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Correction for "Phylogenetic classification of the world's tropical forests, " by J. W. Ferry Slik, Janet Franklin, Víctor Arroyo- Rodríguez, Richard Field, Salomon Aguilar, Nikolay Aguirre, Jorge Ahumada, Shin-Ichiro Aiba, Luciana F. Alves, Anitha K, Andres Avella, Francisco Mora, Gerardo A. Aymard C., Selene B?ez, Patricia Balvanera, Meredith L. Bastian, Jean-François Bastin, Peter J. Bellingham, Eduardo van den Berg, Polyanna da Conceição Bispo, Pascal Boeckx, Katrin Boehning-Gaese, Frans Bongers, Brad Boyle, Fabian Brambach, Francis Q. Brearley, Sandra Brown, Shauna-Lee Chai, Robin L. Chazdon, Shengbin Chen, Phourin Chhang, George Chuyong, Corneille Ewango, Indiana M. Coronado, Jurgi Crist?bal-Azkarate, Heike Culmsee, Kipiro Damas, H. S. Dattaraja, Priya Davidar, Saara J. DeWalt, Hazimah Din, Donald R. Drake, Alvaro Duque, Giselda Durigan, Karl Eichhorn, Eduardo Schmidt Eler, Tsutomu Enoki, Andreas Ensslin, Adand? Belarmain Fandohan, Nina Farwig, Kenneth J. Feeley, Markus Fischer, Olle Forshed, Queila Souza Garcia, Satish Chandra Garkoti, Thomas W. Gillespie, Jean- Francois Gillet, Christelle Gonmadje, Iñigo Granzow-de la Cerda, Daniel M. Griffith, James Grogan, Khalid Rehman Hakeem, David J. Harris, Rhett D. Harrison, Andy Hector, Andreas Hemp, J?rgen Homeier, M. Shah Hussain, Guillermo Ibarra-Manríquez, I. Faridah Hanum, Nobuo Imai, Patrick A. Jansen, Carlos Alfredo Joly, Shijo Joseph, Kuswata Kartawinata, Elizabeth Kearsley, Daniel L. Kelly, Michael Kessler, Timothy J. Killeen, Robert M. Kooyman, Yves Laumonier, Susan G. Laurance, William F. Laurance, Michael J. Lawes, Susan G. Letcher, Jeremy Lindsell, Jon Lovett, Jose Lozada, Xinghui Lu, Anne Mette Lykke, Khairil Bin Mahmud, Ni Putu Diana Mahayani, Asyraf Mansor, Andrew R. Marshall, Emanuel H. Martin, Darley Calderado Leal Matos, Jorge A.Meave, Felipe P. L. Melo, Zhofre Huberto Aguirre Mendoza, Faizah Metali, Vincent P. Medjibe, Jean Paul Metzger, Thiago Metzker, D. Mohandass, Miguel A. Munguía-Rosas, Rodrigo Muñoz, Eddy Nurtjahy, Eddie Lenza de Oliveira, Onrizal, Pia Parolin, Marc Parren, N. Parthasarathy, Ekananda Paudel, Rolando Perez, Eduardo A. P?rez-García, Ulf Pommer, Lourens Poorter, Lan Qi, Maria Teresa F. Piedade, Jos? Roberto Rodrigues Pinto, Axel Dalberg Poulsen, John R. Poulsen, Jennifer S. Powers, Rama Chandra Prasad, Jean-Philippe Puyravaud, Orlando Rangel, Jan Reitsma, Diogo S. B. Rocha, Samir Rolim, Francesco Rovero, Andes Rozak, Kalle Ruokolainen, Ervan Rutishauser, Gemma Rutten, Mohd. NizamMohd. Said, Felipe Z. Saiter, Philippe Saner, Braulio Santos, João Roberto dos Santos, Swapan Kumar Sarker, Christine B. Schmitt, Jochen Schoengart, Mark Schulze, Douglas Sheil, Plinio Sist, Alexandre F. Souza, Wilson Roberto Spironello, Tereza Sposito, Robert Steinmetz, Tariq Stevart, Marcio Seiji Suganuma, Rahayu Sukri, Aisha Sultana, Raman Sukumar, Terry Sunderland, Supriyadi, H. S. Suresh, Eizi Suzuki, Marcelo Tabarelli, Jianwei Tang, Ed V. J. Tanner, Natalia Targhetta, Ida Theilade, Duncan Thomas, Jonathan Timberlake, M?rcio de Morisson Valeriano, Johan van Valkenburg, Tran Van Do, Hoang Van Sam, John H. Vandermeer, Hans Verbeeck, Ole Reidar Vetaas, Victor Adekunle, Simone A. Vieira, Campbell O.Webb, Edward L. Webb, Timothy Whitfeld, Serge Wich, John Williams, Susan Wiser, FlorianWittmann, Xiaobo Yang, C. Yves Adou Yao, Sandra L. Yap, Rakan A. Zahawi, Rahmad Zakaria, and Runguo Zang, which was first published February 5, 2018; 10.1073/pnas.1714977115 (Proc Natl Acad Sci USA 115:1837-1842). The authors note that, due to a printer's error, an author name published incorrectly. The author name Lan Qi should instead appear as Lan Qie. The online version has been corrected.

Extensive forest restoration is a key strategy to meet nature-based sustainable development goals and provide multiple social and environmental benefits 1 . Yet achieving forest restoration at scale requires cost-effective methods 2 . Tree planting in degraded landscapes is a popular but costly forest restoration method that often results in less biodiverse forests when compared to natural regeneration techniques under similar conditions 3 . Here we assess the current spatial distribution of pantropical natural forest (from 2000 to 2016) and use this to present a model of the potential for natural regeneration across tropical forested countries and biomes at a spatial resolution of 30 m. We estimate that an area of 215 million hectares-an area greater than the entire country of Mexico-has potential for natural forest regeneration, representing an above-ground carbon sequestration potential of 23.4 Gt C (range, 21.1-25.7 Gt) over 30 years. Five countries (Brazil, Indonesia, China, Mexico and Colombia) account for 52% of this estimated potential, showcasing the need for targeting restoration initiatives that leverage natural regeneration potential. Our results facilitate broader equitable decision-making processes that capitalize on the widespread opportunity for natural regeneration to help achieve national and global environmental agendas., (© 2024. The Author(s).)

