Rainforest transformation reallocates energy from green to brown food webs.

Terrestrial animal biodiversity is increasingly being lost because of land-use change1,2. However, functional and energetic consequences aboveground and belowground and across trophic levels in megadiverse tropical ecosystems remain largely unknown. To fill this gap, we assessed changes in energy fluxes across 'green' aboveground (canopy arthropods and birds) and 'brown' belowground (soil arthropods and earthworms) animal food webs in tropical rainforests and plantations in Sumatra, Indonesia. Our results showed that most of the energy in rainforests is channelled to the belowground animal food web. Oil palm and rubber plantations had similar or, in the case of rubber agroforest, higher total animal energy fluxes compared to rainforest but the key energetic nodes were distinctly different: in rainforest more than 90% of the total animal energy flux was channelled by arthropods in soil and canopy, whereas in plantations more than 50% of the energy was allocated to annelids (earthworms). Land-use change led to a consistent decline in multitrophic energy flux aboveground, whereas belowground food webs responded with reduced energy flux to higher trophic levels, down to -90%, and with shifts from slow (fungal) to fast (bacterial) energy channels and from faeces production towards consumption of soil organic matter. This coincides with previously reported soil carbon stock depletion3. Here we show that well-documented animal biodiversity declines with tropical land-use change4-6 are associated with vast energetic and functional restructuring in food webs across aboveground and belowground ecosystem compartments.

Valley formation aridifies East Africa and elevates Congo Basin rainfall.

East African aridification during the past 8 million years is frequently invoked as a driver of large-scale shifts in vegetation1 and the evolution of new animal lineages, including hominins2-4. However, evidence for increasing aridity is debated5 and, crucially, the mechanisms leading to dry conditions are unclear6. Here, numerical model experiments show that valleys punctuating the 6,000-km-long East African Rift System (EARS) are central to the development of dry conditions in East Africa. These valleys, including the Turkana Basin in Kenya, cause East Africa to dry by channelling water vapour towards Central Africa, a process that simultaneously enhances rainfall in the Congo Basin rainforest. Without the valleys, the uplift of the rift system leads to a wetter climate in East Africa and a drier climate in the Congo Basin. Results from climate model experiments demonstrate that the detailed tectonic development of Africa has shaped the rainfall distribution, with profound implications for the evolution of African plant and animal lineages.

The carbon sink of secondary and degraded humid tropical forests.

The globally important carbon sink of intact, old-growth tropical humid forests is declining because of climate change, deforestation and degradation from fire and logging1-3. Recovering tropical secondary and degraded forests now cover about 10% of the tropical forest area4, but how much carbon they accumulate remains uncertain. Here we quantify the aboveground carbon (AGC) sink of recovering forests across three main continuous tropical humid regions: the Amazon, Borneo and Central Africa5,6. On the basis of satellite data products4,7, our analysis encompasses the heterogeneous spatial and temporal patterns of growth in degraded and secondary forests, influenced by key environmental and anthropogenic drivers. In the first 20 years of recovery, regrowth rates in Borneo were up to 45% and 58% higher than in Central Africa and the Amazon, respectively. This is due to variables such as temperature, water deficit and disturbance regimes. We find that regrowing degraded and secondary forests accumulated 107 Tg C year-1 (90-130 Tg C year-1) between 1984 and 2018, counterbalancing 26% (21-34%) of carbon emissions from humid tropical forest loss during the same period. Protecting old-growth forests is therefore a priority. Furthermore, we estimate that conserving recovering degraded and secondary forests can have a feasible future carbon sink potential of 53 Tg C year-1 (44-62 Tg C year-1) across the main tropical regions studied.

Increased Amazon carbon emissions mainly from decline in law enforcement.

The Amazon forest carbon sink is declining, mainly as a result of land-use and climate change1-4. Here we investigate how changes in law enforcement of environmental protection policies may have affected the Amazonian carbon balance between 2010 and 2018 compared with 2019 and 2020, based on atmospheric CO2 vertical profiles5,6, deforestation7 and fire data8, as well as infraction notices related to illegal deforestation9. We estimate that Amazonia carbon emissions increased from a mean of 0.24 ± 0.08 PgC year-1 in 2010-2018 to 0.44 ± 0.10 PgC year-1 in 2019 and 0.52 ± 0.10 PgC year-1 in 2020 (± uncertainty). The observed increases in deforestation were 82% and 77% (94% accuracy) and burned area were 14% and 42% in 2019 and 2020 compared with the 2010-2018 mean, respectively. We find that the numbers of notifications of infractions against flora decreased by 30% and 54% and fines paid by 74% and 89% in 2019 and 2020, respectively. Carbon losses during 2019-2020 were comparable with those of the record warm El Niño (2015-2016) without an extreme drought event. Statistical tests show that the observed differences between the 2010-2018 mean and 2019-2020 are unlikely to have arisen by chance. The changes in the carbon budget of Amazonia during 2019-2020 were mainly because of western Amazonia becoming a carbon source. Our results indicate that a decline in law enforcement led to increases in deforestation, biomass burning and forest degradation, which increased carbon emissions and enhanced drying and warming of the Amazon forests.

