Limits to the accurate and generalizable use of soundscapes to monitor biodiversity.

Although eco-acoustic monitoring has the potential to deliver biodiversity insight on vast scales, existing analytical approaches behave unpredictably across studies. We collated 8,023 audio recordings with paired manual avifaunal point counts to investigate whether soundscapes could be used to monitor biodiversity across diverse ecosystems. We found that neither univariate indices nor machine learning models were predictive of species richness across datasets but soundscape change was consistently indicative of community change. Our findings indicate that there are no common features of biodiverse soundscapes and that soundscape monitoring should be used cautiously and in conjunction with more reliable in-person ecological surveys.

Co-limitation towards lower latitudes shapes global forest diversity gradients.

The latitudinal diversity gradient (LDG) is one of the most recognized global patterns of species richness exhibited across a wide range of taxa. Numerous hypotheses have been proposed in the past two centuries to explain LDG, but rigorous tests of the drivers of LDGs have been limited by a lack of high-quality global species richness data. Here we produce a high-resolution (0.025° × 0.025°) map of local tree species richness using a global forest inventory database with individual tree information and local biophysical characteristics from ~1.3 million sample plots. We then quantify drivers of local tree species richness patterns across latitudes. Generally, annual mean temperature was a dominant predictor of tree species richness, which is most consistent with the metabolic theory of biodiversity (MTB). However, MTB underestimated LDG in the tropics, where high species richness was also moderated by topographic, soil and anthropogenic factors operating at local scales. Given that local landscape variables operate synergistically with bioclimatic factors in shaping the global LDG pattern, we suggest that MTB be extended to account for co-limitation by subordinate drivers.

A global evaluation of the effectiveness of voluntary REDD+ projects at reducing deforestation and degradation in the moist tropics.

Reducing emissions from deforestation and forest degradation (REDD+) projects aim to contribute to climate change mitigation by protecting and enhancing carbon stocks in tropical forests, but there have been no systematic global evaluations of their impact. We used a new data set for tropical humid forests and a standardized evaluation approach (based on pixel matching) to quantify the performance of a representative sample of 40 voluntary REDD+ projects in 9 countries certified under the Verified Carbon Standard (VCS). In the first 5 years of implementation, deforestation within project areas was reduced by 47% (95% confidence interval [CI]: 24-68) compared with matched counterfactual pixels, and degradation rates were 58% lower (95% CI: 49-63). Reductions were small in absolute terms but greater in sites located in high-deforestation settings and did not appear to be substantially undermined by leakage activities in forested areas within 10 km of project boundaries. At the 26th Conference of the Parties of the United Nations Framework Convention on Climate Change, the international community renewed its commitment to tackling tropical deforestation as a nature-based solution to climate change. Our results indicate that incentivizing forest conservation through voluntary site-based projects can slow tropical deforestation and highlight the particular importance of prioritizing financing for areas at greater risk of deforestation.

Evaluación Global de la Efectividad de proyectos REDD+ en la Reducción de la Deforestación y Degradación en el Trópico Húmedo Resumen Los proyectos para la reducción de emisiones derivados de la deforestación y degradación de bosques (REDD+) tienen como meta contribuir a la mitigación del cambio climático al protejer y  fomentar la disponibilidad de carbono en bosques tropicales, pero a la fecha no se han realizado evaluaciones globales sistemáticas sobre su impacto. Utilizamos bases de datos recientes sobre bosques tropicales húmedos y un método estandarizado de evaluación (basado en 'emparejamiento' [matching] de pixeles) para cuantificar el desempeño de una muestra representativa de 40 proyectos voluntarios REDD+ , localizados en 9 países y certificados bajo el estándar Verified Carbon Standard (VCS). En los primeros 5 años de implementación, la deforestación en las áreas de los proyectos disminuyó en 47% (IC 95% 24-68) en comparación con los pixeles contrafactuales correspondientes, y las tasas de degradación fueron 58% menos (IC 95% 49-63). Las reducciones fueron pequeñas en términos absolutos pero mayores en sitios con tasas de deforestación elevadas, y no parecieron ser afectadas sustancialmente por efectos de fuga (leakage) en áreas boscosas en un radio de 10 km de los límites del proyecto. En la COP26, la comunidad internacional renovó su compromiso de afrontar la deforestación tropical como una solución al cambio climático basada en la naturaleza. Nuestros resultados indican que incentivar la conservación de bosques mediante proyectos locales voluntarios puede disminuir la deforestación tropical, y resaltan la importancia de priorizar financiamiento en áreas con un mayor riesgo de deforestación.

Tallo: A global tree allometry and crown architecture database.