The interspecific relationship between functional traits and tree seedling performance can be inconsistent, potentially due to site-to-site or microsite variation in environmental conditions. Studies of seedling traits and performance often focus on above-ground traits, despite the importance of below-ground resource acquisition and biomass allocation to above versus below-ground functions. Here we investigate how varying environmental conditions across sites induce intraspecific variation in organ-level (above-ground, below-ground) and biomass allocation traits, affecting interspecific relationships between these traits and seedling performance. We analyzed trait expression for 12 organ-level and three allocation traits and their relationships with height growth (1716 seedlings) and mortality (15,862 seedlings) for 26 tree species across three sites along a forest successional gradient in Costa Rica. We found significant intraspecific differences across sites in all allocation traits, but only in three of seven above-ground and three of five below-ground organ-level traits. Allocation traits were better predictors of seedling performance than organ-level traits. Relationships between allocation traits and both growth and mortality varied among all sites, but for organ-level traits, only relationships with growth varied among sites. These results underscore that biomass allocation plays a key role in the earliest life stages of trees and that site-specific conditions can influence how functional traits mediate seedling establishment during succession.

Secondary tropical forests play an increasingly important role in carbon budgets and biodiversity conservation. Understanding successional trajectories is therefore imperative for guiding forest restoration and climate change mitigation efforts. Forest succession is driven by the demographic strategies-combinations of growth, mortality and recruitment rates-of the tree species in the community. However, our understanding of demographic diversity in tropical tree species stems almost exclusively from old-growth forests. Here, we assembled demographic information from repeated forest inventories along chronosequences in two wet (Costa Rica, Panama) and two dry (Mexico) Neotropical forests to assess whether the ranges of demographic strategies present in a community shift across succession. We calculated demographic rates for >500 tree species while controlling for canopy status to compare demographic diversity (i.e., the ranges of demographic strategies) in early successional (0-30 years), late successional (30-120 years) and old-growth forests using two-dimensional hypervolumes of pairs of demographic rates. Ranges of demographic strategies largely overlapped across successional stages, and early successional stages already covered the full spectrum of demographic strategies found in old-growth forests. An exception was a group of species characterized by exceptionally high mortality rates that was confined to early successional stages in the two wet forests. The range of demographic strategies did not expand with succession. Our results suggest that studies of long-term forest monitoring plots in old-growth forests, from which most of our current understanding of demographic strategies of tropical tree species is derived, are surprisingly representative of demographic diversity in general, but do not replace the need for further studies in secondary forests., (© 2024 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Forests are a substantial terrestrial carbon sink, but anthropogenic changes in land use and climate have considerably reduced the scale of this system 1 . Remote-sensing estimates to quantify carbon losses from global forests 2-5 are characterized by considerable uncertainty and we lack a comprehensive ground-sourced evaluation to benchmark these estimates. Here we combine several ground-sourced 6 and satellite-derived approaches 2,7,8 to evaluate the scale of the global forest carbon potential outside agricultural and urban lands. Despite regional variation, the predictions demonstrated remarkable consistency at a global scale, with only a 12% difference between the ground-sourced and satellite-derived estimates. At present, global forest carbon storage is markedly under the natural potential, with a total deficit of 226 Gt (model range = 151-363 Gt) in areas with low human footprint. Most (61%, 139 Gt C) of this potential is in areas with existing forests, in which ecosystem protection can allow forests to recover to maturity. The remaining 39% (87 Gt C) of potential lies in regions in which forests have been removed or fragmented. Although forests cannot be a substitute for emissions reductions, our results support the idea 2,3,9 that the conservation, restoration and sustainable management of diverse forests offer valuable contributions to meeting global climate and biodiversity targets., (© 2023. The Author(s).)

Abandonment of agricultural lands promotes the global expansion of secondary forests, which are critical for preserving biodiversity and ecosystem functions and services. Such roles largely depend, however, on two essential successional attributes, trajectory and recovery rate, which are expected to depend on landscape-scale forest cover in nonlinear ways. Using a multi-scale approach and a large vegetation dataset (843 plots, 3511 tree species) from 22 secondary forest chronosequences distributed across the Neotropics, we show that successional trajectories of woody plant species richness, stem density and basal area are less predictable in landscapes (4 km radius) with intermediate (40-60%) forest cover than in landscapes with high (greater than 60%) forest cover. This supports theory suggesting that high spatial and environmental heterogeneity in intermediately deforested landscapes can increase the variation of key ecological factors for forest recovery (e.g. seed dispersal and seedling recruitment), increasing the uncertainty of successional trajectories. Regarding the recovery rate, only species richness is positively related to forest cover in relatively small (1 km radius) landscapes. These findings highlight the importance of using a spatially explicit landscape approach in restoration initiatives and suggest that these initiatives can be more effective in more forested landscapes, especially if implemented across spatial extents of 1-4 km radius.

Current policy is driving renewed impetus to restore forests to return ecological function, protect species, sequester carbon and secure livelihoods. Here we assess the contribution of tree planting to ecosystem restoration in tropical and sub-tropical Asia; we synthesize evidence on mortality and growth of planted trees at 176 sites and assess structural and biodiversity recovery of co-located actively restored and naturally regenerating forest plots. Mean mortality of planted trees was 18% 1 year after planting, increasing to 44% after 5 years. Mortality varied strongly by site and was typically ca 20% higher in open areas than degraded forest, with height at planting positively affecting survival. Size-standardized growth rates were negatively related to species-level wood density in degraded forest and plantations enrichment settings. Based on community-level data from 11 landscapes, active restoration resulted in faster accumulation of tree basal area and structural properties were closer to old-growth reference sites, relative to natural regeneration, but tree species richness did not differ. High variability in outcomes across sites indicates that planting for restoration is potentially rewarding but risky and context-dependent. Restoration projects must prepare for and manage commonly occurring challenges and align with efforts to protect and reconnect remaining forest areas. The abstract of this article is available in Bahasa Indonesia in the electronic supplementary material. This article is part of the theme issue 'Understanding forest landscape restoration: reinforcing scientific foundations for the UN Decade on Ecosystem Restoration'.