Direct evidence for phosphorus limitation on Amazon forest productivity.

The productivity of rainforests growing on highly weathered tropical soils is expected to be limited by phosphorus availability1. Yet, controlled fertilization experiments have been unable to demonstrate a dominant role for phosphorus in controlling tropical forest net primary productivity. Recent syntheses have demonstrated that responses to nitrogen addition are as large as to phosphorus2, and adaptations to low phosphorus availability appear to enable net primary productivity to be maintained across major soil phosphorus gradients3. Thus, the extent to which phosphorus availability limits tropical forest productivity is highly uncertain. The majority of the Amazonia, however, is characterized by soils that are more depleted in phosphorus than those in which most tropical fertilization experiments have taken place2. Thus, we established a phosphorus, nitrogen and base cation addition experiment in an old growth Amazon rainforest, with a low soil phosphorus content that is representative of approximately 60% of the Amazon basin. Here we show that net primary productivity increased exclusively with phosphorus addition. After 2 years, strong responses were observed in fine root (+29%) and canopy productivity (+19%), but not stem growth. The direct evidence of phosphorus limitation of net primary productivity suggests that phosphorus availability may restrict Amazon forest responses to CO2 fertilization4, with major implications for future carbon sequestration and forest resilience to climate change.

High aboveground carbon stock of African tropical montane forests.

Tropical forests store 40-50 per cent of terrestrial vegetation carbon1. However, spatial variations in aboveground live tree biomass carbon (AGC) stocks remain poorly understood, in particular in tropical montane forests2. Owing to climatic and soil changes with increasing elevation3, AGC stocks are lower in tropical montane forests compared with lowland forests2. Here we assemble and analyse a dataset of structurally intact old-growth forests (AfriMont) spanning 44 montane sites in 12 African countries. We find that montane sites in the AfriMont plot network have a mean AGC stock of 149.4 megagrams of carbon per hectare (95% confidence interval 137.1-164.2), which is comparable to lowland forests in the African Tropical Rainforest Observation Network4 and about 70 per cent and 32 per cent higher than averages from plot networks in montane2,5,6 and lowland7 forests in the Neotropics, respectively. Notably, our results are two-thirds higher than the Intergovernmental Panel on Climate Change default values for these forests in Africa8. We find that the low stem density and high abundance of large trees of African lowland forests4 is mirrored in the montane forests sampled. This carbon store is endangered: we estimate that 0.8 million hectares of old-growth African montane forest have been lost since 2000. We provide country-specific montane forest AGC stock estimates modelled from our plot network to help to guide forest conservation and reforestation interventions. Our findings highlight the need for conserving these biodiverse9,10 and carbon-rich ecosystems.

Unveiling African rainforest composition and vulnerability to global change.

Africa is forecasted to experience large and rapid climate change1 and population growth2 during the twenty-first century, which threatens the world's second largest rainforest. Protecting and sustainably managing these African forests requires an increased understanding of their compositional heterogeneity, the environmental drivers of forest composition and their vulnerability to ongoing changes. Here, using a very large dataset of 6 million trees in more than 180,000 field plots, we jointly model the distribution in abundance of the most dominant tree taxa in central Africa, and produce continuous maps of the floristic and functional composition of central African forests. Our results show that the uncertainty in taxon-specific distributions averages out at the community level, and reveal highly deterministic assemblages. We uncover contrasting floristic and functional compositions across climates, soil types and anthropogenic gradients, with functional convergence among types of forest that are floristically dissimilar. Combining these spatial predictions with scenarios of climatic and anthropogenic global change suggests a high vulnerability of the northern and southern forest margins, the Atlantic forests and most forests in the Democratic Republic of the Congo, where both climate and anthropogenic threats are expected to increase sharply by 2085. These results constitute key quantitative benchmarks for scientists and policymakers to shape transnational conservation and management strategies that aim to provide a sustainable future for central African forests.

How deregulation, drought and increasing fire impact Amazonian biodiversity.

