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Published online: 15 April 2019 ; On the African continent, the population is expected to expand fourfold in the next century, which will increasingly impact the global carbon cycle and biodiversity conservation. Therefore, it is of vital importance to understand how carbon stocks and community assembly recover after slash‐and‐burn events in tropical second growth forests. We inventoried a chronosequence of 15 1‐ha plots in lowland tropical forest of the central Congo Basin and evaluated changes in aboveground and soil organic carbon stocks and in tree species diversity, functional composition, and community‐weighted functional traits with succession. We aimed to track long‐term recovery trajectories of species and carbon stocks in secondary forests, comparing 5 to 200 + year old secondary forest with reference primary forest. Along the successional gradient, the functional composition followed a trajectory from resource acquisition to resource conservation, except for nitrogen‐related leaf traits. Despite a fast, initial recovery of species diversity and functional composition, there were still important structural and carbon stock differences between old growth secondary and pristine forest, which suggests that a full recovery of secondary forests might take much longer than currently shown. As such, the aboveground carbon stocks of 200 + year old forest were only 57% of those in the pristine reference forest, which suggests a slow recovery of aboveground carbon stocks, although more research is needed to confirm this observation. The results of this study highlight the need for more in‐depth studies on forest recovery in Central Africa, to gain insight into the processes that control biodiversity and carbon stock recovery. ; Federal Science Policy Office, Belgium ; Peer Review

Reliable tree height-diameter (H-D) allometric equations are a key tool for the estimation of forest productivity and Above Ground Biomass (AGB). Most existing H-D allometric equations developed for the tropical region are based on large-scale multi-species datasets, and their use to derive information on productivity and AGB at the species level is prone to uncertainties. The single-species H-D allometric equations available are mainly focused on monocultures or stands with simple tree species mixtures and did not account for the site effects. Here we measured the height and diameter of 2,288 trees of the emergent tree species Pericopsis elata (Harms) Meeuwen in the Democratic Republic of the Congo (DRC) and in Cameroon. We first examined how accurate multispecies H-D allometric equations are in predicting the total height of P. elata. We then tested whether single-species H-D allometric equations vary between sites. We developed the first H-D allometric equation of P. elata and tested whether and how stand-level and environmental variables induce changes in H-D allometric relationship of P. elata at the regional level. We additionally evaluated whether tree-level variables are important at the local level where climate and stand development stage are expected to be less variable. We found that pantropical, regional and local H-D allometric equations significantly underestimate the total height of P. elata. The local multi-species H-D allometric equation developed for Yangambi showed the highest underestimation in all the studied sites. This result supports the need for an H-D allometric equation specific for P. elata. The species-level H-D allometric equation developed showed significant underestimations for trees from the disturbed and undisturbed forests in DRC, while overestimations were observed for similar sites in Cameroon. Using a mixed-effect H-D allometric equation, we showed that even within a single species, a substantial variation exists between sites. This variation showed to be driven by the differences in the maximum asymptotic height (Hmax) between sites. We found that P. elata trees are taller and attain higher Hmax in DRC than in Cameroon. The basal area showed to be a significant covariate accounting for the site effects at the regional-scale where climate variables showed minor effects. However, at the local-scale, none of climate or stand variables showed to be significant. Local-scale variation showed to be associated with differences in light availability, highlighting the potential of management options that shape the local environment in driving species productivity.