There has never been a more pressing and opportune time for science and practice to collaborate towards restoration of the world's forests. Multiple uncertainties remain for achieving successful, long-term forest landscape restoration (FLR). In this article, we use expert knowledge and literature review to identify knowledge gaps that need closing to advance restoration practice, as an introduction to a landmark theme issue on FLR and the UN Decade on Ecosystem Restoration. Aligned with an Adaptive Management Cycle for FLR, we identify 15 essential science advances required to facilitate FLR success for nature and people. They highlight that the greatest science challenges lie in the conceptualization, planning and assessment stages of restoration, which require an evidence base for why, where and how to restore, at realistic scales. FLR and underlying sciences are complex, requiring spatially explicit approaches across disciplines and sectors, considering multiple objectives, drivers and trade-offs critical for decision-making and financing. The developing tropics are a priority region, where scientists must work with stakeholders across the Adaptive Management Cycle. Clearly communicated scientific evidence for action at the outset of restoration planning will enable donors, decision makers and implementers to develop informed objectives, realistic targets and processes for accountability. This article paves the way for 19 further articles in this theme issue, with author contributions from across the world. This article is part of the theme issue 'Understanding forest landscape restoration: reinforcing scientific foundations for the UN Decade on Ecosystem Restoration'.

Given the importance of species diversity as a tool for assessing recovery during forest regeneration and active restoration, robust approaches for assessing changes in tree species diversity over time are urgently needed. We assessed changes in tree species diversity during natural regeneration over 12-20 years in eight 1-ha monitoring plots in NE Costa Rica, six second-growth forests and two old-growth reference forests. We used diversity profiles to show successional trajectories in measures of observed, asymptotic and standardized tree diversity and evenness as well as sample completeness. We randomly subsampled 1-ha plot data to evaluate how well smaller spatial subsamples would have captured temporal trajectories. Annual surveys in eight 1-ha plots were missing substantial numbers of rare or infrequent species. Older second-growth sites showed consistent declines in tree diversity, whereas younger sites showed fluctuating patterns or increases. Subsample areas of 0.5 ha or greater were sufficient to infer the diversity of abundant species, but smaller subsamples failed to capture temporal trajectories of species richness and yielded positively biased estimates of evenness. In tropical forest regions with high levels of diversity, species diversity from small sample plots should be assessed using methods that incorporate abundance information and that standardize for sample coverage. This article is part of the theme issue 'Understanding forest landscape restoration: reinforcing scientific foundations for the UN Decade on Ecosystem Restoration'.

Forests that regrow naturally on abandoned fields are important for restoring biodiversity and ecosystem services, but can they also preserve the distinct regional tree floras? Using the floristic composition of 1215 early successional forests (≤20 years) in 75 human-modified landscapes across the Neotropic realm, we identified 14 distinct floristic groups, with a between-group dissimilarity of 0.97. Floristic groups were associated with location, bioregions, soil pH, temperature seasonality, and water availability. Hence, there is large continental-scale variation in the species composition of early successional forests, which is mainly associated with biogeographic and environmental factors but not with human disturbance indicators. This floristic distinctiveness is partially driven by regionally restricted species belonging to widespread genera. Early secondary forests contribute therefore to restoring and conserving the distinctiveness of bioregions across the Neotropical realm, and forest restoration initiatives should use local species to assure that these distinct floras are maintained.

Natural forest regrowth is a cost-effective, nature-based solution for biodiversity recovery, yet different socioenvironmental factors can lead to variable outcomes. A critical knowledge gap in forest restoration planning is how to predict where natural forest regrowth is likely to lead to high levels of biodiversity recovery, which is an indicator of conservation value and the potential provisioning of diverse ecosystem services. We sought to predict and map landscape-scale recovery of species richness and total abundance of vertebrates, invertebrates, and plants in tropical and subtropical second-growth forests to inform spatial restoration planning. First, we conducted a global meta-analysis to quantify the extent to which recovery of species richness and total abundance in second-growth forests deviated from biodiversity values in reference old-growth forests in the same landscape. Second, we employed a machine-learning algorithm and a comprehensive set of socioenvironmental factors to spatially predict landscape-scale deviation and map it. Models explained on average 34% of observed variance in recovery (range 9-51%). Landscape-scale biodiversity recovery in second-growth forests was spatially predicted based on socioenvironmental landscape factors (human demography, land use and cover, anthropogenic and natural disturbance, ecosystem productivity, and topography and soil chemistry); was significantly higher for species richness than for total abundance for vertebrates (median range-adjusted predicted deviation 0.09 vs. 0.34) and invertebrates (0.2 vs. 0.35) but not for plants (which showed a similar recovery for both metrics [0.24 vs. 0.25]); and was positively correlated for total abundance of plant and vertebrate species (Pearson r = 0.45, p = 0.001). Our approach can help identify tropical and subtropical forest landscapes with high potential for biodiversity recovery through natural forest regrowth., (© 2021 Society for Conservation Biology.)

Tropical forests disappear rapidly because of deforestation, yet they have the potential to regrow naturally on abandoned lands. We analyze how 12 forest attributes recover during secondary succession and how their recovery is interrelated using 77 sites across the tropics. Tropical forests are highly resilient to low-intensity land use; after 20 years, forest attributes attain 78% (33 to 100%) of their old-growth values. Recovery to 90% of old-growth values is fastest for soil (<1 decade) and plant functioning (<2.5 decades), intermediate for structure and species diversity (2.5 to 6 decades), and slowest for biomass and species composition (>12 decades). Network analysis shows three independent clusters of attribute recovery, related to structure, species diversity, and species composition. Secondary forests should be embraced as a low-cost, natural solution for ecosystem restoration, climate change mitigation, and biodiversity conservation.

