Abstract. Forest ecosystem models based on heuristic water stress functions poorly predict tropical forest response to drought because they do not capture the diversity of hydraulic traits (including variation in tree size) observed in tropical forests. We developed a Richards’ equation-based model of plant hydraulics in which all parameters of its constitutive equations are biologically-interpretable and measureable plant hydraulic traits (e.g. turgor loss point πtlp, bulk elastic modulus ε, hydraulic capacitance Cft, xylem hydraulic conductivity ks,max, water potential at 50 % loss of conductivity for both xylem (P50,x) and stomata (P50,gs), and the leaf:sapwood area ratio Al:As). We embedded this plant hydraulics model within a forest simulator (TFS) that modeled individual tree light environments and their upper boundary condition (transpiration) as well as provided a means for parameterizing individual variation in hydraulic traits. We synthesized literature and existing databases to parameterize all hydraulic traits as a function of stem and leaf traits wood density (WD), leaf mass per area (LMA) and photosynthetic capacity (Amax) and evaluated the coupled model’s (TFS-Hydro) predictions against diurnal and seasonal variability in stem and leaf water potential as well as stand-scaled sap flux. Our hydraulic trait synthesis revealed coordination among leaf and xylem hydraulic traits and statistically significant relationships of most hydraulic traits with more easily measured plant traits. Using the most informative empirical trait-trait relationships derived from this synthesis, the TFS-Hydro model parameterization is capable of representing patterns of coordination and trade-offs in hydraulic traits. TFS-Hydro successfully captured individual variation in leaf and stem water potential due to increasing tree size and light environment, with model representation of hydraulic architecture and plant traits exerting primary and secondary controls, respectively, on the fidelity of model predictions. The plant hydraulics model made substantial improvements to simulations of total ecosystem transpiration under control conditions, but the absence of a vertically stratified soil hydrology model precluded improvements to the simulation of drought response. Remaining uncertainties and limitations of the trait paradigm for plant hydraulics modeling are highlighted.
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Abstract. Accurately predicting the response of Amazonia to climate change is important for predicting changes across the globe. However, changes in multiple climatic factors simultaneously may result in complex non-linear responses, which are difficult to predict using vegetation models. Using leaf and canopy scale observations, this study evaluated the capability of five vegetation models (CLM3.5, ED2, JULES, SiB3, and SPA) to simulate the responses of canopy and leaf scale productivity to changes in temperature and drought in an Amazonian forest. The models did not agree as to whether gross primary productivity (GPP) was more sensitive to changes in temperature or precipitation. There was greater model–data consistency in the response of net ecosystem exchange to changes in temperature, than in the response to temperature of leaf area index (LAI), net photosynthesis (An) and stomatal conductance (gs). Modelled canopy scale fluxes are calculated by scaling leaf scale fluxes to LAI, and therefore in this study similarities in modelled ecosystem scale responses to drought and temperature were the result of inconsistent leaf scale and LAI responses among models. Across the models, the response of an to temperature was more closely linked to stomatal behaviour than biochemical processes. Consequently all the models predicted that GPP would be higher if tropical forests were 5 °C colder, closer to the model optima for gs. There was however no model consistency in the response of the An–gs relationship when temperature changes and drought were introduced simultaneously. The inconsistencies in the An–gs relationships amongst models were caused by to non-linear model responses induced by simultaneous drought and temperature change. To improve the reliability of simulations of the response of Amazonian rainforest to climate change the mechanistic underpinnings of vegetation models need more complete validation to improve accuracy and consistency in the scaling of processes from leaf to canopy.
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Abstract. Accurately representing the response of ecosystems to environmental change in land surface models (LSM) is crucial to making accurate predictions of future climate. Many LSMs do not correctly capture plant respiration and growth fluxes, particularly in response to extreme climatic events. This is in part due to the unrealistic assumption that total plant carbon expenditure (PCE) is always equal to gross carbon accumulation by photosynthesis. We present and evaluate a simple model of labile carbon storage and utilisation (SUGAR), designed to be integrated into an LSM, that allows simulated plant respiration and growth to vary independently of photosynthesis. SUGAR buffers simulated PCE against seasonal variation in photosynthesis, producing more constant (less variable) predictions of plant growth and respiration relative to an LSM that does not represent labile carbon storage. This allows the model to more accurately capture observed carbon fluxes at a large-scale drought experiment in a tropical moist forest in the Amazon, relative to the Joint UK Land Environment Simulator LSM (JULES). SUGAR is designed to improve the representation of carbon storage in LSMs and provides a simple framework that allows new processes to be integrated as the empirical understanding of carbon storage in plants improves. The study highlights the need for future research into carbon storage and allocation in plants, particularly in response to extreme climate events such as drought.
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AbstractTropical forests face increasing climate risk1,2, yet our ability to predict their response to climate change is limited by poor understanding of their resistance to water stress. Although xylem embolism resistance thresholds (for example, $$\varPsi $$
. Ψ
. 50) and hydraulic safety margins (for example, HSM50) are important predictors of drought-induced mortality risk3–5, little is known about how these vary across Earth’s largest tropical forest. Here, we present a pan-Amazon, fully standardized hydraulic traits dataset and use it to assess regional variation in drought sensitivity and hydraulic trait ability to predict species distributions and long-term forest biomass accumulation. Parameters $$\varPsi $$
. Ψ
. 50 and HSM50 vary markedly across the Amazon and are related to average long-term rainfall characteristics. Both $$\varPsi $$
. Ψ
. 50 and HSM50 influence the biogeographical distribution of Amazon tree species. However, HSM50 was the only significant predictor of observed decadal-scale changes in forest biomass. Old-growth forests with wide HSM50 are gaining more biomass than are low HSM50 forests. We propose that this may be associated with a growth–mortality trade-off whereby trees in forests consisting of fast-growing species take greater hydraulic risks and face greater mortality risk. Moreover, in regions of more pronounced climatic change, we find evidence that forests are losing biomass, suggesting that species in these regions may be operating beyond their hydraulic limits. Continued climate change is likely to further reduce HSM50 in the Amazon6,7, with strong implications for the Amazon carbon sink.

Future increases in drought severity and frequency are predicted to have substantial impacts on plant function and survival. However, there is considerable uncertainty concerning what drought adjustment is and whether plants can adjust to sustained drought. This review focuses on woody plants and synthesises the evidence for drought adjustment in a selection of key above-ground and below-ground plant traits. We assess whether evaluating the drought adjustment of single traits, or selections of traits that operate on the same plant functional axis (e.g. photosynthetic traits) is sufficient, or whether a multi-trait approach, integrating across multiple axes, is required. We conclude that studies on drought adjustments in woody plants might overestimate the capacity for adjustment to drier environments if spatial studies along gradients are used, without complementary experimental approaches. We provide evidence that drought adjustment is common in above-ground and below-ground traits; however, whether this is adaptive and sufficient to respond to future droughts remains uncertain for most species. To address this uncertainty, we must move towards studying trait integration within and across multiple axes of plant function (e.g. above-ground and below-ground) to gain a holistic view of drought adjustments at the whole-plant scale and how these influence plant survival.

While existing moratoria in Indonesia and Malaysia should preclude continued large-scale expansion of palm oil production into new areas of South-East Asian tropical peatland, existing plantations in the region remain a globally significant source of atmospheric carbon due to drainage driven decomposition of peatland soils. Previous studies have made clear the direct link between drainage depth and peat carbon decomposition and significant reductions in the emission rate of CO2 can be made by raising water tables nearer to the soil surface. However, the impact of such changes on palm fruit yield is not well understood and will be a critical consideration for plantation managers. Here we take advantage of very high frequency, long-term monitoring of canopy-scale carbon exchange at a mature oil palm plantation in Malaysian Borneo to investigate the relationship between drainage level and photosynthetic uptake and consider the confounding effects of light quality and atmospheric vapour pressure deficit. Canopy modelling from our dataset demonstrated that palms were exerting significantly greater stomatal control at deeper water table depths (WTD) and the optimum WTD for photosynthesis was found to be between 0.3 and 0.4 m below the soil surface. Raising WTD to this level, from the industry typical drainage level of 0.6 m, could increase photosynthetic uptake by 3.6 % and reduce soil surface emission of CO2 by 11 %. Our study site further showed that despite being poorly drained compared to other planting blocks at the same plantation, monthly fruit bunch yield was, on average, 14 % greater. While these results are encouraging, and at least suggest that raising WTD closer to the soil surface to reduce emissions is unlikely to produce significant yield penalties, our results are limited to a single study site and more work is urgently needed to confirm these results at other plantations.