Data capturing multiple axes of tree size and shape, such as a tree's stem diameter, height and crown size, underpin a wide range of ecological research-from developing and testing theory on forest structure and dynamics, to estimating forest carbon stocks and their uncertainties, and integrating remote sensing imagery into forest monitoring programmes. However, these data can be surprisingly hard to come by, particularly for certain regions of the world and for specific taxonomic groups, posing a real barrier to progress in these fields. To overcome this challenge, we developed the Tallo database, a collection of 498,838 georeferenced and taxonomically standardized records of individual trees for which stem diameter, height and/or crown radius have been measured. These data were collected at 61,856 globally distributed sites, spanning all major forested and non-forested biomes. The majority of trees in the database are identified to species (88%), and collectively Tallo includes data for 5163 species distributed across 1453 genera and 187 plant families. The database is publicly archived under a CC-BY 4.0 licence and can be access from: https://doi.org/10.5281/zenodo.6637599. To demonstrate its value, here we present three case studies that highlight how the Tallo database can be used to address a range of theoretical and applied questions in ecology-from testing the predictions of metabolic scaling theory, to exploring the limits of tree allometric plasticity along environmental gradients and modelling global variation in maximum attainable tree height. In doing so, we provide a key resource for field ecologists, remote sensing researchers and the modelling community working together to better understand the role that trees play in regulating the terrestrial carbon cycle.

Functional susceptibility of tropical forests to climate change.

Tropical forests are some of the most biodiverse ecosystems in the world, yet their functioning is threatened by anthropogenic disturbances and climate change. Global actions to conserve tropical forests could be enhanced by having local knowledge on the forests' functional diversity and functional redundancy as proxies for their capacity to respond to global environmental change. Here we create estimates of plant functional diversity and redundancy across the tropics by combining a dataset of 16 morphological, chemical and photosynthetic plant traits sampled from 2,461 individual trees from 74 sites distributed across four continents together with local climate data for the past half century. Our findings suggest a strong link between climate and functional diversity and redundancy with the three trait groups responding similarly across the tropics and climate gradient. We show that drier tropical forests are overall less functionally diverse than wetter forests and that functional redundancy declines with increasing soil water and vapour pressure deficits. Areas with high functional diversity and high functional redundancy tend to better maintain ecosystem functioning, such as aboveground biomass, after extreme weather events. Our predictions suggest that the lower functional diversity and lower functional redundancy of drier tropical forests, in comparison with wetter forests, may leave them more at risk of shifting towards alternative states in face of further declines in water availability across tropical regions.

The number of tree species on Earth.

One of the most fundamental questions in ecology is how many species inhabit the Earth. However, due to massive logistical and financial challenges and taxonomic difficulties connected to the species concept definition, the global numbers of species, including those of important and well-studied life forms such as trees, still remain largely unknown. Here, based on global ground-sourced data, we estimate the total tree species richness at global, continental, and biome levels. Our results indicate that there are ∼73,000 tree species globally, among which ∼9,000 tree species are yet to be discovered. Roughly 40% of undiscovered tree species are in South America. Moreover, almost one-third of all tree species to be discovered may be rare, with very low populations and limited spatial distribution (likely in remote tropical lowlands and mountains). These findings highlight the vulnerability of global forest biodiversity to anthropogenic changes in land use and climate, which disproportionately threaten rare species and thus, global tree richness.

Arbuscular mycorrhizal trees influence the latitudinal beta-diversity gradient of tree communities in forests worldwide.

Arbuscular mycorrhizal (AM) and ectomycorrhizal (EcM) associations are critical for host-tree performance. However, how mycorrhizal associations correlate with the latitudinal tree beta-diversity remains untested. Using a global dataset of 45 forest plots representing 2,804,270 trees across 3840 species, we test how AM and EcM trees contribute to total beta-diversity and its components (turnover and nestedness) of all trees. We find AM rather than EcM trees predominantly contribute to decreasing total beta-diversity and turnover and increasing nestedness with increasing latitude, probably because wide distributions of EcM trees do not generate strong compositional differences among localities. Environmental variables, especially temperature and precipitation, are strongly correlated with beta-diversity patterns for both AM trees and all trees rather than EcM trees. Results support our hypotheses that latitudinal beta-diversity patterns and environmental effects on these patterns are highly dependent on mycorrhizal types. Our findings highlight the importance of AM-dominated forests for conserving global forest biodiversity.

Recovery of logged forest fragments in a human-modified tropical landscape during the 2015-16 El Niño.

