern part), which annually receives at least 1600 mm of precipitation. Tibouchina pulchra was found to clearly indicate forest regeneration as almost all individuals were found within or adjacent to forests regrown after 1962. By contrast, Cecropia hololeuca was found to indicate older disturbed forests, with all individuals almost exclusively found in forest fragments already present in 1962. At the regional scale, using the dominance maps of both species, we show that at least 4.3% of the current region's natural forests have regrown after 1962 (Tibouchina dominated, ∼ 4757 ha) and that ∼ 9% of the old natural forests have experienced significant disturbance (Cecropia dominated).

Evolutionary diversity is associated with wood productivity in Amazonian forests.

Higher levels of taxonomic and evolutionary diversity are expected to maximize ecosystem function, yet their relative importance in driving variation in ecosystem function at large scales in diverse forests is unknown. Using 90 inventory plots across intact, lowland, terra firme, Amazonian forests and a new phylogeny including 526 angiosperm genera, we investigated the association between taxonomic and evolutionary metrics of diversity and two key measures of ecosystem function: aboveground wood productivity and biomass storage. While taxonomic and phylogenetic diversity were not important predictors of variation in biomass, both emerged as independent predictors of wood productivity. Amazon forests that contain greater evolutionary diversity and a higher proportion of rare species have higher productivity. While climatic and edaphic variables are together the strongest predictors of productivity, our results show that the evolutionary diversity of tree species in diverse forest stands also influences productivity. As our models accounted for wood density and tree size, they also suggest that additional, unstudied, evolutionarily correlated traits have significant effects on ecosystem function in tropical forests. Overall, our pan-Amazonian analysis shows that greater phylogenetic diversity translates into higher levels of ecosystem function: tropical forest communities with more distantly related taxa have greater wood productivity.

Compositional response of Amazon forests to climate change.

Most of the planet's diversity is concentrated in the tropics, which includes many regions undergoing rapid climate change. Yet, while climate-induced biodiversity changes are widely documented elsewhere, few studies have addressed this issue for lowland tropical ecosystems. Here we investigate whether the floristic and functional composition of intact lowland Amazonian forests have been changing by evaluating records from 106 long-term inventory plots spanning 30 years. We analyse three traits that have been hypothesized to respond to different environmental drivers (increase in moisture stress and atmospheric CO2 concentrations): maximum tree size, biogeographic water-deficit affiliation and wood density. Tree communities have become increasingly dominated by large-statured taxa, but to date there has been no detectable change in mean wood density or water deficit affiliation at the community level, despite most forest plots having experienced an intensification of the dry season. However, among newly recruited trees, dry-affiliated genera have become more abundant, while the mortality of wet-affiliated genera has increased in those plots where the dry season has intensified most. Thus, a slow shift to a more dry-affiliated Amazonia is underway, with changes in compositional dynamics (recruits and mortality) consistent with climate-change drivers, but yet to significantly impact whole-community composition. The Amazon observational record suggests that the increase in atmospheric CO2 is driving a shift within tree communities to large-statured species and that climate changes to date will impact forest composition, but long generation times of tropical trees mean that biodiversity change is lagging behind climate change.

Tropical land carbon cycle responses to 2015/16 El Niño as recorded by atmospheric greenhouse gas and remote sensing data.

The outstanding tropical land climate characteristic over the past decades is rapid warming, with no significant large-scale precipitation trends. This warming is expected to continue but the effects on tropical vegetation are unknown. El Niño-related heat peaks may provide a test bed for a future hotter world. Here we analyse tropical land carbon cycle responses to the 2015/16 El Niño heat and drought anomalies using an atmospheric transport inversion. Based on the global atmospheric CO2 and fossil fuel emission records, we find no obvious signs of anomalously large carbon release compared with earlier El Niño events, suggesting resilience of tropical vegetation. We find roughly equal net carbon release anomalies from Amazonia and tropical Africa, approximately 0.5 PgC each, and smaller carbon release anomalies from tropical East Asia and southern Africa. Atmospheric CO anomalies reveal substantial fire carbon release from tropical East Asia peaking in October 2015 while fires contribute only a minor amount to the Amazonian carbon flux anomaly. Anomalously large Amazonian carbon flux release is consistent with downregulation of primary productivity during peak negative near-surface water anomaly (October 2015 to March 2016) as diagnosed by solar-induced fluorescence. Finally, we find an unexpected anomalous positive flux to the atmosphere from tropical Africa early in 2016, coincident with substantial CO release.This article is part of a discussion meeting issue 'The impact of the 2015/2016 El Niño on the terrestrial tropical carbon cycle: patterns, mechanisms and implications'.

Recent intensification of Amazon flooding extremes driven by strengthened Walker circulation.

The Amazon basin is the largest watershed on Earth. Although the variability of the Amazon hydrological cycle has been increasing since the late 1990s, its underlying causes have remained elusive. We use water levels in the Amazon River to quantify changes in extreme events and then analyze their cause. Despite continuing research emphasis on droughts, the largest change over recent decades is a marked increase in very severe floods. Increased flooding is linked to a strengthening of the Walker circulation, resulting from strong tropical Atlantic warming and tropical Pacific cooling. Atlantic warming due to combined anthropogenic and natural factors has contributed to enhance the change in atmospheric circulation. Whether this anomalous increase in flooding will last depends on the evolution of the tropical inter-ocean temperature difference.

Pervasive Rise of Small-scale Deforestation in Amazonia.

Understanding forest loss patterns in Amazonia, the Earth's largest rainforest region, is critical for effective forest conservation and management. Following the most detailed analysis to date, spanning the entire Amazon and extending over a 14-year period (2001-2014), we reveal significant shifts in deforestation dynamics of Amazonian forests. Firstly, hotspots of Amazonian forest loss are moving away from the southern Brazilian Amazon to Peru and Bolivia. Secondly, while the number of new large forest clearings (>50 ha) has declined significantly over time (46%), the number of new small clearings (<1 ha) increased by 34% between 2001-2007 and 2008-2014. Thirdly, we find that small-scale low-density forest loss expanded markedly in geographical extent during 2008-2014. This shift presents an important and alarming new challenge for forest conservation, despite reductions in overall deforestation rates.

Erratum: Large emissions from floodplain trees close the Amazon methane budget.

This corrects the article DOI: 10.1038/nature24639.