Reliable tree height-diameter (H-D) allometric equations are a key tool for the estimation of forest productivity and Above Ground Biomass (AGB). Most existing H-D allometric equations developed for the tropical region are based on large-scale multi-species datasets, and their use to derive information on productivity and AGB at the species level is prone to uncertainties. The single-species H-D allometric equations available are mainly focused on monocultures or stands with simple tree species mixtures and did not account for the site effects. Here we measured the height and diameter of 2,288 trees of the emergent tree species Pericopsis elata (Harms) Meeuwen in the Democratic Republic of the Congo (DRC) and in Cameroon. We first examined how accurate multispecies H-D allometric equations are in predicting the total height of P. elata. We then tested whether single-species H-D allometric equations vary between sites. We developed the first H-D allometric equation of P. elata and tested whether and how stand-level and environmental variables induce changes in H-D allometric relationship of P. elata at the regional level. We additionally evaluated whether tree-level variables are important at the local level where climate and stand development stage are expected to be less variable. We found that pantropical, regional and local H-D allometric equations significantly underestimate the total height of P. elata. The local multi-species H-D allometric equation developed for Yangambi showed the highest underestimation in all the studied sites. This result supports the need for an H-D allometric equation specific for P. elata. The species-level H-D allometric equation developed showed significant underestimations for trees from the disturbed and undisturbed forests in DRC, while overestimations were observed for similar sites in Cameroon. Using a mixed-effect H-D allometric equation, we showed that even within a single species, a substantial variation exists between sites. This variation showed to be driven by the differences in the maximum asymptotic height (Hmax) between sites. We found that P. elata trees are taller and attain higher Hmax in DRC than in Cameroon. The basal area showed to be a significant covariate accounting for the site effects at the regional-scale where climate variables showed minor effects. However, at the local-scale, none of climate or stand variables showed to be significant. Local-scale variation showed to be associated with differences in light availability, highlighting the potential of management options that shape the local environment in driving species productivity. ; Peer reviewed

En Afrique, la phénologie reproductive des arbres tropicaux, majoritairement annuelle, semble présenter une certaine régularité. Cette étude documente les variations intra- et interannuelles de la phénologie reproductive des arbres de la forêt du Mayombe à partir des données historiques de la Réserve de Luki (République démocratique du Congo). Le diamètre de reproduction des espèces exploitées pour le bois, encore largement méconnu, a été également examiné pour les espèces suffisamment représentées dans les données. Le suivi phénologique de 3 642 arbres appartenant à 158 espèces et 39 familles a été réalisé tous les 10 jours de 1948 à 1957. Les statistiques circulaires ont été utilisées pour tester le synchronisme de la phénologie entre arbres, à l'échelle de la communauté, pour la forêt dans son ensemble, et individuellement pour 87 espèces, dont 35 espèces bien représentées (n ≥ 20 arbres), 16 espèces commerciales et 36 autres espèces. Des régressions logistiques ont permis de déterminer le diamètre de fructification (minimum et régulier) de ces espèces. Pour la majorité des espèces, la floraison était régulière, annuelle et largement saisonnière (81,6 %, 71 espèces). Les pics étaient plus marqués pour la floraison que pour la fructification, plus étalée dans le temps, bien que significativement agrégée temporellement. La majorité des arbres et des espèces fleurissaient entre décembre et février, pendant la petite saison sèche, bien que des fleurs et des fruits étaient observables toute l'année à l'échelle de la communauté. Seules 13 espèces ont montré une relation significative entre le diamètre et la reproduction, parmi lesquelles sept espèces de canopée, cinq de sous-bois et une héliophile. Pour ces espèces, la moyenne du diamètre minimum de reproduction était de 17,3 cm. ; Peer reviewed

A diversity of phenological strategies has been reported for tropical tree species. Defoliation and seasonal dormancy of cambial activity inform us on how trees cope with water stress during the dry season, or maximize the use of resources during the rainy season. Here, we study the matching between leaf phenology (unfolding and shedding) and cambial activity for Prioria balsamifera, a key timber species in the Democratic Republic of Congo. In particular, we (i) evaluated the seasonality of cambial activity and synchrony of phenology among trees in response to climate and (ii) identified the seasonality of leaf phenology and its relation with cambial phenology. The study was conducted in the Luki Man and Biosphere Reserve, located in the Mayombe forest at the southern margin of the Congo Basin. Historic defoliation data were collected every ten days using weekly crown observations whereas recent observations involved time-lapse cameras. Cambial pinning was performed on ten trees during 20 months and radius dendrometers were installed on three trees during 13 months. Tree rings were measured on cores from 13 trees and growth synchrony was evaluated. We found that P. balsamifera defoliates annually with a peak observed at the end of the dry season and the beginning of the rainy season. The new leaves unfolded shortly after shedding of the old leaves. The peak defoliation dates varied across years from September 12 to November 14 and the fraction of number of trees that defoliated at a given time was found to be negatively correlated with annual rainfall and temperature; during the dry season, when precipitation and temperatures are the lowest. Wood formation (radial growth), was found to be highly seasonal, with cambial dormancy occurring during the dry season and growth starting at the beginning of the rainy season. Individual ring-width series did not cross date well. The within species variability of leaf phenology and cambial rhythms provides indication about resistance of the population against climatic changes.