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'.

Atmospheric conditions are expected to become warmer and drier in the future, but little is known about how evaporative demand influences forest structure and function independently from soil moisture availability, and how fast-response variables (such as canopy water potential and stomatal conductance) may mediate longer-term changes in forest structure and function in response to climate change. We used two tropical rainforest sites with different temperatures and vapour pressure deficits (VPD), but nonlimiting soil water supply, to assess the impact of evaporative demand on ecophysiological function and forest structure. Common species between sites allowed us to test the extent to which species composition, relative abundance and intraspecific variability contributed to site-level differences. The highest VPD site had lower midday canopy water potentials, canopy conductance (gc ), annual transpiration, forest stature, and biomass, while the transpiration rate was less sensitive to changes in VPD; it also had different height-diameter allometry (accounting for 51% of the difference in biomass between sites) and higher plot-level wood density. Our findings suggest that increases in VPD, even in the absence of soil water limitation, influence fast-response variables, such as canopy water potentials and gc , potentially leading to longer-term changes in forest stature resulting in reductions in biomass.

Tropical forests take up more carbon (C) from the atmosphere per annum by photosynthesis than any other type of vegetation. Phosphorus (P) limitations to C uptake are paramount for tropical and subtropical forests around the globe. Yet the generality of photosynthesis-P relationships underlying these limitations are in question, and hence are not represented well in terrestrial biosphere models. Here we demonstrate the dependence of photosynthesis and underlying processes on both leaf N and P concentrations. The regulation of photosynthetic capacity by P was similar across four continents. Implementing P constraints in the ORCHIDEE-CNP model, gross photosynthesis was reduced by 36% across the tropics and subtropics relative to traditional N constraints and unlimiting leaf P. Our results provide a quantitative relationship for the P dependence for photosynthesis for the front-end of global terrestrial C models that is consistent with canopy leaf measurements.

Fine-scale topographic-edaphic gradients are common in tropical forests and drive species spatial turnover and marked changes in forest structure and function. We evaluate how hydraulic traits of tropical tree species relate to vertical and horizontal spatial niche specialization along such a gradient. Along a topographic-edaphic gradient with uniform climate in Borneo, we measured six key hydraulic traits in 156 individuals of differing heights in 13 species of Dipterocarpaceae. We investigated how hydraulic traits relate to habitat, tree height and their interaction on this gradient. Embolism resistance increased in trees on sandy soils but did not vary with tree height. By contrast, water transport capacity increased on sandier soils and with increasing tree height. Habitat and height only interact for hydraulic efficiency, with slope for height changing from positive to negative from the clay-rich to the sandier soil. Habitat type influenced trait-trait relationships for all traits except wood density. Our data reveal that variation in the hydraulic traits of dipterocarps is driven by a combination of topographic-edaphic conditions, tree height and taxonomic identity. Our work indicates that hydraulic traits play a significant role in shaping forest structure across topographic-edaphic and vertical gradients and may contribute to niche specialization among dipterocarp species.

. There is high potential for ecosystem restoration across tropical savannah-dominated regions, but the benefits that could be gained from this restoration are rarely assessed. This study focuses on the Brazilian Cerrado, a highly species-rich savannah-dominated region, as an exemplar to review potential restoration benefits using three metrics: net biomass gains, plant species richness and ability to connect restored and native vegetation. Localized estimates of the most appropriate restoration vegetation type (grassland, savannah, woodland/forest) for pasturelands are produced. Carbon sequestration potential is significant for savannah and woodland/forest restoration in the seasonally dry tropics (net biomass gains of 58.2 ± 37.7 and 130.0 ± 69.4 Mg ha
. −1
. ). Modelled restoration species richness gains were highest in the central and south-east of the Cerrado for savannahs and grasslands, and in the west and north-west for woodlands/forests. The potential to initiate restoration projects across the whole of the Cerrado is high and four hotspot areas are identified. We demonstrate that landscape restoration across all vegetation types within heterogeneous tropical savannah-dominated regions can maximize biodiversity and carbon gains. However, conservation of existing vegetation is essential to minimizing the cost and improving the chances of restoration success.
.
. This article is part of the theme issue ‘Understanding forest landscape restoration: reinforcing scientific foundations for the UN Decade on Ecosystem Restoration’.

Native vegetation across the Brazilian Cerrado is highly heterogeneous and biodiverse and provides important ecosystem services, including carbon and water balance regulation, however, land-use changes have been extensive. Conservation and restoration of native vegetation is essential and could be facilitated by detailed landcover maps. Here, across a large case study region in Goiás State, Brazil (1.1 Mha), we produced physiognomy level maps of native vegetation (n = 8) and other landcover types (n = 5). Seven different classification schemes using different combinations of input satellite imagery were used, with a Random Forest classifier and 2-stage approach implemented within Google Earth Engine. Overall classification accuracies ranged from 88.6-92.6% for native and non-native vegetation at the formation level (stage-1), and 70.7-77.9% for native vegetation at the physiognomy level (stage-2), across the seven different classifications schemes. The differences in classification accuracy resulting from varying the input imagery combination and quality control procedures used were small. However, a combination of seasonal Sentinel-1 (C-band synthetic aperture radar) and Sentinel-2 (surface reflectance) imagery resulted in the most accurate classification at a spatial resolution of 20 m. Classification accuracies when using Landsat-8 imagery were marginally lower, but still reasonable. Quality control procedures that account for vegetation burning when selecting vegetation reference data may also improve classification accuracy for some native vegetation types. Detailed landcover maps, produced using freely available satellite imagery and upscalable techniques, will be important tools for understanding vegetation functioning at the landscape scale and for implementing restoration projects.

Terrestrial ecosystems absorb over one-quarter of anthropogenic carbon dioxide (CO2) released into the atmosphere each year. Heterotrophic soil microbial respiration, associated with the decomposition of organic matter, contributes to approximately half of the CO2 released from the terrestrial biosphere to the atmosphere per year. However, with increasing global temperatures, there is the potential for soil microbial respiration to increase, resulting in a substantial release of CO2 into the atmosphere, therefore contributing extensively to the positive land C-climate feedback and accelerating climate change. In the short-term (days-years), heterotrophic soil microbial respiration is strongly and positively related to temperature. In the long-term (years-decades), however, the positive response of soil microbial respiration to warming declines, which could be caused by direct (acclimation, evolution and species sorting) or indirect (e.g. substrate availability, moisture) effects of warming. The primary focus on whole soil microbial community responses has made it difficult to identify potential mechanisms involved in controlling long-term warming responses, thus the response of respiration to warming in the long term remains controversial. To address this uncertainty, it is necessary to study individual species and gradually more diverse and complex decomposer communities. To this end, the respiratory thermal response of wood decay basidiomycetes, the dominant decomposers of wood, were investigated. Respiration rates of individual species and two- and three-species assemblages of basidiomycetes and semi-natural wood decay communities, decomposing beech wood (Fagus sylvatica), were measured during a 90-day cooling approach. In addition, a warming approach was applied to the two- and three-species assemblages and semi-natural wood decay communities. The direction of any thermal response (decreased temperature sensitivity of respiration (compensatory), increased temperature sensitivity of respiration (enhancing), and no change in the temperature sensitivity of respiration (no response)) was determined. To increase our understanding of the respiratory thermal responses, the growth response of the basidiomycete species to temperature was also measured.

Following cooling, individual species of basidiomycetes showed an overall enhancing response, with no compensatory responses identified. Two-species assemblages and the three-species assemblage showed no thermal responses overall to cooling, but with some evidence of compensatory and enhancing responses. Semi-natural wood decomposing communities showed no thermal response overall to cooling, with more enhancing than compensatory responses detected. With warming, two-species assemblages showed no thermal response overall, with more compensatory than enhancing responses detected, whereas the three-species assemblage that was dominated by one species towards the end of incubation demonstrated a compensatory response overall. The compensatory responses from the two- and three-species assemblages were likely caused by exceeding the optimum temperature for growth for some species or by the differences in the progression of the species interactions. Therefore, there was little evidence of compensatory responses that would decrease the temperature sensitivity of respiration. Semi-natural wood decay communities showed no thermal response overall to warming, but some enhancing responses were identified.

