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Funding: Data collection was largely funded by the UK Natural Environment Research Council (NERC) project TREMOR (NE/N004655/1) to D.G., E.G. and O.P., with further funds from Coordenação de Aperfeiçoamento de Pessoal de Nível Superior—Brasil (CAPES, finance code 001) to J.V.T. and a University of Leeds Climate Research Bursary Fund to J.V.T. D.G., E.G. and O.P. acknowledge further support from a NERC-funded consortium award (ARBOLES, NE/S011811/1). This paper is an outcome of J.V.T.’s doctoral thesis, which was sponsored by CAPES (GDE 99999.001293/2015-00). J.V.T. was previously supported by the NERC-funded ARBOLES project (NE/S011811/1) and is supported at present by the Swedish Research Council Vetenskapsrådet (grant no. 2019-03758 to R.M.). E.G., O.P. and D.G. acknowledge support from NERC-funded BIORED grant (NE/N012542/1). O.P. acknowledges support from an ERC Advanced Grant and a Royal Society Wolfson Research Merit Award. R.S.O. was supported by a CNPq productivity scholarship, the São Paulo Research Foundation (FAPESP-Microsoft 11/52072-0) and the US Department of Energy, project GoAmazon (FAPESP 2013/50531-2). M.M. acknowledges support from MINECO FUN2FUN (CGL2013-46808-R) and DRESS (CGL2017-89149-C2-1-R). C.S.-M., F.B.V. and P.R.L.B. were financed by Coordenação de Aperfeiçoamento de Pessoal de Nível Superior—Brasil (CAPES, finance code 001). C.S.-M. received a scholarship from the Brazilian National Council for Scientific and Technological Development (CNPq 140353/2017-8) and CAPES (science without borders 88881.135316/2016-01). Y.M. acknowledges the Gordon and Betty Moore Foundation and ERC Advanced Investigator Grant (GEM-TRAITS, 321131) for supporting the Global Ecosystems Monitoring (GEM) network (gem.tropicalforests.ox.ac.uk), within which some of the field sites (KEN, TAM and ALP) are nested. The authors thank Brazil–USA Collaborative Research GoAmazon DOE-FAPESP-FAPEAM (FAPESP 2013/50533-5 to L.A.) and National Science Foundation (award DEB-1753973 to L. Alves). They thank Serrapilheira Serra-1709-18983 (to M.H.) and CNPq-PELD/POPA-441443/2016-8 (to L.G.) (P.I. Albertina Lima). They thank all the colleagues and grants mentioned elsewhere [8,36] that established, identified and measured the Amazon forest plots in the RAINFOR network analysed here. The authors particularly thank J. Lyod, S. Almeida, F. Brown, B. Vicenti, N. Silva and L. Alves. This work is an outcome approved Research Project no. 19 from ForestPlots.net, a collaborative initiative developed at the University of Leeds that unites researchers and the monitoring of their permanent plots from the world’s tropical forests [61]. The authros thank A. Levesley, K. Melgaço Ladvocat and G. Pickavance for ForestPlots.net management. They thank Y. Wang and J. Baker, respectively, for their help with the map and with the climatic data. The authors acknowledge the invaluable help of M. Brum for kindly providing the comparison of vulnerability curves based on PAD and on PLC shown in this manuscript. They thank J. Martinez-Vilalta for his comments on an early version of this manuscript. The authors also thank V. Hilares and the Asociación para la Investigación y Desarrollo Integral (AIDER, Puerto Maldonado, Peru); V. Saldaña and Instituto de Investigaciones de la Amazonía Peruana (IIAP) for local field campaign support in Peru; E. Chavez and Noel Kempff Natural History Museum for local field campaign support in Bolivia; ICMBio, INPA/NAPPA/LBA COOMFLONA (Cooperativa mista da Flona Tapajós) and T. I. Bragança-Marituba for the research support. Tropical 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, Ψ50) 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 Ψ50 and HSM50 vary markedly across the Amazon and are related to average long-term rainfall characteristics. Both Ψ50 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. Publisher PDF

The continued functioning of tropical forests under climate change depends on their resilience to drought and heat. However, there is little understanding of how tropical forests will respond to combinations of these stresses, and no field studies to date have explicitly evaluated whether sustained drought alters sensitivity to temperature. We measured the temperature response of net photosynthesis, foliar respiration and the maximum quantum efficiency of photosystem II (F