A diversity of phenological strategies has been reported for tropical tree species. Defoliation and seasonal dormancy of cambial activity inform us on how trees cope with water stress during the dry season, or maximize the use of resources during the rainy season. Here, we study the matching between leaf phenology (unfolding and shedding) and cambial activity for Prioria balsamifera, a key timber species in the Democratic Republic of Congo. In particular, we (i) evaluated the seasonality of cambial activity and synchrony of phenology among trees in response to climate and (ii) identified the seasonality of leaf phenology and its relation with cambial phenology. The study was conducted in the Luki Man and Biosphere Reserve, located in the Mayombe forest at the southern margin of the Congo Basin. Historic defoliation data were collected every ten days using weekly crown observations whereas recent observations involved timelapse cameras. Cambial pinning was performed on ten trees during 20 months and radius dendrometers were installed on three trees during 13 months. Tree rings were measured on cores from 13 trees and growth synchrony was evaluated. We found that P. balsamifera defoliates annually with a peak observed at the end of the dry season and the beginning of the rainy season. The new leaves unfolded shortly after shedding of the old leaves. The peak defoliation dates varied across years from September 12 to November 14 and the fraction of number of trees that defoliated at a given time was found to be negatively correlated with annual rainfall and temperature; during the dry season, when precipitation and temperatures are the lowest. Wood formation (radial growth), was found to be highly seasonal, with cambial dormancy occurring during the dry season and growth starting at the beginning of the rainy season. Individual ringwidth series did not cross date well. The within species variability of leaf phenology and cambial rhythms provides indication about resistance of the population against climatic changes. ; Peer reviewed

This is the final version of the article. Available from Wiley via the DOI in this record. ; Quantifying the relationship between tree diameter and height is a key component of efforts to estimate biomass and carbon stocks in tropical forests. Although substantial site-to-site variation in height-diameter allometries has been documented, the time consuming nature of measuring all tree heights in an inventory plot means that most studies do not include height, or else use generic pan-tropical or regional allometric equations to estimate height. Using a pan-tropical dataset of 73 plots where at least 150 trees had in-field ground-based height measurements, we examined how the number of trees sampled affects the performance of locally derived height-diameter allometries, and evaluated the performance of different methods for sampling trees for height measurement. Using cross-validation, we found that allometries constructed with just 20 locally measured values could often predict tree height with lower error than regional or climate-based allometries (mean reduction in prediction error = 0.46 m). The predictive performance of locally derived allometries improved with sample size, but with diminishing returns in performance gains when more than 40 trees were sampled. Estimates of stand-level biomass produced using local allometries to estimate tree height show no over- or under-estimation bias when compared with biomass estimates using field measured heights. We evaluated five strategies to sample trees for height measurement, and found that sampling strategies that included measuring the heights of the ten largest diameter trees in a plot outperformed (in terms of resulting in local height-diameter models with low height prediction error) entirely random or diameter size-class stratified approaches. Our results indicate that even limited sampling of heights can be used to refine height-diameter allometries. We recommend aiming for a conservative threshold of sampling 50 trees per location for height measurement, and including the ten trees with the largest diameter in this sample. ; This paper is a product of the RAINFOR, AfriTRON and T-FORCES networks, for which we are indebted to the hundreds of institutions, field assistants and local communities across many countries that have supported and hosted fieldwork. The three networks have been supported by the Natural Environment Research Council (NERC) Urgency Grants and NERC Consortium Grants "AMAZONICA" (NE/F005806/1), "TROBIT" (NE/D005590/1) and "BIO-RED" (NE/N012542/1), a NERC New Investigators Grant, a European Research Council grant ("Tropical Forests in the Changing Earth System"), the Gordon and Betty Moore Foundation, the David and Lucile Packard Foundation, the European Union's Seventh Framework Programme (283080, "GEOCARBON"; 282664, "AMAZALERT"), the Royal Society and Gabon's National Parks Agency (ANPN). R.J.W.B. is funded by a NERC research fellowship (grant ref: NE/I021160/1). S.L.L. was supported by a Royal Society University Research Fellowship, ERC Advanced Grant and a Phillip Leverhulme Prize. O.L.P. is supported by an ERC Advanced Grant and a Royal Society Wolfson Research Merit Award. L.F.B. was supported by a NERC studentship, RGS-IBG Henrietta Hutton Grant and Royal Society Dudley Stamp Award. R.H. and M.C. were supported through the long-term research development project no. RVO 67985939 and a KBFSC research fellowship (2011, to R.H.). M. Svátek was funded by the Ministry of Education, Youth and Sports of the Czech Republic (grant number INGO II LG15051). We thank Georgia Pickavance for assistance with database curation, and Natacha Nssi Bengone, Sylvester Chenikan, Eric Chezeaux, Armandu Daniels, Jean-Louis Doucet, Kath Jeffery, Edi Mirmanto, Abel Monteagudo-Mendoza, Faustin Mpanya Lukasu, Reuben Nilus, Guido Pardo, Lourens Poorter, Sylvester Tan, Marisol Toledo, Armando Torres-Lezama, John Tshibamba Mukendi, Richard Tshombe, Geertje van der Heijden, Lee White, Hannsjoerg Woell and John Woods, Gabon's National Parks Agency (ANPN), the Forest Development Authority of Liberia and Wildlife Conservation Society-Democratic Republic of Congo for assistance with access to datasets. We thank an anonymous reviewer for constructive comments on this manuscript.