One-third of all Neotropical forests are secondary forests that regrow naturally after agricultural use through secondary succession. We need to understand better how and why succession varies across environmental gradients and broad geographic scales. Here, we analyze functional recovery using community data on seven plant characteristics (traits) of 1,016 forest plots from 30 chronosequence sites across the Neotropics. By analyzing communities in terms of their traits, we enhance understanding of the mechanisms of succession, assess ecosystem recovery, and use these insights to propose successful forest restoration strategies. Wet and dry forests diverged markedly for several traits that increase growth rate in wet forests but come at the expense of reduced drought tolerance, delay, or avoidance, which is important in seasonally dry forests. Dry and wet forests showed different successional pathways for several traits. In dry forests, species turnover is driven by drought tolerance traits that are important early in succession and in wet forests by shade tolerance traits that are important later in succession. In both forests, deciduous and compound-leaved trees decreased with forest age, probably because microclimatic conditions became less hot and dry. Our results suggest that climatic water availability drives functional recovery by influencing the start and trajectory of succession, resulting in a convergence of community trait values with forest age when vegetation cover builds up. Within plots, the range in functional trait values increased with age. Based on the observed successional trait changes, we indicate the consequences for carbon and nutrient cycling and propose an ecologically sound strategy to improve forest restoration success., Competing Interests: The authors declare no competing interest.

The UN Decade on Ecosystem Restoration offers immense potential to return hundreds of millions of hectares of degraded tropical landscapes to functioning ecosystems. Well-designed restoration can tackle multiple Sustainable Development Goals, driving synergistic benefits for biodiversity, ecosystem services, agricultural and timber production, and local livelihoods at large spatial scales. To deliver on this potential, restoration efforts must recognise and reduce trade-offs among objectives, and minimize competition with food production and conservation of native ecosystems. Restoration initiatives also need to confront core environmental challenges of climate change and inappropriate planting in savanna biomes, be robustly funded over the long term, and address issues of poor governance, inadequate land tenure, and socio-cultural disparities in benefits and costs. Tackling these issues using the landscape approach is vital to realising the potential for restoration to break the cycle of land degradation and poverty, and deliver on its core environmental and social promises., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2021 Elsevier Inc. All rights reserved.)

Tropical forest regeneration after abandonment of former agricultural land depends critically on the input of tree seeds, yet seed dispersal is increasingly disrupted in contemporary human-modified landscapes. Here, we introduce the concept of seed-rain-successional feedbacks as a deterministic process in which seed rain is shaped by successional dynamics internal to a forest site and that acts to reinforce priority effects. We used a combination of time series and chronosequence approaches to investigate how the quantity and taxonomic and functional composition of seed rain change during succession and to evaluate the strength of seed-rain-successional feedbacks, relative to other deterministic and stochastic mechanisms, in secondary wet forests of Costa Rica. We found that both successional niches and seed-rain-successional feedbacks shaped successional trajectories in the seed rain. Determinism due to successional niche assembly was supported by the increasing convergence of community structure to that of a mature forest, in terms of both functional and taxonomic composition. With successional age, the proportions of large-seeded, shade-tolerant species in the seed rain increased, whereas the proportion of animal-dispersed species did not change significantly. Seed-rain-successional feedbacks increased in strength with successional age, as the proportion of immigrant seeds (species not locally represented in the site) decreased with successional age, and the composition of the seed rain became more similar to that of the adult trees at the forest site. The deterministic assembly generated by seed-rain-successional feedback likely contributed to the increasing divergence of secondary forest sites from each other during succession. To the extent that human modification of tropical forest landscapes reduces connectivity via factors such as forest cover loss, our results suggest that seed-rain-successional feedbacks are likely to increasingly shape regeneration trajectories in and amplify floristic heterogeneity among tropical secondary forests., (© 2021 by the Ecological Society of America.)

This study describes the 35-year progression of activities in the Pontal do Paranapanema region of São Paulo State, Brazil. These activities began as a research project on the conservation ecology of the highly endangered Black Lion Tamarin and broadened into a landscape-scale restoration and conservation project involving the active participation of hundreds of landless families that colonized the region. Rather than viewing these colonists as a threat, a non-governmental organization arose to address their needs, providing training and support livelihoods. Local communities were engaged in conservation and restoration activities focused on studying the movement patterns of endangered species, environmental education programmes, planting native trees along riparian corridors, establishing coffee agroforestry plantings and initiating community-managed nurseries for the production of local native seedlings and non-native fruit trees. Farmers gained knowledge, income and food security, and developed a sense of ownership and shared responsibility for protecting wildlife, conserving forest fragments and restoring forests. Land sharing and restoring forest functions within an agricultural landscape matrix created new opportunities for people and endangered wildlife. We explore how key factors and partnerships critically influenced the landscape trajectory and conclude with lessons learned that may be relevant to sustainable landscape initiatives in other contexts., Competing Interests: The authors have no competing interests., (© 2020 The Authors.)

Extensive ecosystem restoration is increasingly seen as being central to conserving biodiversity 1 and stabilizing the climate of the Earth 2 . Although ambitious national and global targets have been set, global priority areas that account for spatial variation in benefits and costs have yet to be identified. Here we develop and apply a multicriteria optimization approach that identifies priority areas for restoration across all terrestrial biomes, and estimates their benefits and costs. We find that restoring 15% of converted lands in priority areas could avoid 60% of expected extinctions while sequestering 299 gigatonnes of CO 2 -30% of the total CO 2 increase in the atmosphere since the Industrial Revolution. The inclusion of several biomes is key to achieving multiple benefits. Cost effectiveness can increase up to 13-fold when spatial allocation is optimized using our multicriteria approach, which highlights the importance of spatial planning. Our results confirm the vast potential contributions of restoration to addressing global challenges, while underscoring the necessity of pursuing these goals synergistically.

Managed tropical forests are a globally important carbon pool, but the effects of logging and thinning intensities on long-term biomass dynamics are poorly known. We investigated the demographic mechanisms of above-ground biomass recovery over 48 years in an Australian tropical forest following four silvicultural treatments: selective logging only as a control and selective logging followed by low-, medium- and high-intensity thinning. Initial biomass recovery rates following thinning were poor predictors of the long-term changes. Initial biomass recovery from 1969 to 1973 was slow and was largely concentrated on an increase in the biomass of residual stems. From 1973 to 1997, above ground biomass (AGB) increased almost linearly, with a similar slope for all sites. From 1997 to 2015, the rate of biomass accumulation slowed, especially for the L treatment. All thinning treatments stimulated more recruitment and regrowth of non-harvested remaining trees compared to the untreated control. Biomass at both the low and medium intensity treatments has almost fully recovered to 98% and 97% of pre-logging biomass levels respectively. The predicted times of complete above-ground biomass recovery for the logging only and high intensity treatments are 55 and 77 years respectively. The slower biomass recovery at the logging only site was largely due to increased mortality in the last measurement period. The slower recovery of the high intensity site was due to a combination of a higher initial reduction in biomass from thinning and the increased mortality in the last measurement period. The high mortality rates in the most recent measurement period are likely due to the impacts of two cyclones that impacted the study site. Our results suggest that it will take at least around 50 years for this site to recover to its pre-harvest biomass, much longer than many of the cutting cycles currently used in tropical forest management., (Copyright © 2020 Elsevier B.V. All rights reserved.)