The findings showed that the temperature sensitivity of wood decomposition was increased when basidiomycetes were grown alone, however, this was reduced during competitive interspecific interactions between basidiomycetes and species in wood decomposing communities. With increasing global temperatures, individual species, growing alone during the early colonisation and decomposition of wood and in decay columns in stable wood communities, may increase their respiration, but simple communities of interacting basidiomycetes and more diverse wood decomposing communities in natural systems, will more often cause no change in the temperature sensitivity of respiration overall. However, the species present in communities will most likely determine the direction and strength of respiratory responses to temperature, and thus the overall temperature sensitivity of respiration of wood decay communities. With limited evidence for compensatory responses and more evidence of enhancing responses detected, it is, thus, considered unlikely that the temperature sensitivity of wood decomposition will decline as global temperatures rise. Therefore, there remains the potential for a positive feedback to climate change through increased wood decomposition with warming.

Future climate change predictions for tropical forests highlight increased frequency and intensity of extreme drought events. However, it remains unclear whether large and small trees have differential strategies to tolerate drought due to the different niches they occupy. The future of tropical forests is ultimately dependent on the capacity of small trees (

Background: the Cerrado of central Brazil—the world’s largest Neotropical savanna – is comprised of a mosaic of highly heterogeneous vegetation growing on an extremely diverse geologic and geomorphologic background. Geomorphic processes under stable tectonic and climatic conditions facilitated the development of diverse edaphic properties, which interact with disturbance events to form unique vegetation types. Scope: in this review, we detail how the geophysical environment affects soil formation and evaluate the mechanisms through which edaphic conditions control vegetation structure, floristic diversity and functional diversity. Conclusion: the influence of geomorphic processes on edaphic properties has a marked impact on the ecology and evolution of plant communities. Species exhibit morphological and physiological adaptations that optimise their successful establishment in particular soil conditions. Furthermore, fire disturbance alters these soil-vegetation associations further regulating the structural nature of these communities. Therefore, we propose an integrative view where edaphic, chemical and physical properties act as modulators of vegetation stands, and these conditions interact with the fire regime. The knowledge of plant edaphic niches, their functional traits related to resource acquisition and use, as well as the interaction of edaphic properties and disturbance regimes is paramount to research planning, conservation, and successful restoration of the full diversity of Cerrado vegetation types.

Seed markets are vital to scaling up ecosystem restoration in the Brazilian Cerrado, home of the world’s most species-rich grasslands and savannas. We compiled lists of species traded by four major Cerrado seed supply systems to investigate the representativeness of the species currently available for seed-based restoration. We also identified whether dominant ground-layer species are being sourced for seed production. Seeds from 263 Cerrado species can be purchased for restoration, of which 68% are trees, particularly legumes (24%). 63% of the traded species were found in only one seed supply system. The five most dominant graminoids of the Cerrado ground layer were available for sale, but two additional species uncommon in old-growth areas represented 44% of the sales of a key seed trader in Central Brazil. The expansion of Cerrado seed supply systems should be supported to further increase the number of species on the market. Sourcing seeds from a diversity of herbaceous species is central to facilitating the restoration of species-rich grasslands and savannas in the Cerrado. Recovering the diversity and functioning of old-growth open ecosystems through seeds will depend on increasing the supply and demand for species typical of Cerrado’s ground layer.

Plant transpiration links physiological responses of vegetation to water supply and demand with hydrological, energy, and carbon budgets at the land-atmosphere interface. However, despite being the main land evaporative flux at the global scale, transpiration and its response to environmental drivers are currently not well constrained by observations. Here we introduce the first global compilation of whole-plant transpiration data from sap flow measurements (SAPFLUXNET, https://sapfluxnet.creaf.cat/, last access: 8 June 2021). We harmonized and quality-controlled individual datasets supplied by contributors worldwide in a semi-automatic data workflow implemented in the R programming language. Datasets include sub-daily time series of sap flow and hydrometeorological drivers for one or more growing seasons, as well as metadata on the stand characteristics, plant attributes, and technical details of the measurements. SAPFLUXNET contains 202 globally distributed datasets with sap flow time series for 2714 plants, mostly trees, of 174 species. SAPFLUXNET has a broad bioclimatic coverage, with woodland/shrubland and temperate forest biomes especially well represented (80% of the datasets). The measurements cover a wide variety of stand structural characteristics and plant sizes. The datasets encompass the period between 1995 and 2018, with 50% of the datasets being at least 3 years long. Accompanying radiation and vapour pressure deficit data are available for most of the datasets, while on-site soil water content is available for 56% of the datasets. Many datasets contain data for species that make up 90% or more of the total stand basal area, allowing the estimation of stand transpiration in diverse ecological settings. SAPFLUXNET adds to existing plant trait datasets, ecosystem flux networks, and remote sensing products to help increase our understanding of plant water use, plant responses to drought, and ecohydrological processes. SAPFLUXNET version 0.1.5 is freely available from the Zenodo repository (10.5281/zenodo.3971689; Poyatos et al. 2020a). The "sapfluxnetr"R package-designed to access, visualize, and process SAPFLUXNET data-is available from CRAN.

Species loss leads to changes in ecosystem function and services, impacting human well‐being. Although biodiversity restoration is pivotal to circumvent this situation, the techniques for restoring old‐growth savannas are still limited and the restoration outcomes remain unpredictable. Here, we use a trait‐based approach to understand the functional outcomes of ecological restoration via direct seeding in a Brazilian savanna (cerrado, hereafter neotropical savanna). We compared the functional composition from woody and non‐woody component, total biomass, and biomass allocation of a restored relative to a degraded savanna (abandoned pasture) dominated by exotic grasses and a well‐preserved old‐growth native savanna. We found that the functional composition of restored communities was very similar to those dominated by exotic grasses, both characterized by a greater dominance of species with acquisitive traits, higher above‐ground biomass, and lower investment in root biomass. In contrast, the native vegetation exhibited a dominance of conservative traits and higher investment in belowground rather than aboveground biomass. Even though the acquisitive traits in the restored savanna allow a fast aboveground biomass accumulation and soil cover, the lower belowground biomass investment in the restored savanna may limit its resistance and resilience to droughts and fires. Our findings suggest that restoration efforts in neotropical savanna should focus on fostering the establishment of slow‐growing species to recover the ecosystem properties provided by the high biodiversity in neotropical savannas.

Abstract. Drought is predicted to increase in the future due to climate change, bringing with it myriad impacts on ecosystems. Plants respond to drier soils by reducing stomatal conductance in order to conserve water and avoid hydraulic damage. Despite the importance of plant drought responses for the global carbon cycle and local and regional climate feedbacks, land surface models are unable to capture observed plant responses to soil moisture stress. We assessed the impact of soil moisture stress on simulated gross primary productivity (GPP) and latent energy flux (LE) in the Joint UK Land Environment Simulator (JULES) vn4.9 on seasonal and annual timescales and evaluated 10 different representations of soil moisture stress in the model. For the default configuration, GPP was more realistic in temperate biome sites than in the tropics or high-latitude (cold-region) sites, while LE was best simulated in temperate and high-latitude (cold) sites. Errors that were not due to soil moisture stress, possibly linked to phenology, contributed to model biases for GPP in tropical savanna and deciduous forest sites. We found that three alternative approaches to calculating soil moisture stress produced more realistic results than the default parameterization for most biomes and climates. All of these involved increasing the number of soil layers from 4 to 14 and the soil depth from 3.0 to 10.8 m. In addition, we found improvements when soil matric potential replaced volumetric water content in the stress equation (the “soil14_psi” experiments), when the critical threshold value for inducing soil moisture stress was reduced (“soil14_p0”), and when plants were able to access soil moisture in deeper soil layers (“soil14_dr*2”). For LE, the biases were highest in the default configuration in temperate mixed forests, with overestimation occurring during most of the year. At these sites, reducing soil moisture stress (with the new parameterizations mentioned above) increased LE and increased model biases but improved the simulated seasonal cycle and brought the monthly variance closer to the measured variance of LE. Further evaluation of the reason for the high bias in LE at many of the sites would enable improvements in both carbon and energy fluxes with new parameterizations for soil moisture stress. Increasing the soil depth and plant access to deep soil moisture improved many aspects of the simulations, and we recommend these settings in future work using JULES or as a general way to improve land surface carbon and water fluxes in other models. In addition, using soil matric potential presents the opportunity to include plant functional type-specific parameters to further improve modeled fluxes.