Biodiversity contributes to the ecological and climatic stability of the Amazon Basin1,2, but is increasingly threatened by deforestation and fire3,4. Here we quantify these impacts over the past two decades using remote-sensing estimates of fire and deforestation and comprehensive range estimates of 11,514 plant species and 3,079 vertebrate species in the Amazon. Deforestation has led to large amounts of habitat loss, and fires further exacerbate this already substantial impact on Amazonian biodiversity. Since 2001, 103,079-189,755 km2 of Amazon rainforest has been impacted by fires, potentially impacting the ranges of 77.3-85.2% of species that are listed as threatened in this region5. The impacts of fire on the ranges of species in Amazonia could be as high as 64%, and greater impacts are typically associated with species that have restricted ranges. We find close associations between forest policy, fire-impacted forest area and their potential impacts on biodiversity. In Brazil, forest policies that were initiated in the mid-2000s corresponded to reduced rates of burning. However, relaxed enforcement of these policies in 2019 has seemingly begun to reverse this trend: approximately 4,253-10,343 km2 of forest has been impacted by fire, leading to some of the most severe potential impacts on biodiversity since 2009. These results highlight the critical role of policy enforcement in the preservation of biodiversity in the Amazon.

Forest mosaics, not savanna corridors, dominated in Southeast Asia during the Last Glacial Maximum.

The dominant paradigm is that large tracts of Southeast Asia's lowland rainforests were replaced with a "savanna corridor" during the cooler, more seasonal climates of the Last Glacial Maximum (LGM) (23,000 to 19,000 y ago). This interpretation has implications for understanding the resilience of Asia's tropical forests to projected climate change, implying a vulnerability to "savannization". A savanna corridor is also an important foundation for archaeological interpretations of how humans moved through and settled insular Southeast Asia and Australia. Yet an up-to-date, multiproxy, and empirical examination of the palaeoecological evidence for this corridor is lacking. We conducted qualitative and statistical analyses of 59 palaeoecological records across Southeast Asia to test the evidence for LGM savannization and clarify the relationships between methods, biogeography, and ecological change in the region from the start of Late Glacial Period (119,000 y ago) to the present. The pollen records typically show montane forest persistence during the LGM, while δ13C biomarker proxies indicate the expansion of C4-rich grasslands. We reconcile this discrepancy by hypothesizing the expansion of montane forest in the uplands and replacement of rainforest with seasonally dry tropical forest in the lowlands. We also find that smooth forest transitions between 34,000 and 2,000 y ago point to the capacity of Southeast Asia's ecosystems both to resist and recover from climate stressors, suggesting resilience to savannization. Finally, the timing of ecological change observed in our combined datasets indicates an 'early' onset of the LGM in Southeast Asia from ~30,000 y ago.

Environmental drivers of megafauna and hominin extinction in Southeast Asia.

Southeast Asia has emerged as an important region for understanding hominin and mammalian migrations and extinctions. High-profile discoveries have shown that Southeast Asia has been home to at least five members of the genus Homo1-3. Considerable turnover in Pleistocene megafauna has previously been linked with these hominins or with climate change4, although the region is often left out of discussions of megafauna extinctions. In the traditional hominin evolutionary core of Africa, attempts to establish the environmental context of hominin evolution and its association with faunal changes have long been informed by stable isotope methodologies5,6. However, such studies have largely been neglected in Southeast Asia. Here we present a large-scale dataset of stable isotope data for Southeast Asian mammals that spans the Quaternary period. Our results demonstrate that the forests of the Early Pleistocene had given way to savannahs by the Middle Pleistocene, which led to the spread of grazers and extinction of browsers-although geochronological limitations mean that not all samples can be resolved to glacial or interglacial periods. Savannahs retreated by the Late Pleistocene and had completely disappeared by the Holocene epoch, when they were replaced by highly stratified closed-canopy rainforest. This resulted in the ascendency of rainforest-adapted species as well as Homo sapiens-which has a unique adaptive plasticity among hominins-at the expense of savannah and woodland specialists, including Homo erectus. At present, megafauna are restricted to rainforests and are severely threatened by anthropogenic deforestation.

Temperate rainforests near the South Pole during peak Cretaceous warmth.

The mid-Cretaceous period was one of the warmest intervals of the past 140 million years1-5, driven by atmospheric carbon dioxide levels of around 1,000 parts per million by volume6. In the near absence of proximal geological records from south of the Antarctic Circle, it is disputed whether polar ice could exist under such environmental conditions. Here we use a sedimentary sequence recovered from the West Antarctic shelf-the southernmost Cretaceous record reported so far-and show that a temperate lowland rainforest environment existed at a palaeolatitude of about 82° S during the Turonian-Santonian age (92 to 83 million years ago). This record contains an intact 3-metre-long network of in situ fossil roots embedded in a mudstone matrix containing diverse pollen and spores. A climate model simulation shows that the reconstructed temperate climate at this high latitude requires a combination of both atmospheric carbon dioxide concentrations of 1,120-1,680 parts per million by volume and a vegetated land surface without major Antarctic glaciation, highlighting the important cooling effect exerted by ice albedo under high levels of atmospheric carbon dioxide.