To constrain global warming, we must strongly curtail greenhouse gas emissions and capture excess atmospheric carbon dioxide 1,2 . Regrowing natural forests is a prominent strategy for capturing additional carbon 3 , but accurate assessments of its potential are limited by uncertainty and variability in carbon accumulation rates 2,3 . To assess why and where rates differ, here we compile 13,112 georeferenced measurements of carbon accumulation. Climatic factors explain variation in rates better than land-use history, so we combine the field measurements with 66 environmental covariate layers to create a global, one-kilometre-resolution map of potential aboveground carbon accumulation rates for the first 30 years of natural forest regrowth. This map shows over 100-fold variation in rates across the globe, and indicates that default rates from the Intergovernmental Panel on Climate Change (IPCC) 4,5 may underestimate aboveground carbon accumulation rates by 32 per cent on average and do not capture eight-fold variation within ecozones. Conversely, we conclude that maximum climate mitigation potential from natural forest regrowth is 11 per cent lower than previously reported 3 owing to the use of overly high rates for the location of potential new forest. Although our data compilation includes more studies and sites than previous efforts, our results depend on data availability, which is concentrated in ten countries, and data quality, which varies across studies. However, the plots cover most of the environmental conditions across the areas for which we predicted carbon accumulation rates (except for northern Africa and northeast Asia). We therefore provide a robust and globally consistent tool for assessing natural forest regrowth as a climate mitigation strategy.

Legumes provide an essential service to ecosystems by capturing nitrogen from the atmosphere and delivering it to the soil, where it may then be available to other plants. However, this facilitation by legumes has not been widely studied in global tropical forests. Demographic data from 11 large forest plots (16-60 ha) ranging from 5.25° S to 29.25° N latitude show that within forests, leguminous trees have a larger effect on neighbor diversity than non-legumes. Where soil nitrogen is high, most legume species have higher neighbor diversity than non-legumes. Where soil nitrogen is low, most legumes have lower neighbor diversity than non-legumes. No facilitation effect on neighbor basal area was observed in either high or low soil N conditions. The legume-soil nitrogen positive feedback that promotes tree diversity has both theoretical implications for understanding species coexistence in diverse forests, and practical implications for the utilization of legumes in forest restoration.

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

Over 140 Mha of restoration commitments have been pledged across the global tropics, yet guidance is needed to identify those landscapes where implementation is likely to provide the greatest potential benefits and cost-effective outcomes. By overlaying seven recent, peer-reviewed spatial datasets as proxies for socioenvironmental benefits and feasibility of restoration, we identified restoration opportunities (areas with higher potential return of benefits and feasibility) in lowland tropical rainforest landscapes. We found restoration opportunities throughout the tropics. Areas scoring in the top 10% (i.e., restoration hotspots) are located largely within conservation hotspots (88%) and in countries committed to the Bonn Challenge (73%), a global effort to restore 350 Mha by 2030. However, restoration hotspots represented only a small portion (19.1%) of the Key Biodiversity Area network. Concentrating restoration investments in landscapes with high benefits and feasibility would maximize the potential to mitigate anthropogenic impacts and improve human well-being.

Tropical forests are converted at an alarming rate for agricultural use and pastureland, but also regrow naturally through secondary succession. For successful forest restoration, it is essential to understand the mechanisms of secondary succession. These mechanisms may vary across forest types, but analyses across broad spatial scales are lacking. Here, we analyse forest recovery using 1,403 plots that differ in age since agricultural abandonment from 50 sites across the Neotropics. We analyse changes in community composition using species-specific stem wood density (WD), which is a key trait for plant growth, survival and forest carbon storage. In wet forest, succession proceeds from low towards high community WD (acquisitive towards conservative trait values), in line with standard successional theory. However, in dry forest, succession proceeds from high towards low community WD (conservative towards acquisitive trait values), probably because high WD reflects drought tolerance in harsh early successional environments. Dry season intensity drives WD recovery by influencing the start and trajectory of succession, resulting in convergence of the community WD over time as vegetation cover builds up. These ecological insights can be used to improve species selection for reforestation. Reforestation species selected to establish a first protective canopy layer should, among other criteria, ideally have a similar WD to the early successional communities that dominate under the prevailing macroclimatic conditions.

Regenerating tropical forests have an immense capacity to capture carbon and harbor biodiversity. The recuperation of the nitrogen cycle following disturbance can fuel biomass regeneration, but few studies have evaluated the successional dynamics of nitrogen and nitrogen inputs in tropical forests. We assessed symbiotic and asymbiotic nitrogen fixation, soil inorganic nitrogen concentrations, and tree growth in a well-studied series of five tropical forest plots ranging from 19 yr in age to old-growth forests. Wet-season soil inorganic nitrogen concentrations were high in all plots, peaking in the 29-yr-old plot. Inputs from symbiotic nitrogen fixation declined through succession, while asymbiotic nitrogen fixation peaked in the 37-yr-old plot. Consequently, the dominant nitrogen fixation input switched from symbiotic fixation in the younger plots to asymbiotic fixation in the older plots. Tree growth was highest in the youngest plots and declined through succession. Interestingly, symbiotic nitrogen fixation was negatively correlated with the basal area of nitrogen-fixing trees across our study plots, highlighting the danger in using nitrogen-fixing trees as a proxy for rates of symbiotic nitrogen fixation. Our results demonstrate that the nitrogen cycle has largely recuperated by 19 yr following disturbance, allowing for rapid biomass regeneration at our site. This work provides important insight into the sources and dynamics of nitrogen that support growth and carbon capture in regenerating Neotropical forests., (© 2019 by the Ecological Society of America.)