Restoration techniques tailored to grasslands are needed to improve the effectiveness of restoration in tropical landscapes. In this study, we investigated the joint effects of plant–soil legacies and soil inocula in native and invaded Cerrado grasslands to evaluate whether different microbial origins affect plant–soil feedbacks and the likelihood of restoration. Using two grass species, we measured aboveground biomass, and several plant traits over two growth cycles. Species responded differently to inocula and legacies. The legacy of the invasive Urochloa eminii and invaded soil inocula positively affected mycorrhizal colonization. The legacy of Diectomis fastigiata, a commonly used species in Cerrado restoration, resulted in a negative self‐feedback potentially limiting its effectiveness for restoration. The success of the invasive species was in part due to its broad ecological niche and its ability to cope with a broad range of soil conditions. Our research suggests soil inocula and legacies could be used to aid restoration efforts in the tropics, allowing restoration practitioners to stimulate the growth of species targeting functional traits for a given ecosystem.

A large portion of the terrestrial vegetation carbon stock is stored in the above-ground biomass (AGB) of tropical forests, but the exact amount remains uncertain, partly owing to the lack of measurements. To date, accessible peer-reviewed data are available for just 10 large tropical trees in the Amazon that have been harvested and directly measured entirely via weighing. Here, we harvested four large tropical rainforest trees (stem diameter: 0.6-1.2 m, height: 30-46 m, AGB: 3960-18 584 kg) in intact old-growth forest in East Amazonia, and measured above-ground green mass, moisture content and woody tissue density. We first present rare ecological insights provided by these data, including unsystematic intra-tree variations in density, with both height and radius. We also found the majority of AGB was usually found in the crown, but varied from 42 to 62%. We then compare non-destructive approaches for estimating the AGB of these trees, using both classical allometry and new lidar-based methods. Terrestrial lidar point clouds were collected pre-harvest, on which we fitted cylinders to model woody structure, enabling retrieval of volume-derived AGB. Estimates from this approach were more accurate than allometric counterparts (mean tree-scale relative error: 3% versus 15%), and error decreased when up-scaling to the cumulative AGB of the four trees (1% versus 15%). Furthermore, while allometric error increased fourfold with tree size over the diameter range, lidar error remained constant. This suggests error in these lidar-derived estimates is random and additive. Were these results transferable across forest scenes, terrestrial lidar methods would reduce uncertainty in stand-scale AGB estimates, and therefore advance our understanding of the role of tropical forests in the global carbon cycle.

Plant traits are increasingly being used to improve prediction of plant function, including plant demography. However, the capability of plant traits to predict demographic rates remains uncertain, particularly in the context of trees experiencing a changing climate. Here we present data combining 17 plant traits associated with plant structure, metabolism and hydraulic status, with measurements of long-term mean, maximum and relative growth rates for 176 trees from the world's longest running tropical forest drought experiment. We demonstrate that plant traits can predict mean annual tree growth rates with moderate explanatory power. However, only combinations of traits associated more directly with plant functional processes, rather than more commonly employed traits like wood density or leaf mass per area, yield the power to predict growth. Critically, we observe a shift from growth being controlled by traits related to carbon cycling (assimilation and respiration) in well-watered trees, to traits relating to plant hydraulic stress in drought-stressed trees. We also demonstrate that even with a very comprehensive set of plant traits and growth data on large numbers of tropical trees, considerable uncertainty remains in directly interpreting the mechanisms through which traits influence performance in tropical forests.

Land surface models (LSM) represent a significant source of uncertainty in predictions of future climate. Many LSMs are unable to account for differences between plant carbon assimilation through photosynthesis, and plant carbon expenditure through autotrophic respiration and growth, as they do not comprehensively represent labile non-structural carbohydrate (NSC) stores that allow asynchrony between assimilation and expenditure to occur. This limits the ability of LSMs to accurately capture seasonal and inter-annual variation of ecosystem carbon fluxes in particular during periods of environmental stress.
This thesis discusses the current empirical understanding of NSC, and examines previous representations of NSC storage and utilisation within LSMs. A simple model of NSC designed to decouple plant carbon assimilation and expenditure in LSMs and improve predictions of ecosystem carbon fluxes, is presented. The model is tested at three scales and under varying climatic conditions. First, in simulations across the Amazon rainforest, the model decouples respiration and growth from photosynthesis, resulting in shifts in the seasonal cycle of total carbon expenditure. Then at a tropical drought experiment in Caxiuan˜ a, Brazil, the model allows more accurate predictions of carbon fluxes relative to a LSM that does not represent NSC. Finally, at a global scale, the model is used to highlight the potential importance of NSC in predictions of global terrestrial carbon uptake. The thesis concludes by outlining possible developments for future work.

Tropical forests possess exceptional levels of tree species richness but explaining this diversity has presented a long existing challenge. Habitat niche partitioning provides a hypothesis for species co-existence, whereby species avoid competitive exclusion by partitioning demands on multiple resources within an environment. However, limited understanding concerning how tree function is influenced by multiple environmental variables has limited the support for this hypothesis. This knowledge gap also limits our ability to predict how tropical forest tree communities will respond to environmental change, given multiple dimensions of a species’ niche are likely to be affected.
In this thesis, I investigate the role of niche partitioning in supporting co-existence of species and the turnover of species across edaphic gradients, as well as how long-term changes to the environment from selective logging and drought affect niche space of tropical tree species. I use species distribution models and measurements of leaf physiological traits to determine the key dimensions of tree species’ niches in primary forests.
In chapter 2 I demonstrate niche partitioning is strong within tropical forests with at least 60-86% of abundant species occupying their own unique niche. Species partition a wide range of abiotic environments, including soil nutrient, topographic and light environments, with greater environmental heterogeneity enhancing the scope for niche partitioning. Building on this, in chapter 3 I find that variation in nutrient availability explains more variation in leaf physiology and habitat preferences than light availability of species from the Dipterocarpaceae family that dominates South-East Asian forests. This highlights the importance of edaphic environments in structuring tropical forest communities. I also find different leaf nutrients are related to photosynthetic capacity in different forest types, revealing that multiple different nutrients may limit productivity and affect species distributions in tropical forests.
Many tropical forest tree species are highly specialised with limited ability to adjust their traits between environments, underlining their potential vulnerability to environmental change. In chapter 4 I show seedlings from selectively logged Bornean forests have different community weighted mean trait values, with greater belowground investment in logged forests. These adaptations are sufficient to overcome soil stress and to maintain foliar nutrient concentrations. However, I show seedlings of species that dominate old-growth forests are less able to adapt their traits and experience elevated mortality rates in logged forests. I attribute this to greater soil nutrient limitation as they are unable to maintain leaf nutrient concentrations. Selective logging will therefore likely drive shifts in species composition towards greater dominance of earlier-successional species that have traits capable of surviving in disturbed environments. This could result in local-scale reductions in species diversity and functional diversity, which could reduce long-term resilience to environmental change. In contrast, in Chapter 5 I demonstrate small trees in Amazonian forests are able to respond to changes in their environment following long-term drought conditions. Following mortality of large canopy trees, small trees can respond to increased light availability even under reduced water availability by adjusting resource allocation and by increasing nutrient use efficiency. Despite evidence of resilience to long-term drought conditions, hyper-dominant species show a greater capacity to respond, which may further enhance the dominance of these species under future climates.
In conclusion my results highlight the importance of habitat niche partitioning in structuring tropical forest tree communities and identify key environmental variables that determine species distribution and tree function. My results have important implications for the conservation and restoration of tropical forests under environmental change. Avoidance of environmental homogenisation and changes to as few environmental conditions as possible is likely to be important in maintaining high species diversity in tropical forests and to avoid increased dominance by few generalist species. Many current conservation and restoration projects focus on recovering vegetation, but my research highlights the additional need to maintain and restore soil environments, especially for the long-term persistence of highly specialist species.

Comparisons among methods are essential to validate plant traits measured across studies. However, a rigorous analysis is a complex task that needs to take into account not only the principle of the method and its correct use, but also inherent intraspecific trait variability, something we feel is not fully considered by Sergent et al. (2020). They compared the Bench dehydration, MicroCT, and Pneumatic methods using three long-vesseled species and found divergence among these methods. As a key finding, Sergent and colleagues reported unreliable estimates of Ψ50 for Olea europaea when using the Pneumatic method in a such long-vesseled species. Here, we tested this finding by measuring independently vulnerability curves for O. europaea. Our results reinforce the viability of the Pneumatic method to estimate embolism vulnerability in long-vesseled species, as already found by others. Briefly, we also discuss important procedures when using the Pneumatic method and encourage further experiments, as the only way to know better the limitations of available methods and improve our understanding about plant water relations.