The past 40 years in Southeast Asia have seen about 50% of lowland rainforests converted to oil palm and other plantations, and much of the remaining forest heavily logged. Little is known about how fragmentation influences recovery and whether climate change will hamper restoration. Here, we use repeat airborne LiDAR surveys spanning the hot and dry 2015-16 El Niño Southern Oscillation event to measure canopy height growth across 3,300 ha of regenerating tropical forests spanning a logging intensity gradient in Malaysian Borneo. We show that the drought led to increased leaf shedding and branch fall. Short forest, regenerating after heavy logging, continued to grow despite higher evaporative demand, except when it was located close to oil palm plantations. Edge effects from the plantations extended over 300 metres into the forests. Forest growth on hilltops and slopes was particularly impacted by the combination of fragmentation and drought, but even riparian forests located within 40 m of oil palm plantations lost canopy height during the drought. Our results suggest that small patches of logged forest within plantation landscapes will be slow to recover, particularly as ENSO events are becoming more frequent.

Carbon flux and forest dynamics: Increased deadwood decomposition in tropical rainforest tree-fall canopy gaps.

Tree mortality rates are increasing within tropical rainforests as a result of global environmental change. When trees die, gaps are created in forest canopies and carbon is transferred from the living to deadwood pools. However, little is known about the effect of tree-fall canopy gaps on the activity of decomposer communities and the rate of deadwood decay in forests. This means that the accuracy of regional and global carbon budgets is uncertain, especially given ongoing changes to the structure of rainforest ecosystems. Therefore, to determine the effect of canopy openings on wood decay rates and regional carbon flux, we carried out the first assessment of deadwood mass loss within canopy gaps in old-growth rainforest. We used replicated canopy gaps paired with closed canopy sites in combination with macroinvertebrate accessible and inaccessible woodblocks to experimentally partition the relative contribution of microbes vs. termites to decomposition within contrasting understorey conditions. We show that over a 12 month period, wood mass loss increased by 63% in canopy gaps compared with closed canopy sites and that this increase was driven by termites. Using LiDAR data to quantify the proportion of canopy openings in the study region, we modelled the effect of observed changes in decomposition within gaps on regional carbon flux. Overall, we estimate that this accelerated decomposition increases regional wood decay rate by up to 18.2%, corresponding to a flux increase of 0.27 Mg C ha-1  year-1 that is not currently accounted for in regional carbon budgets. These results provide the first insights into how small-scale disturbances in rainforests can generate hotspots for decomposer activity and carbon fluxes. In doing so, we show that including canopy gap dynamics and their impacts on wood decomposition in forest ecosystems can help improve the predictive accuracy of the carbon cycle in land surface models.

Leech blood-meal invertebrate-derived DNA reveals differences in Bornean mammal diversity across habitats.

The application of metabarcoding to environmental and invertebrate-derived DNA (eDNA and iDNA) is a new and increasingly applied method for monitoring biodiversity across a diverse range of habitats. This approach is particularly promising for sampling in the biodiverse humid tropics, where rapid land-use change for agriculture means there is a growing need to understand the conservation value of the remaining mosaic and degraded landscapes. Here we use iDNA from blood-feeding leeches (Haemadipsa picta) to assess differences in mammalian diversity across a gradient of forest degradation in Sabah, Malaysian Borneo. We screened 557 individual leeches for mammal DNA by targeting fragments of the 16S rRNA gene and detected 14 mammalian genera. We recorded lower mammal diversity in the most heavily degraded forest compared to higher quality twice logged forest. Although the accumulation curves of diversity estimates were comparable across these habitat types, diversity was higher in twice logged forest, with more taxa of conservation concern. In addition, our analysis revealed differences between the community recorded in the heavily logged forest and that of the twice logged forest. By revealing differences in mammal diversity across a human-modified tropical landscape, our study demonstrates the value of iDNA as a noninvasive biomonitoring approach in conservation assessments.

Maximizing the value of forest restoration for tropical mammals by detecting three-dimensional habitat associations.

Tropical forest ecosystems are facing unprecedented levels of degradation, severely compromising habitat suitability for wildlife. Despite the fundamental role biodiversity plays in forest regeneration, identifying and prioritizing degraded forests for restoration or conservation, based on their wildlife value, remains a significant challenge. Efforts to characterize habitat selection are also weakened by simple classifications of human-modified tropical forests as intact vs. degraded, which ignore the influence that three-dimensional (3D) forest structure may have on species distributions. Here, we develop a framework to identify conservation and restoration opportunities across logged forests in Borneo. We couple high-resolution airborne light detection and ranging (LiDAR) and camera trap data to characterize the response of a tropical mammal community to changes in 3D forest structure across a degradation gradient. Mammals were most responsive to covariates that accounted explicitly for the vertical and horizontal characteristics of the forest and actively selected structurally complex environments comprising tall canopies, increased plant area index throughout the vertical column, and the availability of a greater diversity of niches. We show that mammals are sensitive to structural simplification through disturbance, emphasizing the importance of maintaining and enhancing structurally intact forests. By calculating occurrence thresholds of species in response to forest structural change, we identify areas of degraded forest that would provide maximum benefit for multiple high-conservation value species if restored. The study demonstrates the advantages of using LiDAR to map forest structure, rather than relying on overly simplistic classifications of human-modified tropical forests, for prioritizing regions for restoration.