Old-growth tropical forests harbor an immense diversity of tree species but are rapidly being cleared, while secondary forests that regrow on abandoned agricultural lands increase in extent. We assess how tree species richness and composition recover during secondary succession across gradients in environmental conditions and anthropogenic disturbance in an unprecedented multisite analysis for the Neotropics. Secondary forests recover remarkably fast in species richness but slowly in species composition. Secondary forests take a median time of five decades to recover the species richness of old-growth forest (80% recovery after 20 years) based on rarefaction analysis. Full recovery of species composition takes centuries (only 34% recovery after 20 years). A dual strategy that maintains both old-growth forests and species-rich secondary forests is therefore crucial for biodiversity conservation in human-modified tropical landscapes.

International commitments for ecosystem restoration add up to one-quarter of the world's arable land. Fulfilling them would ease global challenges such as climate change and biodiversity decline but could displace food production and impose financial costs on farmers. Here, we present a restoration prioritization approach capable of revealing these synergies and trade-offs, incorporating ecological and economic efficiencies of scale and modelling specific policy options. Using an actual large-scale restoration target of the Atlantic Forest hotspot, we show that our approach can deliver an eightfold increase in cost-effectiveness for biodiversity conservation compared with a baseline of non-systematic restoration. A compromise solution avoids 26% of the biome's current extinction debt of 2,864 plant and animal species (an increase of 257% compared with the baseline). Moreover, this solution sequesters 1 billion tonnes of CO 2 -equivalent (a 105% increase) while reducing costs by US$28 billion (a 57% decrease). Seizing similar opportunities elsewhere would offer substantial contributions to some of the greatest challenges for humankind.

One of the most intriguing questions in plant ecology is which evolutionary strategy allows widely distributed species to increase their ecological range and grow in changing environmental conditions. Phenotypic plasticity and local adaptations are major processes governing species range margins, but little is known about their relative contribution for tree species distribution in tropical forest regions. We investigated the relative role of phenotypic plasticity and local adaptation in the ecological distribution of the widespread palm Euterpe edulis in the Brazilian Atlantic Forest. Genetic sampling and experiments were performed in old-growth remnants of two forest types with higher (Seasonal Semideciduous Forests vs. Submontane Rainforest) and lower biogeographic association and environmental similarities (Submontane Rainforest vs. Restinga Forest). We first assessed the molecular genetic differentiation among populations, focusing on the group of loci potentially under selection in each forest, using single-nucleotide polymorphism (SNPs) outliers. Further, we looked for potential adaptive divergence among populations in a common garden experiment and in reciprocal transplants for two plant development phases: seedling establishment and sapling growth. Analysis with outlier loci indicated that all individuals from the Semideciduous Forest formed a single group, while another group was formed by overlapping individuals from Submontane Rainforest and Restinga Forest. Molecular differentiation was corroborated by reciprocal transplants, which yielded strong evidence of local adaptations for seedling establishment in the biogeographically divergent Rainforest and Semideciduous Forest, but not for Restinga Forest and Submontane Rainforest. Phenotypic plasticity for palm seedling establishment favors range expansion to biogeographically related or recently colonized forest types, while persistence in the newly colonized ecosystem may be favored by local adaptations if climatic conditions diverge over time, reducing gene flow between populations. SNPs obtained by next-generation sequencing can help exploring adaptive genetic variation in tropical trees, which impose several challenges to the use of reciprocal transplants.

The nutrient demands of regrowing tropical forests are partly satisfied by nitrogen-fixing legume trees, but our understanding of the abundance of those species is biased towards wet tropical regions. Here we show how the abundance of Leguminosae is affected by both recovery from disturbance and large-scale rainfall gradients through a synthesis of forest inventory plots from a network of 42 Neotropical forest chronosequences. During the first three decades of natural forest regeneration, legume basal area is twice as high in dry compared with wet secondary forests. The tremendous ecological success of legumes in recently disturbed, water-limited forests is likely to be related to both their reduced leaflet size and ability to fix N 2 , which together enhance legume drought tolerance and water-use efficiency. Earth system models should incorporate these large-scale successional and climatic patterns of legume dominance to provide more accurate estimates of the maximum potential for natural nitrogen fixation across tropical forests.

Tropical secondary forests (TSF) are a global carbon sink of 1.6 Pg C/year. However, TSF carbon uptake is estimated using chronosequence studies that assume differently aged forests can be used to predict change in aboveground biomass density (AGBD) over time. We tested this assumption using two airborne lidar datasets separated by 11.5 years over a Neotropical landscape. Using data from 1998, we predicted canopy height and AGBD within 1.1 and 10.3% of observations in 2009, with higher accuracy for forest height than AGBD and for older TSFs in comparison to younger ones. This result indicates that the space-for-time assumption is robust at the landscape-scale. However, since lidar measurements of secondary tropical forest are rare, we used the 1998 lidar dataset to test how well plot-based studies quantify the mean TSF height and biomass in a landscape. We found that the sample area required to produce estimates of height or AGBD close to the landscape mean is larger than the typical area sampled in secondary forest chronosequence studies. For example, estimating AGBD within 10% of the landscape mean requires more than thirty 0.1 ha plots per age class, and more total area for larger plots. We conclude that under-sampling in ground-based studies may introduce error into estimations of the TSF carbon sink, and that this error can be reduced by more extensive use of lidar measurements., (© 2017 John Wiley & Sons Ltd.)