Abstract


Tropical tree species span a range of life‐history strategies within a fast–slow continuum. The position of a species within this continuum is thought to reflect a negative relationship between growth and storage, with fast‐growing species allocating more carbon to growth and slow‐growing species investing more in storage. For tropical species, the relationship between storage and life‐history strategies has been largely studied on seedlings and less so in adult trees.

We evaluated how stored non‐structural carbohydrates (NSC) vary across adult trees spanning the fast–slow continuum in the Peruvian Amazon by: (a) analysing whole‐tree NSC in two species of contrasting growth and (b) investigating the relationships with key life‐history traits across a broader set of species.

Our results are consistent with a growth–storage trade‐off. The analysis of whole‐tree NSC revealed that the slow‐growing Eschweilera coriacea stored about 2.7 times as much NSC as the fast‐growing Bixa arborea due to markedly higher storage in woody stems and roots. B. arborea also had higher seasonality in NSC, reflecting its strong seasonality in stem growth. Across a range of species, stem starch was negatively related to species growth rate and positively related to wood density.

Given the role of NSC in mediating plants' response to stress, our results suggest that slow‐growing species with greater storage reserves may be more resilient to drought than fast‐growing species.

AbstractThe fate of tropical forests under future climate change is dependent on the capacity of their trees to adjust to drier conditions. The capacity of trees to withstand drought is likely to be determined by traits associated with their hydraulic systems. However, data on whether tropical trees can adjust hydraulic traits when experiencing drought remain rare. We measured plant hydraulic traits (e.g. hydraulic conductivity and embolism resistance) and plant hydraulic system status (e.g. leaf water potential, native embolism and safety margin) on >150 trees from 12 genera (36 species) and spanning a stem size range from 14 to 68 cm diameter at breast height at the world's only long‐running tropical forest drought experiment. Hydraulic traits showed no adjustment following 15 years of experimentally imposed moisture deficit. This failure to adjust resulted in these drought‐stressed trees experiencing significantly lower leaf water potentials, and higher, but variable, levels of native embolism in the branches. This result suggests that hydraulic damage caused by elevated levels of embolism is likely to be one of the key drivers of drought‐induced mortality following long‐term soil moisture deficit. We demonstrate that some hydraulic traits changed with tree size, however, the direction and magnitude of the change was controlled by taxonomic identity. Our results suggest that Amazonian trees, both small and large, have limited capacity to acclimate their hydraulic systems to future droughts, potentially making them more at risk of drought‐induced mortality.

The response of small understory trees to long-term drought is vital in determining the future composition, carbon stocks and dynamics of tropical forests. Long-term drought is, however, also likely to expose understory trees to increased light availability driven by drought-induced mortality. Relatively little is known about the potential for understory trees to adjust their physiology to both decreasing water and increasing light availability. We analysed data on maximum photosynthetic capacity (Jmax , Vcmax ), leaf respiration (Rleaf ), leaf mass per area (LMA), leaf thickness and leaf nitrogen and phosphorus concentrations from 66 small trees across 12 common genera at the world's longest running tropical rainfall exclusion experiment and compared responses to those from 61 surviving canopy trees. Small trees increased Jmax , Vcmax , Rleaf and LMA (71, 29, 32, 15% respectively) in response to the drought treatment, but leaf thickness and leaf nutrient concentrations did not change. Small trees were significantly more responsive than large canopy trees to the drought treatment, suggesting greater phenotypic plasticity and resilience to prolonged drought, although differences among taxa were observed. Our results highlight that small tropical trees have greater capacity to respond to ecosystem level changes and have the potential to regenerate resilient forests following future droughts.

Land surface models (LSMs) typically use empirical functions to represent vegetation responses to soil drought. These functions largely neglect recent advances in plant ecophysiology that link xylem hydraulic functioning with stomatal responses to climate. We developed an analytical stomatal optimization model based on xylem hydraulics (SOX) to predict plant responses to drought. Coupling SOX to the Joint UK Land Environment Simulator (JULES) LSM, we conducted a global evaluation of SOX against leaf- and ecosystem-level observations. SOX simulates leaf stomatal conductance responses to climate for woody plants more accurately and parsimoniously than the existing JULES stomatal conductance model. An ecosystem-level evaluation at 70 eddy flux sites shows that SOX decreases the sensitivity of gross primary productivity (GPP) to soil moisture, which improves the model agreement with observations and increases the predicted annual GPP by 30% in relation to JULES. SOX decreases JULES root-mean-square error in GPP by up to 45% in evergreen tropical forests, and can simulate realistic patterns of canopy water potential and soil water dynamics at the studied sites. SOX provides a parsimonious way to incorporate recent advances in plant hydraulics and optimality theory into LSMs, and an alternative to empirical stress factors.

Xylem vulnerability to embolism represents an important trait to determine species distribution patterns and drought resistance. However, estimating embolism resistance frequently requires time-consuming and ambiguous hydraulic lab measurements. Based on a recently developed pneumatic method, we present and test the "Pneumatron", a device that generates high time-resolution and fully automated vulnerability curves. Embolism resistance is estimated by applying a partial vacuum to extract air from an excised xylem sample, while monitoring the pressure change over time. Although the amount of gas extracted is strongly correlated with the percentage loss of xylem conductivity, validation of the Pneumatron was performed by comparison with the optical method for Eucalyptus camaldulensis leaves. The Pneumatron improved the precision of the pneumatic method considerably, facilitating the detection of small differences in the (percentage of air discharged [PAD] < 0.47%). Hence, the Pneumatron can directly measure the 50% PAD without any fitting of vulnerability curves. PAD and embolism frequency based on the optical method were strongly correlated (r2 = 0.93) for E. camaldulensis. By providing an open source platform, the Pneumatron represents an easy, low-cost, and powerful tool for field measurements, which can significantly improve our understanding of plant-water relations and the mechanisms behind embolism.

Abstract. Accurately representing the response of ecosystems to environmental change in land surface models (LSMs) is crucial to making accurate predictions of future climate. Many LSMs do not correctly capture plant respiration and growth fluxes, particularly in response to extreme climatic events. This is in part due to the unrealistic assumption that total plant carbon expenditure (PCE) is always equal to gross carbon accumulation by photosynthesis. We present and evaluate a simple model of labile carbon storage and utilisation (SUGAR) designed to be integrated into an LSM, which allows simulated plant respiration and growth to vary independent of photosynthesis. SUGAR buffers simulated PCE against seasonal variation in photosynthesis, producing more constant (less variable) predictions of plant growth and respiration relative to an LSM that does not represent labile carbon storage. This allows the model to more accurately capture observed carbon fluxes at a large-scale drought experiment in a tropical moist forest in the Amazon, relative to the Joint UK Land Environment Simulator LSM (JULES). SUGAR is designed to improve the representation of carbon storage in LSMs and provides a simple framework that allows new processes to be integrated as the empirical understanding of carbon storage in plants improves. The study highlights the need for future research into carbon storage and allocation in plants, particularly in response to extreme climate events such as drought.
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Abstract


Whether tropical trees acclimate to long‐term drought stress remains unclear. This uncertainty is amplified if drought stress is accompanied by changes in other drivers such as the increases in canopy light exposure that might be induced by tree mortality or other disturbances.

Photosynthetic capacity, leaf respiration, non‐structural carbohydrate (NSC) storage and stomatal conductance were measured on 162 trees at the world's longest running (15 years) tropical forest drought experiment. We test whether surviving trees have altered strategies for carbon storage and carbon use in the drier and elevated light conditions present following drought‐related tree mortality.

Relative to control trees, the surviving trees experiencing the drought treatment showed functional responses including: (a) moderately reduced photosynthetic capacity; (b) increased total leaf NSC; and (c) a switch from starch to soluble sugars as the main store of branch NSC. This contrasts with earlier findings at this experiment of no change in photosynthetic capacity or NSC storage. The changes detected here only occurred in the subset of drought‐stressed trees with canopies exposed to high radiation and were absent in trees with less‐exposed canopies and also in the community average. In contrast to previous results acquired through less intensive species sampling from this experiment, we also observe no species‐average drought‐induced change in leaf respiration.

Our results suggest that long‐term responses to drought stress are strongly influenced by a tree's full‐canopy light environment and therefore that disturbance‐induced changes in stand density and dynamics are likely to substantially impact tropical forest responses to climate change. We also demonstrate that, while challenging, intensive sampling is essential in tropical forests to avoid sampling biases caused by limited taxonomic coverage.