This is the final version. Available from the National Academy of Sciences via the DOI in this record. ; The input data and R code are available on ForestPlots (https://doi.org/10.5521/forestplots.net/2021_4). ; The responses of tropical forests to environmental change are critical uncertainties in predicting the future impacts of climate change. The positive phase of the 2015-2016 El Niño Southern Oscillation resulted in unprecedented heat and low precipitation in the tropics with substantial impacts on the global carbon cycle. The role of African tropical forests is uncertain as their responses to short-term drought and temperature anomalies have yet to be determined using on-the-ground measurements. African tropical forests may be particularly sensitive because they exist in relatively dry conditions compared with Amazonian or Asian forests, or they may be more resistant because of an abundance of drought-adapted species. Here, we report responses of structurally intact old-growth lowland tropical forests inventoried within the African Tropical Rainforest Observatory Network (AfriTRON). We use 100 long-term inventory plots from six countries each measured at least twice prior to and once following the 2015-2016 El Niño event. These plots experienced the highest temperatures and driest conditions on record. The record temperature did not significantly reduce carbon gains from tree growth or significantly increase carbon losses from tree mortality, but the record drought did significantly decrease net carbon uptake. Overall, the long-term biomass increase of these forests was reduced due to the El Niño event, but these plots remained a live biomass carbon sink (0.51 ± 0.40 Mg C ha-1 y-1) despite extreme environmental conditions. Our analyses, while limited to African tropical forests, suggest they may be more resistant to climatic extremes than Amazonian and Asian forests. ; Natural Environment Research Council (NERC) ; Natural Environment Research Council (NERC) ; European Research Council (ERC) ; The Royal Society ; Belgian Science Policy Office (BELSPO) ; Belgian Science Policy Office (BELSPO) ; Belgian Science Policy Office (BELSPO) ; Belgian Science Policy Office (BELSPO) ; Flemish Interuniversity Council VLIR-UOS ; Flemish Interuniversity Council VLIR-UOS ; Natural Environment Research Council (NERC) ; The Gordon and Betty Moore Foundation ; European Union ; Natural Environment Research Council (NERC) ; Natural Environment Research Council (NERC) ; Natural Environment Research Council (NERC) ; Gabon's National Parks Agency ; Leverhulme Trust ; The David and Lucile Packard Foundation ; CIFOR

This is the final version. Available on open access from Wiley via the DOI in this record ; The full author list is available in the online article at the DOI in this record ; Aim: Palms are an iconic, diverse and often abundant component of tropical ecosystems that provide many ecosystem services. Being monocots, tree palms are evolutionarily, morphologically and physiologically distinct from other trees, and these differences have important consequences for ecosystem services (e.g., carbon sequestration and storage) and in terms of responses to climate change. We quantified global patterns of tree palm relative abundance to help improve understanding of tropical forests and reduce uncertainty about these ecosystems under climate change. Location: Tropical and subtropical moist forests. Time period: Current. Major taxa studied: Palms (Arecaceae). Methods: We assembled a pantropical dataset of 2,548 forest plots (covering 1,191 ha) and quantified tree palm (i.e., ≥10 cm diameter at breast height) abundance relative to co-occurring non-palm trees. We compared the relative abundance of tree palms across biogeographical realms and tested for associations with palaeoclimate stability, current climate, edaphic conditions and metrics of forest structure. Results: On average, the relative abundance of