Mixed tree plantings and natural regeneration are the main restoration approaches for recovering tropical forests worldwide. Despite substantial differences in implementation costs between these methods, little is known regarding how they differ in terms of ecological outcomes, which is key information for guiding decision making and cost-effective restoration planning. Here, we compared the early ecological outcomes of natural regeneration and tree plantations for restoring the Brazilian Atlantic Forest in agricultural landscapes. We assessed and compared vegetation structure and composition in young (7-20 yr old) mixed tree plantings (PL), second-growth tropical forests established on former pastures (SGp), on former Eucalyptus spp. plantations (SGe), and in old-growth reference forests (Ref). We sampled trees with diameter at breast height (DBH) 1-5 cm (saplings) and trees at DBH > 5 cm (trees) in a total of 32 20 × 45 m plots established in these landscapes. Overall, the ecological outcomes of natural regeneration and restoration plantations were markedly different. SGe forests showed higher abundance of large (DBH > 20 cm) nonnative species, of which 98% were resprouting Eucalyptus trees, than SGp and PL, and higher total aboveground biomass; however, aboveground biomass of native species was higher in PL than in SGe. PL forests had lower abundance of native saplings and lianas than both naturally established second-growth forests, and lower proportion of animal dispersed saplings than SGe, probably due to higher isolation from native forest remnants. Rarefied species richness of trees was lower in SGp, intermediate in SGe and Ref and higher in PL, whereas rarefied species richness of saplings was higher in SG than in Ref. Species composition differed considerably among regeneration types. Although these forests are inevitably bound to specific landscape contexts and may present varying outcomes as they develop through longer time frames, the ecological particularities of forests established through different restoration approaches indicate that naturally established forests may not show similar outcomes to mixed tree plantings. The results of this study underscore the importance that restoration decisions need to be based on more robust expectations of outcomes that allow for a better analysis of the cost-effectiveness of different restoration approaches before scaling-up forest restoration in the tropics., (© 2017 by the Ecological Society of America.)

Knowledge about the biogeographic affinities of the world's tropical forests helps to better understand regional differences in forest structure, diversity, composition, and dynamics. Such understanding will enable anticipation of region-specific responses to global environmental change. Modern phylogenies, in combination with broad coverage of species inventory data, now allow for global biogeographic analyses that take species evolutionary distance into account. Here we present a classification of the world's tropical forests based on their phylogenetic similarity. We identify five principal floristic regions and their floristic relationships: ( i ) Indo-Pacific, ( ii ) Subtropical, ( iii ) African, ( iv ) American, and ( v ) Dry forests. Our results do not support the traditional neo- versus paleotropical forest division but instead separate the combined American and African forests from their Indo-Pacific counterparts. We also find indications for the existence of a global dry forest region, with representatives in America, Africa, Madagascar, and India. Additionally, a northern-hemisphere Subtropical forest region was identified with representatives in Asia and America, providing support for a link between Asian and American northern-hemisphere forests., Competing Interests: Conflict of interest statement: V.A.-R., K.B.-G., B.B., F.Q.B., N.F., M.K., W.F.L., S. G. Letcher, C.B.S., D.S., T. Stevart, and S. Wiser have coauthored papers with Jens-Christian Svenning in the past 48 months. A.M.L. and Jens-Christian Svenning are both affiliated with Aarhus University., (Copyright © 2018 the Author(s). Published by PNAS.)

Is active restoration the best approach to achieve ecological restoration success (the return to a reference condition, that is, old-growth forest) when compared to natural regeneration in tropical forests? Our meta-analysis of 133 studies demonstrated that natural regeneration surpasses active restoration in achieving tropical forest restoration success for all three biodiversity groups (plants, birds, and invertebrates) and five measures of vegetation structure (cover, density, litter, biomass, and height) tested. Restoration success for biodiversity and vegetation structure was 34 to 56% and 19 to 56% higher in natural regeneration than in active restoration systems, respectively, after controlling for key biotic and abiotic factors (forest cover, precipitation, time elapsed since restoration started, and past disturbance). Biodiversity responses were based primarily on ecological metrics of abundance and species richness (74%), both of which take far less time to achieve restoration success than similarity and composition. This finding challenges the widely held notion that natural forest regeneration has limited conservation value and that active restoration should be the default ecological restoration strategy. The proposition that active restoration achieves greater restoration success than natural regeneration may have arisen because previous comparisons lacked controls for biotic and abiotic factors; we also did not find any difference between active restoration and natural regeneration outcomes for vegetation structure when we did not control for these factors. Future policy priorities should align the identified patterns of biophysical and ecological conditions where each or both restoration approaches are more successful, cost-effective, and compatible with socioeconomic incentives for tropical forest restoration.

Whether successional forests converge towards an equilibrium in species composition remains an elusive question, hampered by high idiosyncrasy in successional dynamics. Based on long-term tree monitoring in second-growth (SG) and old-growth (OG) forests in Costa Rica, we show that patterns of convergence between pairs of forest stands depend upon the relative abundance of species exhibiting distinct responses to the successional gradient. For instance, forest generalists contributed to convergence between SG and OG forests, whereas rare species and old-growth specialists were a source of divergence. Overall, opposing trends in taxonomic similarity among different subsets of species nullified each other, producing a net outcome of stasis over time. Our results offer an explanation for the limited convergence observed between pairwise communities and suggest that rare species and old-growth specialists may be prone to dispersal limitation, while the dynamics of generalists and second-growth specialists are more predictable, enhancing resilience in tropical secondary forests., (© 2017 John Wiley & Sons Ltd/CNRS.)

Estimating the species, phylogenetic, and functional diversity of a community is challenging because rare species are often undetected, even with intensive sampling. The Good-Turing frequency formula, originally developed for cryptography, estimates in an ecological context the true frequencies of rare species in a single assemblage based on an incomplete sample of individuals. Until now, this formula has never been used to estimate undetected species, phylogenetic, and functional diversity. Here, we first generalize the Good-Turing formula to incomplete sampling of two assemblages. The original formula and its two-assemblage generalization provide a novel and unified approach to notation, terminology, and estimation of undetected biological diversity. For species richness, the Good-Turing framework offers an intuitive way to derive the non-parametric estimators of the undetected species richness in a single assemblage, and of the undetected species shared between two assemblages. For phylogenetic diversity, the unified approach leads to an estimator of the undetected Faith's phylogenetic diversity (PD, the total length of undetected branches of a phylogenetic tree connecting all species), as well as a new estimator of undetected PD shared between two phylogenetic trees. For functional diversity based on species traits, the unified approach yields a new estimator of undetected Walker et al.'s functional attribute diversity (FAD, the total species-pairwise functional distance) in a single assemblage, as well as a new estimator of undetected FAD shared between two assemblages. Although some of the resulting estimators have been previously published (but derived with traditional mathematical inequalities), all taxonomic, phylogenetic, and functional diversity estimators are now derived under the same framework. All the derived estimators are theoretically lower bounds of the corresponding undetected diversities; our approach reveals the sufficient conditions under which the estimators are nearly unbiased, thus offering new insights. Simulation results are reported to numerically verify the performance of the derived estimators. We illustrate all estimators and assess their sampling uncertainty with an empirical dataset for Brazilian rain forest trees. These estimators should be widely applicable to many current problems in ecology, such as the effects of climate change on spatial and temporal beta diversity and the contribution of trait diversity to ecosystem multi-functionality., (© 2017 by the Ecological Society of America.)