Resilience of Spanish forests to recent droughts and climate change.

A widespread increase in forest cover is underway in northern Mediterranean forests because of land abandonment and decreased wood demand, but the resilience of these successional forests to climate change remains unresolved. Here we use 18-year time series of canopy greenness derived from satellite imagery (NDVI) to evaluate the impacts of climate change on Spain's forests. Specifically, we analyzed how NDVI was influenced by the climatic water balance (i.e. Standardized Precipitation-Evapotranspiration Index, SPEI), using monthly time-series extracted from 3,100 pixels of forest, categorized into ten forest types. The forests increased in leaf area index by 0.01 per year on average (from 1.7 in 2000 to 1.9 in 2017) but there was enormous variation among years related to climatic water balance. Forest types varied in response to drought events: those dominated by drought-avoiding species showed strong covariance between greenness and SPEI, while those dominated by drought-tolerant species showed weak covariance. Native forests usually recovered more than 80% of greenness within the 18 months and the remainder within 5 years, but plantations of Eucalyptus were less resilient. Management to increase the resilience of forests-a key goal of forestry in the Mediterranean region-appears to have had a positive effect: canopy greenness within protected forests was more resilient to drought than within non-protected forests. In conclusion, many of Spain's successional forests have been resilient to drought over the past 18 years, from the perspective of space. Future studies will need to combine remote sensing with field-based analyses of physiological tolerances and mortality processes to understand how Mediterranean forests will respond to the rapid climate change predicted for this region in the coming decades.

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.

Partial river flow recovery with forest age is rare in the decades following establishment.

Forest regeneration and expansion are occurring in many countries, with 80 million ha established from 2000 to 2012 under the Bonn accord and 17.5 million ha established from 1990 to 2005 according to the Food and Agriculture Organisation. Multiple reviews have linked increasing forest cover with reduced river flow and potentially detrimental effects downstream. Previous reviews have investigated trends in river flow response over time, but the influence of forest age remains uncertain. Partial river flow recovery (towards non-forested conditions) has been reported in decades following forest establishment, but the role of climate in driving these trends has not been explored. Here, we evaluate river flow trends in 43 studies following forest establishment, which provide sufficient information to distinguish the effects of ageing forests from variable climate. Our meta-analysis supports previous findings showing that forestation reduces annual river flow (by 23% after 5 years and 38% after 25 years) with greater reductions in catchments with higher mean annual precipitation, larger increases in forest cover, and which were idle, rather than agricultural land, prior to forestation. The impact of forests on river flow is sensitive to annual precipitation and potential evapotranspiration, but responses are highly variable. Forests affect river flow less when annual precipitation is low, and sensitivity to precipitation decreases as catchment aridity increases. The majority of catchments demonstrated persistent river flow declines after forest establishment. However, nine catchments showed partial flow recovery after an initial decrease, with peak flow reductions at an average age of 15 and across a range of tree species. The mean rate of recovery was 34 mm/year over 5 years. Partial flow recovery with forest age cannot be commonly expected, however, and forestation programmes should take into account that changes to annual river flow are likely to persist for up to five decades.

How do trees respond to species mixing in experimental compared to observational studies?

For decades, ecologists have investigated the effects of tree species diversity on tree productivity at different scales and with different approaches ranging from observational to experimental study designs. Using data from five European national forest inventories (16,773 plots), six tree species diversity experiments (584 plots), and six networks of comparative plots (169 plots), we tested whether tree species growth responses to species mixing are consistent and therefore transferrable between those different research approaches. Our results confirm the general positive effect of tree species mixing on species growth (16% on average) but we found no consistency in species-specific responses to mixing between any of the three approaches, even after restricting comparisons to only those plots that shared similar mixtures compositions and forest types. These findings highlight the necessity to consider results from different research approaches when selecting species mixtures that should maximize positive forest biodiversity and functioning relationships.

Reconciling the contribution of environmental and stochastic structuring of tropical forest diversity through the lens of imaging spectroscopy.