A free Plain Language Summary can be found within the Supporting Information of this article.

The absorption of atmospheric water directly into leaves enables plants to alleviate the water stress caused by low soil moisture, hydraulic resistance in the xylem and the effect of gravity on the water column, while enabling plants to scavenge small inputs of water from leaf-wetting events. By increasing the availability of water, and supplying it from the top of the canopy (in a direction facilitated by gravity), foliar uptake (FU) may be a significant process in determining how forests interact with climate, and could alter our interpretation of current metrics for hydraulic stress and sensitivity. FU has not been reported for lowland tropical rainforests; we test whether FU occurs in six common Amazonian tree genera in lowland Amazônia, and make a first estimation of its contribution to canopy-atmosphere water exchange. We demonstrate that FU occurs in all six genera and that dew-derived water may therefore be used to "pay" for some morning transpiration in the dry season. Using meteorological and canopy wetness data, coupled with empirically derived estimates of leaf conductance to FU (kfu ), we estimate that the contribution by FU to annual transpiration at this site has a median value of 8.2% (103 mm/year) and an interquartile range of 3.4%-15.3%, with the biggest sources of uncertainty being kfu and the proportion of time the canopy is wet. Our results indicate that FU is likely to be a common strategy and may have significant implications for the Amazon carbon budget. The process of foliar water uptake may also have a profound impact on the drought tolerance of individual Amazonian trees and tree species, and on the cycling of water and carbon, regionally and globally.

Biomass and area ratios between leaves, stems and roots regulate many physiological and ecological processes. The Huber value Hv (sapwood area/leaf area ratio) is central to plant water balance and drought responses. However, its coordination with key plant functional traits is poorly understood, and prevents developing trait-based prediction models. Based on theoretical arguments, we hypothesise that global patterns in Hv of terminal woody branches can be predicted from variables related to plant trait spectra, that is plant hydraulics and size and leaf economics. Using a global compilation of 1135 species-averaged Hv , we show that Hv varies over three orders of magnitude. Higher Hv are seen in short small-leaved low-specific leaf area (SLA) shrubs with low Ks in arid relative to tall large-leaved high-SLA trees with high Ks in moist environments. All traits depend on climate but climatic correlations are stronger for explanatory traits than Hv. Negative isometry is found between Hv and Ks , suggesting a compensation to maintain hydraulic supply to leaves across species. This work identifies the major global drivers of branch sapwood/leaf area ratios. Our approach based on widely available traits facilitates the development of accurate models of above-ground biomass allocation and helps predict vegetation responses to drought.

Tropical ecosystems harbour the highest concentrations of biodiversity on Earth and play a pivotal role in the global carbon cycle, yet deforestation and degradation continue unabated in many regions, with net forest loss at 5.5 million ha yr-1 between 2010 and 2015. Protected areas offer a partial solution to this problem, with a growing body of evidence demonstrating their effectiveness for habitat conservation in the dense forests of Amazonia, Central Africa and Southeast Asia. Despite containing over a quarter of global biodiversity hotspots and being low density but significant carbon stores, tropical drylands have received far less attention in conservation terms, and research into protected areas in these ecosystems is far more limited. The overall effectiveness of protected areas in different dryland regions, and the factors influencing performance, are less understood. By measuring protected area performance as a function of aboveground biomass change, this study investigated the effectiveness of protected areas in the savannah belt of Nigeria, a country with a long history of environmental degradation. L-band Synthetic Aperture Radar (SAR), a form of remote sensing that penetrates the vegetation canopy, provided a means of consistently monitoring aboveground biomass change over time. Twenty-one areas, ranging in size from 117,000 ha to 608,410 ha, and offering varying levels of protection according to IUCN designations, were selected, with aboveground biomass changes between 2007 and 2017 determined by subjecting L-band SAR data to a novel approach called ‘Biomass Matching’. The combination of SAR and Biomass Matching allowed aboveground biomass changes within these protected areas to be detected and estimated without the need for supplementary field data, which is usually required to calibrate such remote sensing data. All but four protected areas experienced increases in aboveground biomass over the study period, with mean change being +1.22 Mg ha-1, compared to +0.26 Mg ha-1 for a set of twelve similar unprotected areas. Furthermore, their performance was affected by an array of factors, though accessibility and management efficacy were deemed the most influential. These results suggest that, with appropriate monitoring and resourcing, protected areas in Nigerian dry forests and savannahs can provide effective habitat conservation, though more inaccessible areas will inherently perform better.

Abstract


Tropical forests and peatlands provide important ecological, climate and socio‐economic benefits from the local to the global scale. However, these ecosystems and their associated benefits are threatened by anthropogenic activities, including agricultural conversion, timber harvesting, peatland drainage and associated fire. Here, we identify key challenges, and provide potential solutions and future directions to meet forest and peatland conservation and restoration goals in Indonesia, with a particular focus on Kalimantan.

Through a round‐table, dual‐language workshop discussion and literature evaluation, we recognized 59 political, economic, legal, social, logistical and research challenges, for which five key underlying factors were identified. These challenges relate to the 3Rs adopted by the Indonesian Peatland Restoration Agency (Rewetting, Revegetation and Revitalization), plus a fourth R that we suggest is essential to incorporate into (peatland) conservation planning: Reducing Fires.

Our analysis suggests that (a) all challenges have potential for impact on activities under all 4Rs, and many are inter‐dependent and mutually reinforcing, implying that narrowly focused solutions are likely to carry a higher risk of failure; (b) addressing challenges relating to Rewetting and Reducing Fire is critical for achieving goals in all 4Rs, as is considering the local socio‐political situation and acquiring local government and community support; and (c) the suite of challenges faced, and thus conservation interventions required to address these, will be unique to each project, depending on its goals and prevailing local environmental, social and political conditions.

With this in mind, we propose an eight‐step adaptive management framework, which could support projects in both Indonesia and other tropical areas to identify and overcome their specific conservation and restoration challenges.

A free Plain Language Summary can be found within the Supporting Information of this article.

The Amazon rainforest presents high temperatures and annual precipitation, although there are large interannual variations in these meteorological elements. Air temperature (Tar) and relative air humidity (RH) in and above a forest are the result of complex energy exchanges through the processes of reflection, transmission, and absorption of solar energy. This study was carried out in the Caxiuanã National Forest, Pará, Brasil, and the objective was to evaluate the seasonal variability of air temperature and humidity from a vertical profile analysis within the forest with measures heights of 2, 16, 28 and 42 m, with readings taken every 30 minutes from 2012 to 2016. The results indicated a great seasonality in these meteorological elements, since the highest temperatures occurred at the canopy level (28 m), and the lowest ones were observed near the surface (2 m) due to the attenuation of solar radiation inside the forest. The highest RH values were observed near the surface (2 m), and the lowest values occurred above the canopy, due to the higher wind speeds at this level. These results indicate a large spatial-temporal variability of these meteorological elements, which influence the behavior of living organisms that inhabit that forest environment.

Meteorological extreme events such as El Niño events are expected to affect tropical forest net primary production (NPP) and woody growth, but there has been no large-scale empirical validation of this expectation. We collected a large high-temporal resolution dataset (for 1-13 years depending upon location) of more than 172 000 stem growth measurements using dendrometer bands from across 14 regions spanning Amazonia, Africa and Borneo in order to test how much month-to-month variation in stand-level woody growth of adult tree stems (NPPstem) can be explained by seasonal variation and interannual meteorological anomalies. A key finding is that woody growth responds differently to meteorological variation between tropical forests with a dry season (where monthly rainfall is less than 100 mm), and aseasonal wet forests lacking a consistent dry season. In seasonal tropical forests, a high degree of variation in woody growth can be predicted from seasonal variation in temperature, vapour pressure deficit, in addition to anomalies of soil water deficit and shortwave radiation. The variation of aseasonal wet forest woody growth is best predicted by the anomalies of vapour pressure deficit, water deficit and shortwave radiation. In total, we predict the total live woody production of the global tropical forest biome to be 2.16 Pg C yr-1, with an interannual range 1.96-2.26 Pg C yr-1 between 1996-2016, and with the sharpest declines during the strong El Niño events of 1997/8 and 2015/6. There is high geographical variation in hotspots of El Niño-associated impacts, with weak impacts in Africa, and strongly negative impacts in parts of Southeast Asia and extensive regions across central and eastern Amazonia. Overall, there is high correlation (r = -0.75) between the annual anomaly of tropical forest woody growth and the annual mean of the El Niño 3.4 index, driven mainly by strong correlations with anomalies of soil water deficit, vapour pressure deficit and shortwave radiation.This article is part of the discussion meeting issue 'The impact of the 2015/2016 El Niño on the terrestrial tropical carbon cycle: patterns, mechanisms and implications'.