Estimating the spatial amplification of damage caused by degradation in the Amazon.

The Amazon rainforests have been undergoing unprecedented levels of human-induced disturbances. In addition to local impacts, such changes are likely to cascade following the eastern-western atmospheric flow generated by trade winds. We propose a model of spatial and temporal interactions created by this flow to estimate the spread of effects from local disturbances to downwind locations along atmospheric trajectories. The spatial component captures cascading effects propagated by neighboring regions, while the temporal component captures the persistence of local disturbances. Importantly, all these network effects can be described by a single matrix, acting as a spatial multiplier that amplifies local forest disturbances. This matrix holds practical implications for policymakers as they can use it to easily map where the damage of an initial forest disturbance is amplified and propagated to. We identify regions that are likely to cause the largest impact throughout the basin and those that are the most vulnerable to shocks caused by remote deforestation. On average, the presence of cascading effects mediated by winds in the Amazon doubles the impact of an initial damage. However, there is heterogeneity in this impact. While damage in some regions does not propagate, in others, amplification can reach 250%. Since we only account for spillovers mediated by winds, our multiplier of 2 should be seen as a lower bound.

El Niño-Southern Oscillation forcing on carbon and water cycling in a Bornean tropical rainforest.

Carbon dioxide and water vapor exchanges between tropical forest canopies and the atmosphere through photosynthesis, respiration, and evapotranspiration (ET) influence carbon and water cycling at the regional and global scales. Their inter- and intra-annual variations are sensitive to seasonal rhythms and longer-timescale tropical climatic events. In the present study, we assessed the El Niño-Southern Oscillation (ENSO) influence on ET and on the net ecosystem exchange (NEE), using eddy-covariance flux observations in a Bornean rainforest over a 10-y period (2010-2019) that included several El Niño and La Niña events. From flux model inversions, we inferred ecophysiological properties, notably the canopy stomatal conductance and "big-leaf" maximum carboxylation rate (Vcmax25_BL). Mean ET values were similar between ENSO phases (El Niño, La Niña, and neutral conditions). Conversely, the mean net ecosystem productivity was highest during La Niña events and lowest during El Niño events. Combining Shapley additive explanation calculations for nine controlling factors with a machine-learning algorithm, we determined that the primary factors for ET and NEE in the La Niña and neutral phases were incoming shortwave solar radiation and Vcmax25_BL, respectively, but that canopy stomatal conductance was the most significant factor for both ET and NEE in the El Niño phase. A combined stomatal-photosynthesis model approach further indicated that Vcmax25_BL differences between ENSO phases were the most significant controlling factor for canopy photosynthesis, emphasizing the strong need to account for ENSO-induced ecophysiological factor variations in model projections of the long-term carbon balance in Southeast Asian tropical rainforests.

Recurrent droughts increase risk of cascading tipping events by outpacing adaptive capacities in the Amazon rainforest.

Tipping elements are nonlinear subsystems of the Earth system that have the potential to abruptly shift to another state if environmental change occurs close to a critical threshold with large consequences for human societies and ecosystems. Among these tipping elements may be the Amazon rainforest, which has been undergoing intensive anthropogenic activities and increasingly frequent droughts. Here, we assess how extreme deviations from climatological rainfall regimes may cause local forest collapse that cascades through the coupled forest-climate system. We develop a conceptual dynamic network model to isolate and uncover the role of atmospheric moisture recycling in such tipping cascades. We account for heterogeneity in critical thresholds of the forest caused by adaptation to local climatic conditions. Our results reveal that, despite this adaptation, a future climate characterized by permanent drought conditions could trigger a transition to an open canopy state particularly in the southern Amazon. The loss of atmospheric moisture recycling contributes to one-third of the tipping events. Thus, by exceeding local thresholds in forest adaptive capacity, local climate change impacts may propagate to other regions of the Amazon basin, causing a risk of forest shifts even in regions where critical thresholds have not been crossed locally.

Increasing and widespread vulnerability of intact tropical rainforests to repeated droughts.