More than half of the world's tropical forests are currently recovering from human land use, and this regenerating biomass now represents the largest carbon (C)-capturing potential on Earth. How quickly these forests regenerate is now a central concern for both conservation and global climate-modeling efforts. Symbiotic nitrogen-fixing trees are thought to provide much of the nitrogen (N) required to fuel tropical secondary regrowth and therefore to drive the rate of forest regeneration, yet we have a poor understanding of how these N fixers influence the trees around them. Do they promote forest growth, as expected if the new N they fix facilitates neighboring trees? Or do they suppress growth, as expected if competitive inhibition of their neighbors is strong? Using 17 consecutive years of data from tropical rainforest plots in Costa Rica that range from 10 y since abandonment to old-growth forest, we assessed how N fixers influenced the growth of forest stands and the demographic rates of neighboring trees. Surprisingly, we found no evidence that N fixers facilitate biomass regeneration in these forests. At the hectare scale, plots with more N-fixing trees grew slower. At the individual scale, N fixers inhibited their neighbors even more strongly than did nonfixing trees. These results provide strong evidence that N-fixing trees do not always serve the facilitative role to neighboring trees during tropical forest regeneration that is expected given their N inputs into these systems., Competing Interests: The authors declare no conflict of interest.

Old-growth tropical forests are being extensively deforested and fragmented worldwide. Yet forest recovery through succession has led to an expansion of secondary forests in human-modified tropical landscapes (HMTLs). Secondary forests thus emerge as a potential repository for tropical biodiversity, and also as a source of essential ecosystem functions and services in HMTLs. Such critical roles are controversial, however, as they depend on successional, landscape and socio-economic dynamics, which can vary widely within and across landscapes and regions. Understanding the main drivers of successional pathways of disturbed tropical forests is critically needed for improving management, conservation, and restoration strategies. Here, we combine emerging knowledge from tropical forest succession, forest fragmentation and landscape ecology research to identify the main driving forces shaping successional pathways at different spatial scales. We also explore causal connections between land-use dynamics and the level of predictability of successional pathways, and examine potential implications of such connections to determine the importance of secondary forests for biodiversity conservation in HMTLs. We show that secondary succession (SS) in tropical landscapes is a multifactorial phenomenon affected by a myriad of forces operating at multiple spatio-temporal scales. SS is relatively fast and more predictable in recently modified landscapes and where well-preserved biodiversity-rich native forests are still present in the landscape. Yet the increasing variation in landscape spatial configuration and matrix heterogeneity in landscapes with intermediate levels of disturbance increases the uncertainty of successional pathways. In landscapes that have suffered extensive and intensive human disturbances, however, succession can be slow or arrested, with impoverished assemblages and reduced potential to deliver ecosystem functions and services. We conclude that: (i) succession must be examined using more comprehensive explanatory models, providing information about the forces affecting not only the presence but also the persistence of species and ecological groups, particularly of those taxa expected to be extirpated from HMTLs; (ii) SS research should integrate new aspects from forest fragmentation and landscape ecology research to address accurately the potential of secondary forests to serve as biodiversity repositories; and (iii) secondary forest stands, as a dynamic component of HMTLs, must be incorporated as key elements of conservation planning; i.e. secondary forest stands must be actively managed (e.g. using assisted forest restoration) according to conservation goals at broad spatial scales., (© 2015 Cambridge Philosophical Society.)

We present a historical overview of forest concepts and definitions, linking these changes with distinct perspectives and management objectives. Policies dealing with a broad range of forest issues are often based on definitions created for the purpose of assessing global forest stocks, which do not distinguish between natural and planted forests or reforests, and which have not proved useful in assessing national and global rates of forest regrowth and restoration. Implementing and monitoring forest and landscape restoration requires additional approaches to defining and assessing forests that reveal the qualities and trajectories of forest patches in a spatially and temporally dynamic landscape matrix. New technologies and participatory assessment of forest states and trajectories offer the potential to operationalize such definitions. Purpose-built and contextualized definitions are needed to support policies that successfully protect, sustain, and regrow forests at national and global scales. We provide a framework to illustrate how different management objectives drive the relative importance of different aspects of forest state, dynamics, and landscape context.

Re-establishing connectivity between protected areas isolated by habitat clearing is a key conservation goal in the humid tropics. In northeastern Costa Rica, payments for environmental services (PES) and a government ban on deforestation have subsidized forest protection and reforestation in the San Juan-La Selva Biological Corridor (SJLSBC), resulting in a decline in mature forest loss and the expansion of tree plantations. We use field studies and graph models to assess how conservation efforts have altered functional connectivity over the last 25 years for four species of insectivorous understory birds. Field playback studies assessed how reforestation habitat quality affected the willingness of Myrmeciza exsul, Henicorhina leucosticta, Thamnophilus atrinucha, and Glyphorynchus spirurus to travel outside forest habitat for territorial defense. Observed travel distances were greatest in nonnative and native tree plantations with high understory stem density, regardless of overstory composition. In contrast, tree plantations with low stem density had travel responses comparable to open pasture for three of the four bird species. We modeled landscape connectivity for each species using graph models based on varying possible travel distances in tree plantations, gallery forests, and pastures. From 1986 to 2011, connectivity for all species declined in the SJLSBC landscape (5825 km 2 ) by 14% to 21% despite only a 4.9% net loss in forest area and the rapid expansion of tree plantations over 2% of the landscape. Plantation placement in the landscape limited their potential facilitation of connectivity because they were located either far from forest cover or within already contiguous forest areas. We mapped current connectivity bottlenecks and identified priority areas for future reforestation. We estimate that reforestation of priority areas could improve connectivity by 2% with only a 1% gain in forest cover, an impressive gain given the small area reforested. Results indicate key locations where spatial targeting of PES within the SJLSBC study region would protect existing forest connectivity and enhance the connectivity benefits of reforestation., (© 2016 by the Ecological Society of America.)