tree palms was more than five times larger between Neotropical locations and other biogeographical realms. Tree palms were absent in most locations outside the Neotropics but present in >80% of Neotropical locations. The relative abundance of tree palms was more strongly associated with local conditions (e.g., higher mean annual precipitation, lower soil fertility, shallower water table and lower plot mean wood density) than metrics of long-term climate stability. Life-form diversity also influenced the patterns; palm assemblages outside the Neotropics comprise many non-tree (e.g., climbing) palms. Finally, we show that tree palms can influence estimates of above-ground biomass, but the magnitude and direction of the effect require additional work. Conclusions: Tree palms are not only quintessentially tropical, but they are also overwhelmingly Neotropical. Future work to understand the contributions of tree palms to biomass estimates and carbon cycling will be particularly crucial in Neotropical forests. ; Natur og Univers, Det Frie Forskningsråd ; European Union Horizon 2020 ; Brazilian National Research Council ; Natural Environment Research Council (NERC) ; Vetenskapsrådet ; Coordenação de Aperfeiçoamento de Pessoal de Nível Superior ; Fundação de Amparo à Pesquisa do Estado de São Paulo ; Villum Fonden

This is the final version. Available on open access from Wiley via the DOI in this record ; Most of the planet's diversity is concentrated in the tropics, which includes many regions undergoing rapid climate change. Yet, while climate-induced biodiversity changes are widely documented elsewhere, few studies have addressed this issue for lowland tropical ecosystems. Here we investigate whether the floristic and functional composition of intact lowland Amazonian forests have been changing by evaluating records from 106 long-term inventory plots spanning 30 years. We analyse three traits that have been hypothesized to respond to different environmental drivers (increase in moisture stress and atmospheric CO2 concentrations): maximum tree size, biogeographic water-deficit affiliation and wood density. Tree communities have become increasingly dominated by large-statured taxa, but to date there has been no detectable change in mean wood density or water deficit affiliation at the community level, despite most forest plots having experienced an intensification of the dry season. However, among newly recruited trees, dry-affiliated genera have become more abundant, while the mortality of wet-affiliated genera has increased in those plots where the dry season has intensified most. Thus, a slow shift to a more dry-affiliated Amazonia is underway, with changes in compositional dynamics (recruits and mortality) consistent with climate-change drivers, but yet to significantly impact whole-community composition. The Amazon observational record suggests that the increase in atmospheric CO2 is driving a shift within tree communities to large-statured species and that climate changes to date will impact forest composition, but long generation times of tropical trees mean that biodiversity change is lagging behind climate change. ; Support for RAINFOR has come from the Natural Environment Research Council (NERC) Urgency Grants and NERC Consortium Grants "AMAZONICA" (NE/F005806/1), "TROBIT" (NE/D005590/1) and "BIO‐RED" (NE/N012542/1), a European Research Council (ERC) grant (T‐FORCES, "Tropical Forests in the Changing Earth System"), the Gordon and Betty Moore Foundation, the European Union's Seventh Framework Programme (282664, "AMAZALERT") and the Royal Society (CH160091). OLP was supported by an ERC Advanced Grant and a Royal Society Wolfson Research Merit Award. KGD was supported by a Leverhulme Trust International Academic Fellowship. This paper is part of the PhD of AE‐M, which was funded by the ERC T‐FORCES grant. AE‐M is currently supported by T‐FORCES and the NERC project "TREMOR" (NE/N004655/1).