The current generation of dynamic global vegetation models (DGVMs) lacks a mechanistic representation of vegetation responses to soil drought, impairing their ability to accurately predict Earth system responses to future climate scenarios and climatic anomalies, such as El Niño events. We propose a simple numerical approach to model plant responses to drought coupling stomatal optimality theory and plant hydraulics that can be used in dynamic global vegetation models (DGVMs). The model is validated against stand-scale forest transpiration (E) observations from a long-term soil drought experiment and used to predict the response of three Amazonian forest sites to climatic anomalies during the twentieth century. We show that our stomatal optimization model produces realistic stomatal responses to environmental conditions and can accurately simulate how tropical forest E responds to seasonal, and even long-term soil drought. Our model predicts a stronger cumulative effect of climatic anomalies in Amazon forest sites exposed to soil drought during El Niño years than can be captured by alternative empirical drought representation schemes. The contrasting responses between our model and empirical drought factors highlight the utility of hydraulically-based stomatal optimization models to represent vegetation responses to drought and climatic anomalies in DGVMs.This article is part of a discussion meeting issue 'The impact of the 2015/2016 El Niño on the terrestrial tropical carbon cycle: patterns, mechanisms and implications'.

Are short-term responses by tropical rainforest to drought (e.g. during El Niño) sufficient to predict changes over the long-term, or from repeated drought? Using the world's only long-term (16-year) drought experiment in tropical forest we examine predictability from short-term measurements (1-2 years). Transpiration was maximized in droughted forest: it consumed all available throughfall throughout the 16 years of study. Leaf photosynthetic capacity [Formula: see text] was maintained, but only when averaged across tree size groups. Annual transpiration in droughted forest was less than in control, with initial reductions (at high biomass) imposed by foliar stomatal control. Tree mortality increased after year three, leading to an overall biomass loss of 40%; over the long-term, the main constraint on transpiration was thus imposed by the associated reduction in sapwood area. Altered tree mortality risk may prove predictable from soil and plant hydraulics, but additional monitoring is needed to test whether future biomass will stabilize or collapse. Allocation of assimilate differed over time: stem growth and reproductive output declined in the short-term, but following mortality-related changes in resource availability, both showed long-term resilience, with partial or full recovery. Understanding and simulation of these phenomena and related trade-offs in allocation will advance more effectively through greater use of optimization and probabilistic modelling approaches.This article is part of a discussion meeting issue 'The impact of the 2015/2016 El Niño on the terrestrial tropical carbon cycle: patterns, mechanisms and implications'.

In the tropics, research, conservation and public attention focus on rain forests, but this neglects that half of the global tropics have a seasonally dry climate. These regions are home to dry forests and savannas (Figures 1 and 2), and are the focus of this Primer. The attention given to rain forests is understandable. Their high species diversity, sheer stature and luxuriance thrill biologists today as much as they did the first explorers in the Age of Discovery. Although dry forest and savanna may make less of a first impression, they support a fascinating diversity of plant strategies to cope with stress and disturbance including fire, drought and herbivory. Savannas played a fundamental role in human evolution, and across Africa and India they support iconic megafauna. Pennington et al. introduce seasonally dry biomes in the tropics – savannahs and dry forests.

Faster growth in tropical trees is usually associated with higher mortality rates, but the mechanisms underlying this relationship are poorly understood. In this study, we investigate how tree growth patterns are linked with environmental conditions and hydraulic traits, by monitoring the cambial growth of 9 tropical cloud forest tree species coupled with numerical simulations using an optimization model. We find that fast-growing trees have lower xylem safety margins than slow-growing trees and this pattern is not necessarily linked to differences in stomatal behaviour or environmental conditions when growth occurs. Instead, fast-growing trees have xylem vessels that are more vulnerable to cavitation and lower density wood. We propose the growth - xylem vulnerability trade-off represents a wood hydraulic economics spectrum similar to the classic leaf economic spectrum, and show through numerical simulations that this trade-off can emerge from the coordination between growth rates, wood density, and xylem vulnerability to cavitation. Our results suggest that vulnerability to hydraulic failure might be related with the growth-mortality trade-off in tropical trees, determining important life history differences. These findings are important in furthering our understanding of xylem hydraulic functioning and its implications on plant carbon economy.

AbstractHigh levels of species diversity hamper current understanding of how tropical forests may respond to environmental change. In the tropics, water availability is a leading driver of the diversity and distribution of tree species, suggesting that many tropical taxa may be physiologically incapable of tolerating dry conditions, and that their distributions along moisture gradients can be used to predict their drought tolerance. While this hypothesis has been explored at local and regional scales, large continental-scale tests are lacking. We investigate whether the relationship between drought-induced mortality and distributions holds continentally by relating experimental and observational data of drought-induced mortality across the Neotropics to the large-scale bioclimatic distributions of 115 tree genera. Across the different experiments, genera affiliated to wetter climatic regimes show higher drought-induced mortality than dry-affiliated ones, even after controlling for phylogenetic relationships. This pattern is stronger for adult trees than for saplings or seedlings, suggesting that the environmental filters exerted by drought impact adult tree survival most strongly. Overall, our analysis of experimental, observational, and bioclimatic data across neotropical forests suggests that increasing moisture-stress is indeed likely to drive significant changes in floristic composition.

AbstractTranspiration from the Amazon rainforest generates an essential water source at a global and local scale. However, changes in rainforest function with climate change can disrupt this process, causing significant reductions in precipitation across Amazonia, and potentially at a global scale. We report the only study of forest transpiration following a long‐term (&gt;10 year) experimental drought treatment in Amazonian forest. After 15 years of receiving half the normal rainfall, drought‐related tree mortality caused total forest transpiration to decrease by 30%. However, the surviving droughted trees maintained or increased transpiration because of reduced competition for water and increased light availability, which is consistent with increased growth rates. Consequently, the amount of water supplied as rainfall reaching the soil and directly recycled as transpiration increased to 100%. This value was 25% greater than for adjacent nondroughted forest. If these drought conditions were accompanied by a modest increase in temperature (e.g. 1.5°C), water demand would exceed supply, making the forest more prone to increased tree mortality.

The seasonal climate drivers of the carbon cycle in tropical forests remain poorly known, although these forests account for more carbon assimilation and storage than any other terrestrial ecosystem. Based on a unique combination of seasonal pan-tropical data sets from 89 experimental sites (68 include aboveground wood productivity measurements and 35 litter productivity measurements), their associated canopy photosynthetic capacity (enhanced vegetation index, EVI) and climate, we ask how carbon assimilation and aboveground allocation are related to climate seasonality in tropical forests and how they interact in the seasonal carbon cycle. We found that canopy photosynthetic capacity seasonality responds positively to precipitation when rainfall is &lt; 2000ĝ€-mmĝ€-yrĝ'1 (water-limited forests) and to radiation otherwise (light-limited forests). On the other hand, independent of climate limitations, wood productivity and litterfall are driven by seasonal variation in precipitation and evapotranspiration, respectively. Consequently, light-limited forests present an asynchronism between canopy photosynthetic capacity and wood productivity. First-order control by precipitation likely indicates a decrease in tropical forest productivity in a drier climate in water-limited forest, and in current light-limited forest with future rainfall &lt; 2000ĝ€-mmĝ€-yrĝ'1. Author(s) 2016.