Both niche and stochastic dispersal processes structure the extraordinary diversity of tropical plants, but determining their relative contributions has proven challenging. We address this question using airborne imaging spectroscopy to estimate canopy ß-diversity for an extensive region of a Bornean rainforest and challenge these data with models incorporating niches and dispersal. We show that remotely sensed and field-derived estimates of pairwise dissimilarity in community composition are closely matched, proving the applicability of imaging spectroscopy to provide ß-diversity data for entire landscapes of over 1000 ha containing contrasting forest types. Our model reproduces the empirical data well and shows that the ecological processes maintaining tropical forest diversity are scale dependent. Patterns of ß-diversity are shaped by stochastic dispersal processes acting locally whilst environmental processes act over a wider range of scales.

Author Correction: Limited capacity of tree growth to mitigate the global greenhouse effect under predicted warming.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Forest fragmentation in China and its effect on biodiversity.

Land-use change is fragmenting natural ecosystems, with major consequences for biodiversity. This paper reviews fragmentation trends - historical and current - in China, the fourth largest country on Earth, and explores its consequences. Remote sensing makes it possible to track land-use change at a global scale and monitor fragmentation of dwindling natural landscapes. Yet few studies have linked fragmentation mapped remotely with impacts on biodiversity within human-modified landscapes. Recent reforestation programs have caused substantial increases in forest cover but have not stopped fragmentation, because the new forests are mostly monocultures that further fragment China's remnant old-growth lowland forests that harbour the highest levels of biodiversity. Fragmentation - and associated biodiversity declines - is unevenly distributed in China's forests, being most problematic where agricultural expansion is occurring in the southwest and northeast, serious in the densely populated eastern regions where urbanisation and transport infrastructure are modifying landscapes, but less of a problem in other regions. Analyses of temporal trends show that the drivers of forest fragmentation are shifting from mainly agricultural expansion to urbanisation and infrastructure development. Most of China's old-growth forests persist in small, isolated fragments from which many native species have disappeared, on land unsuitable for human utilisation. Fragmentation throughout China is likely to have major consequences on biodiversity conservation, but few studies have considered these large-scale processes at the national level. Our review fills this research gap and puts forward a systematic perspective relevant to China and beyond.

Limited capacity of tree growth to mitigate the global greenhouse effect under predicted warming.

It is generally accepted that animal heartbeat and lifespan are often inversely correlated, however, the relationship between productivity and longevity has not yet been described for trees growing under industrial and pre-industrial climates. Using 1768 annually resolved and absolutely dated ring width measurement series from living and dead conifers that grew in undisturbed, high-elevation sites in the Spanish Pyrenees and the Russian Altai over the past 2000 years, we test the hypothesis of grow fast-die young. We find maximum tree ages are significantly correlated with slow juvenile growth rates. We conclude, the interdependence between higher stem productivity, faster tree turnover, and shorter carbon residence time, reduces the capacity of forest ecosystems to store carbon under a climate warming-induced stimulation of tree growth at policy-relevant timescales.

A critique of general allometry-inspired models for estimating forest carbon density from airborne LiDAR.

There is currently much interest in developing general approaches for mapping forest aboveground carbon density using structural information contained in airborne LiDAR data. The most widely utilized model in tropical forests assumes that aboveground carbon density is a compound power function of top of canopy height (a metric easily derived from LiDAR), basal area and wood density. Here we derive the model in terms of the geometry of individual tree crowns within forest stands, showing how scaling exponents in the aboveground carbon density model arise from the height-diameter (H-D) and projected crown area-diameter (C-D) allometries of individual trees. We show that a power function relationship emerges when the C-D scaling exponent is close to 2, or when tree diameters follow a Weibull distribution (or other specific distributions) and are invariant across the landscape. In addition, basal area must be closely correlated with canopy height for the approach to work. The efficacy of the model was explored for a managed uneven-aged temperate forest in Ontario, Canada within which stands dominated by sugar maple (Acer saccharum Marsh.) and mixed stands were identified. A much poorer goodness-of-fit was obtained than previously reported for tropical forests (R2 = 0.29 vs. about 0.83). Explanations for the poor predictive power on the model include: (1) basal area was only weakly correlated with top canopy height; (2) tree size distributions varied considerably across the landscape; (3) the allometry exponents are affected by variation in species composition arising from timber management and soil conditions; and (4) the C-D allometric power function was far from 2 (1.28). We conclude that landscape heterogeneity in forest structure and tree allometry reduces the accuracy of general power-function models for predicting aboveground carbon density in managed forests. More studies in different forest types are needed to understand the situations in which power functions of LiDAR height are appropriate for modelling forest carbon stocks.