Tropical forests are global centres of biodiversity and carbon storage. Many tropical countries aspire to protect forest to fulfil biodiversity and climate mitigation policy targets, but the conservation strategies needed to achieve these two functions depend critically on the tropical forest tree diversity-carbon storage relationship. Assessing this relationship is challenging due to the scarcity of inventories where carbon stocks in aboveground biomass and species identifications have been simultaneously and robustly quantified. Here, we compile a unique pan-tropical dataset of 360 plots located in structurally intact old-growth closed-canopy forest, surveyed using standardised methods, allowing a multi-scale evaluation of diversity-carbon relationships in tropical forests. Diversity-carbon relationships among all plots at 1 ha scale across the tropics are absent, and within continents are either weak (Asia) or absent (Amazonia, Africa). A weak positive relationship is detectable within 1 ha plots, indicating that diversity effects in tropical forests may be scale dependent. The absence of clear diversity-carbon relationships at scales relevant to conservation planning means that carbon-centred conservation strategies will inevitably miss many high diversity ecosystems. As tropical forests can have any combination of tree diversity and carbon stocks both require explicit consideration when optimising policies to manage tropical carbon and biodiversity. ; This paper is a product of the RAINFOR, AfriTRON and T-FORCES networks, for which we are hugely indebted to hundreds of institutions, field assistants and local communities across many countries that have hosted fieldwork. The three networks have been supported by a European Research Council (ERC) grant ("T-FORCES" - Tropical Forests in the Changing Earth System), the Gordon and Betty Moore Foundation, the David and Lucile Packard Foundation, the European Union's Seventh Framework Programme (283080, 'GEOCARBON'; 282664, 'AMAZALERT'), and Natural Environment Research Council (NERC) Urgency Grants and NERC Consortium Grants 'AMAZONICA' (NE/F005806/1) and 'TROBIT' (NE/D005590/1), 'BIO-RED' (NE/N012542/1) and a NERC New Investigators Grant, the Royal Society, the Centre for International Forestry (CIFOR) and Gabon's National Parks Agency (ANPN). Additional data were included from the Tropical Ecology Assessment and Monitoring (TEAM) Network, a collaboration between Conservation International, the Missouri Botanical Garden, the Smithsonian Institution and the Wildlife Conservation Society, and partly funded by these institutions, the Gordon and Betty Moore Foundation, and other donors. J.T. was supported by a NERC PhD Studentship with CASE sponsorship from UNEP-WCMC. R.J.W.B. is funded by a NERC research fellowship (grant ref: NE/I021160/1). S.L.L. was supported by a Royal Society University Research Fellowship, ERC Advanced Grant (T-FORCES) and a Phillip Leverhulme Prize. O.L.P. is supported by an ERC Advanced Grant (T-FORCES) and a Royal Society Wolfson Research Merit Award. L.F.B. was supported by a NERC studentship and RGS-IBG Henrietta Hutton Grant. We thank the National Council for Science and Technology Development of Brazil (CNPq) for support to Project Cerrado/Amazonia Transition (PELD/403725/2012-7), Project Phytogeography of Amazonia/Cerrado Transition (CNPq/PPBio/457602/2012-0) and Productivity Grant to B.S.M and B.H.M-J. Funding for plots in the Udzungwa Mountains (Tanzania) was obtained from the Leverhulme Trust under the Valuing the Arc project. We thank the ANPN (Gabon), WCS-Congo and WCS-DR Congo, Marien Ngouabi University and the University of Kisangani for logistical support in Africa, and the Tropenbos Kalimantan project (ITCI plots) and WWF (KUB plots) for providing data from Asia. This study is contribution number 706 to the Technical Series (TS) of the BDFFP – (INPA-STRI). For assistance with access to datasets we thank Adriana Prieto, Agustín Rudas, Alejandro Araujo-Murakami, Alexander G. Parada Gutierrez, Anand Roopsind, Atila Alves de Oliveira, Claudinei Oliveira dos Santos, C. E. Timothy Paine, David Neill, Eliana Jimenez-Rojas, Freddy Ramirez Arevalo, Hannsjoerg Woell, Iêda Leão do Amaral, Irina Mendoza Polo, Isau Huamantupa-Chuquimaco, Julien Engel, Kathryn Jeffery, Luzmila Arroyo, Michael D. Swaine, Nallaret Davila Cardozo, Natalino Silva, Nigel C. A. Pitman, Niro Higuchi, Raquel Thomas, Renske van Ek, Richard Condit, Rodolfo Vasquez Martinez, Timothy J. Killeen, Walter A. Palacios, Wendeson Castro. We thank Georgina Mace and Jon Lloyd for comments on the manuscript. We thank our deceased colleagues, Samuel Almeida, Kwaku Duah, Alwyn Gentry, and Sandra Patiño, for their invaluable contributions to this work and our wider understanding of tropical forest ecology.