Forest ecosystem models based on heuristic water stress functions poorly predict tropical forest response to drought partly because they do not capture the diversity of hydraulic traits (including variation in tree size) observed in tropical forests. We developed a continuous porous media approach to modeling plant hydraulics in which all parameters of the constitutive equations are biologically interpretable and measurable plant hydraulic traits (e.g. turgor loss point πtlp, bulk elastic modulus ϵ, hydraulic capacitance Cft, xylem hydraulic conductivity ks,max, water potential at 50% loss of conductivity for both xylem (P50,x) and stomata (P50,gs), and the leafg: sapwood area ratio Al: As). We embedded this plant hydraulics model within a trait forest simulator (TFS) that models light environments of individual trees and their upper boundary conditions (transpiration), as well as providing a means for parameterizing variation in hydraulic traits among individuals. We synthesized literature and existing databases to parameterize all hydraulic traits as a function of stem and leaf traits, including wood density (WD), leaf mass per area (LMA), and photosynthetic capacity (Amax), and evaluated the coupled model (called TFS v.1-Hydro) predictions, against observed diurnal and seasonal variability in stem and leaf water potential as well as stand-scaled sap flux. Our hydraulic trait synthesis revealed coordination among leaf and xylem hydraulic traits and statistically significant relationships of most hydraulic traits with more easily measured plant traits. Using the most informative empirical trait-trait relationships derived from this synthesis, TFS v.1-Hydro successfully captured individual variation in leaf and stem water potential due to increasing tree size and light environment, with model representation of hydraulic architecture and plant traits exerting primary and secondary controls, respectively, on the fidelity of model predictions. The plant hydraulics model made substantial improvements to simulations of total ecosystem transpiration. Remaining uncertainties and limitations of the trait paradigm for plant hydraulics modeling are highlighted.

Summary


The tropics are predicted to become warmer and drier, and understanding the sensitivity of tree species to drought is important for characterizing the risk to forests of climate change. This study makes use of a long‐term drought experiment in the Amazon rainforest to evaluate the role of leaf‐level water relations, leaf anatomy and their plasticity in response to drought in six tree genera.

The variables (osmotic potential at full turgor, turgor loss point, capacitance, elastic modulus, relative water content and saturated water content) were compared between seasons and between plots (control and through‐fall exclusion) enabling a comparison between short‐ and long‐term plasticity in traits. Leaf anatomical traits were correlated with water relation parameters to determine whether water relations differed among tissues.

The key findings were: osmotic adjustment occurred in response to the long‐term drought treatment; species resistant to drought stress showed less osmotic adjustment than drought‐sensitive species; and water relation traits were correlated with tissue properties, especially the thickness of the abaxial epidermis and the spongy mesophyll.

These findings demonstrate that cell‐level water relation traits can acclimate to long‐term water stress, and highlight the limitations of extrapolating the results of short‐term studies to temporal scales associated with climate change.

Summary


Leaf dark respiration (Rdark) represents an important component controlling the carbon balance in tropical forests. Here, we test how nitrogen (N) and phosphorus (P) affect Rdark and its relationship with photosynthesis using three widely separated tropical forests which differ in soil fertility.

Rdark was measured on 431 rainforest canopy trees, from 182 species, in French Guiana, Peru and Australia. The variation in Rdark was examined in relation to leaf N and P content, leaf structure and maximum photosynthetic rates at ambient and saturating atmospheric CO2 concentration.

We found that the site with the lowest fertility (French Guiana) exhibited greater rates of Rdark per unit leaf N, P and photosynthesis. The data from Australia, for which there were no phylogenetic overlaps with the samples from the South American sites, yielded the most distinct relationships of Rdark with the measured leaf traits.

Our data indicate that no single universal scaling relationship accounts for variation in Rdark across this large biogeographical space. Variability between sites in the absolute rates of Rdark and the Rdark : photosynthesis ratio were driven by variations in N‐ and P‐use efficiency, which were related to both taxonomic and environmental variability.

Summary: Leaf dark respiration (R dark ) is an important yet poorly quantified component of the global carbon cycle. Given this, we analyzed a new global database of R dark and associated leaf traits. Data for 899 species were compiled from 100 sites (from the Arctic to the tropics). Several woody and nonwoody plant functional types (PFTs) were represented. Mixed-effects models were used to disentangle sources of variation in R dark. Area-based R dark at the prevailing average daily growth temperature (T) of each site increased only twofold from the Arctic to the tropics, despite a 20°C increase in growing T (8-28°C). By contrast, R dark at a standard T (25°C, R dark25 ) was threefold higher in the Arctic than in the tropics, and twofold higher at arid than at mesic sites. Species and PFTs at cold sites exhibited higher R dark25 at a given photosynthetic capacity (V cmax25 ) or leaf nitrogen concentration ([N]) than species at warmer sites. R dark25 values at any given V cmax25 or [N] were higher in herbs than in woody plants. The results highlight variation in R dark among species and across global gradients in T and aridity. In addition to their ecological significance, the results provide a framework for improving representation of R dark in terrestrial biosphere models (TBMs) and associated land-surface components of Earth system models (ESMs).

Leaf dark respiration (Rdark ) is an important yet poorly quantified component of the global carbon cycle. Given this, we analyzed a new global database of Rdark and associated leaf traits. Data for 899 species were compiled from 100 sites (from the Arctic to the tropics). Several woody and nonwoody plant functional types (PFTs) were represented. Mixed-effects models were used to disentangle sources of variation in Rdark. Area-based Rdark at the prevailing average daily growth temperature (T) of each site increased only twofold from the Arctic to the tropics, despite a 20°C increase in growing T (8-28°C). By contrast, Rdark at a standard T (25°C, Rdark (25) ) was threefold higher in the Arctic than in the tropics, and twofold higher at arid than at mesic sites. Species and PFTs at cold sites exhibited higher Rdark (25) at a given photosynthetic capacity (Vcmax (25) ) or leaf nitrogen concentration ([N]) than species at warmer sites. Rdark (25) values at any given Vcmax (25) or [N] were higher in herbs than in woody plants. The results highlight variation in Rdark among species and across global gradients in T and aridity. In addition to their ecological significance, the results provide a framework for improving representation of Rdark in terrestrial biosphere models (TBMs) and associated land-surface components of Earth system models (ESMs).

Accurately predicting the response of Amazonia to climate change is important for predicting climate change across the globe. Changes in multiple climatic factors simultaneously result in complex non-linear ecosystem responses, which are difficult to predict using vegetation models. Using leaf- and canopy-scale observations, this study evaluated the capability of five vegetation models (Community Land Model version 3.5 coupled to the Dynamic Global Vegetation model - CLM3.5-DGVM; Ecosystem Demography model version 2 - ED2; the Joint UK Land Environment Simulator version 2.1 - JULES; Simple Biosphere model version 3 - SiB3; and the soil-plant-atmosphere model - SPA) to simulate the responses of leaf- and canopy-scale productivity to changes in temperature and drought in an Amazonian forest. The models did not agree as to whether gross primary productivity (GPP) was more sensitive to changes in temperature or precipitation, but all the models were consistent with the prediction that GPP would be higher if tropical forests were 5 °C cooler than current ambient temperatures. There was greater model-data consistency in the response of net ecosystem exchange (NEE) to changes in temperature than in the response to temperature by net photosynthesis (An), stomatal conductance (gs) and leaf area index (LAI). Modelled canopy-scale fluxes are calculated by scaling leaf-scale fluxes using LAI. At the leaf-scale, the models did not agree on the temperature or magnitude of the optimum points of An, Vcmax or gs, and model variation in these parameters was compensated for by variations in the absolute magnitude of simulated LAI and how it altered with temperature. Across the models, there was, however, consistency in two leaf-scale responses: (1) change in an with temperature was more closely linked to stomatal behaviour than biochemical processes; and (2) intrinsic water use efficiency (IWUE) increased with temperature, especially when combined with drought. These results suggest that even up to fairly extreme temperature increases from ambient levels (+6 °C), simulated photosynthesis becomes increasingly sensitive to gs and remains less sensitive to biochemical changes. To improve the reliability of simulations of the response of Amazonian rainforest to climate change, the mechanistic underpinnings of vegetation models need to be validated at both leaf- and canopy-scales to improve accuracy and consistency in the quantification of processes within and across an ecosystem.

© 2015 Macmillan Publishers Limited. All rights reserved.Stomatal conductance (g s) is a key land-surface attribute as it links transpiration, the dominant component of global land evapotranspiration, and photosynthesis, the driving force of the global carbon cycle. Despite the pivotal role of g s in predictions of global water and carbon cycle changes, a global-scale database and an associated globally applicable model of g s that allow predictions of stomatal behaviour are lacking. Here, we present a database of globally distributed g s obtained in the field for a wide range of plant functional types (PFTs) and biomes. We find that stomatal behaviour differs among PFTs according to their marginal carbon cost of water use, as predicted by the theory underpinning the optimal stomatal model and the leaf and wood economics spectrum. We also demonstrate a global relationship with climate. These findings provide a robust theoretical framework for understanding and predicting the behaviour of g s across biomes and across PFTs that can be applied to regional, continental and global-scale modelling of ecosystem productivity, energy balance and ecohydrological processes in a future changing climate.