Intact tropical rainforests have been exposed to severe droughts in recent decades, which may threaten their integrity, their ability to sequester carbon, and their capacity to provide shelter for biodiversity. However, their response to droughts remains uncertain due to limited high-quality, long-term observations covering extensive areas. Here, we examined how the upper canopy of intact tropical rainforests has responded to drought events globally and during the past 3 decades. By developing a long pantropical time series (1992 to 2018) of monthly radar satellite observations, we show that repeated droughts caused a sustained decline in radar signal in 93%, 84%, and 88% of intact tropical rainforests in the Americas, Africa, and Asia, respectively. Sudden decreases in radar signal were detected around the 1997-1998, 2005, 2010, and 2015 droughts in tropical Americas; 1999-2000, 2004-2005, 2010-2011, and 2015 droughts in tropical Africa; and 1997-1998, 2006, and 2015 droughts in tropical Asia. Rainforests showed similar low resistance (the ability to maintain predrought condition when drought occurs) to severe droughts across continents, but American rainforests consistently showed the lowest resilience (the ability to return to predrought condition after the drought event). Moreover, while the resistance of intact tropical rainforests to drought is decreasing, albeit weakly in tropical Africa and Asia, forest resilience has not increased significantly. Our results therefore suggest the capacity of intact rainforests to withstand future droughts is limited. This has negative implications for climate change mitigation through forest-based climate solutions and the associated pledges made by countries under the Paris Agreement.

Climatic and biotic factors influencing regional declines and recovery of tropical forest biomass from the 2015/16 El Niño.

The 2015/16 El Niño brought severe drought and record-breaking temperatures in the tropics. Here, using satellite-based L-band microwave vegetation optical depth, we mapped changes of above-ground biomass (AGB) during the drought and in subsequent years up to 2019. Over more than 60% of drought-affected intact forests, AGB reduced during the drought, except in the wettest part of the central Amazon, where it declined 1 y later. By the end of 2019, only 40% of AGB reduced intact forests had fully recovered to the predrought level. Using random-forest models, we found that the magnitude of AGB losses during the drought was mainly associated with regionally distinct patterns of soil water deficits and soil clay content. For the AGB recovery, we found strong influences of AGB losses during the drought and of [Formula: see text]. [Formula: see text] is a parameter related to canopy structure and is defined as the ratio of two relative height (RH) metrics of Geoscience Laser Altimeter System (GLAS) waveform data-RH25 (25% energy return height) and RH100 (100% energy return height; i.e., top canopy height). A high [Formula: see text] may reflect forests with a tall understory, thick and closed canopy, and/or without degradation. Such forests with a high [Formula: see text] ([Formula: see text] ≥ 0.3) appear to have a stronger capacity to recover than low-[Formula: see text] ones. Our results highlight the importance of forest structure when predicting the consequences of future drought stress in the tropics.

Phylogeographic analysis of the Bantu language expansion supports a rainforest route.

The Bantu expansion transformed the linguistic, economic, and cultural composition of sub-Saharan Africa. However, the exact dates and routes taken by the ancestors of the speakers of the more than 500 current Bantu languages remain uncertain. Here, we use the recently developed "break-away" geographical diffusion model, specially designed for modeling migrations, with "augmented" geographic information, to reconstruct the Bantu language family expansion. This Bayesian phylogeographic approach with augmented geographical data provides a powerful way of linking linguistic, archaeological, and genetic data to test hypotheses about large language family expansions. We compare four hypotheses: an early major split north of the rainforest; a migration through the Sangha River Interval corridor around 2,500 BP; a coastal migration around 4,000 BP; and a migration through the rainforest before the corridor opening, at 4,000 BP. Our results produce a topology and timeline for the Bantu language family, which supports the hypothesis of an expansion through Central African tropical forests at 4,420 BP (4,040 to 5,000 95% highest posterior density interval), well before the Sangha River Interval was open.

Linking land-use and land-cover transitions to their ecological impact in the Amazon.

Human activities pose a major threat to tropical forest biodiversity and ecosystem services. Although the impacts of deforestation are well studied, multiple land-use and land-cover transitions (LULCTs) occur in tropical landscapes, and we do not know how LULCTs differ in their rates or impacts on key ecosystem components. Here, we quantified the impacts of 18 LULCTs on three ecosystem components (biodiversity, carbon, and soil), based on 18 variables collected from 310 sites in the Brazilian Amazon. Across all LULCTs, biodiversity was the most affected ecosystem component, followed by carbon stocks, but the magnitude of change differed widely among LULCTs and individual variables. Forest clearance for pasture was the most prevalent and high-impact transition, but we also identified other LULCTs with high impact but lower prevalence (e.g., forest to agriculture). Our study demonstrates the importance of considering multiple ecosystem components and LULCTs to understand the consequences of human activities in tropical landscapes.