The incidence and severity of global mangrove mortality due to drought is increasing. Yet, little is understood of the capacity of mangroves to show long-term acclimation of leaf water relations to severe drought. We tested for differences between mid-dry season leaf water relations in two cooccurring mangroves, Aegiceras corniculatum and Rhizophora stylosa before a severe drought (a heatwave combined with low rainfall) and after its relief by the wet season. Consistent with ecological stress memory, the legacy of severe drought enhanced salinity tolerance in the subsequent dry season through coordinated adjustments that reduced the leaf water potential at the turgor loss point and increased cell wall rigidity. These adjustments enabled maintenance of turgor and relative water content with increasing salinity. As most canopy growth occurs during the wet season, acclimation to the ‘memory’ of higher salinity in the previous dry season enables greater leaf function with minimal adjustments, as long-lived leaves progress from wet through dry seasons. However, declining turgor safety margins - the difference between soil water potential and leaf water potential at turgor loss - implied increasing limitation to water use with increasing salinity. Thus, plasticity in leaf water relations contributes fundamentally to mangrove function under varying salinity regimes.

Climate warming is causing compositional changes in Andean tropical montane forests (TMFs). These shifts are hypothesised to result from differential responses to warming of cold- and warm-affiliated species, with the former experiencing mortality and the latter migrating upslope. The thermal acclimation potential of Andean TMFs remains unknown.Along a 2000 m Andean altitudinal gradient, we planted individuals of cold- and warm-affiliated species (under common soil and irrigation), exposing them to the hot and cold extremes of their thermal niches, respectively. We measured the response of net photosynthesis (Anet), photosynthetic capacity and leaf dark respiration (Rdark) to warming/cooling, 5 months after planting.In all species, Anet and photosynthetic capacity at 25°C were highest when growing at growth temperatures (Tg) closest to their thermal means, declining with warming and cooling in cold-affiliated and warm-affiliated species, respectively. When expressed at Tg, photosynthetic capacity and Rdark remained unchanged in cold-affiliated species, but the latter decreased in warm-affiliated counterparts. Rdark at 25°C increased with temperature in all species, but remained unchanged when expressed at Tg.Both species groups acclimated to temperature, but only warm-affiliated species decreased Rdark to photosynthetic capacity ratio at Tg as temperature increased. This could confer them a competitive advantage under future warming.

1. Occult precipitation events (fog, dew, light-rain) can alter plant water and nutritional status, both directly through the aerial uptake of surface water and nutrients, and indirectly via redistribution of atmospheric resources to the soil. However, current frameworks that explain niche segregation, species interactions, and coexistence still consider that ground-rooted plants obtain resources almost exclusively via root absorption from soil.2. Here, we expand the plant hydrological niches model to incorporate both soil and atmospheric resource-axes, thus providing a more complete picture of how ground-rooted terrestrial plants obtain, remobilize, share, and compete for water and soluble nutrients.3. First, we describe how plants with different water acquisition strategies access directly or indirectly atmospheric resources. Then, we discuss how the use of such resources may promote spatiotemporal niche segregation, contributing to shape species distribution and abundance within plant communities. We illustrate this argument with examples from arid, mesic, and wet vegetation types. Finally, we examine how climate and land-use changes may influence plant hydrological niches, potentially altering community structure.4. Synthesis: Understanding how available atmospheric resources influences niche segregation in plant communities is a crucial step towards better predictions of species responses (e.g., changes in distribution, abundance and interactions) to climate change.

Tropical montane forests are among the most productive ecosystems within the tropical region and store a significant amount of carbon in live biomass. With ongoing global climate warming, tropical climates are also getting warmer. The productivity and climate feedbacks of future tropical montane forests depend on the ability of trees to acclimate their photosynthetic metabolism to these new, warmer conditions. However, knowledge of acclimation ability of photosynthesis and its underlying biochemical processes to warming in trees grown under natural field conditions is currently limited due to data scarcity. To reduce this knowledge gap, we used two separate field experiments located in Colombia (with 15 species) and Rwanda (16 species), and for each experiment, tree species were grown at three different sites along an elevation gradient differing in ambient air temperature. At all three sites, we measured the responses of net CO2 assimilation at different CO2 concentration (50 to 2000 ppm) and at different leaf temperatures (15 to 40 °C) in ≈ 3 years old trees. We used these data to derive key photosynthetic biochemical parameters (maximum Rubisco carboxylation capacity - Vcmax and maximum electron transport rate - Jmax) and their temperature sensitivity, as well as the thermal optimum of net photosynthesis (ToptA). We show that tropical montane tree species from the two continents are generally able to acclimate their ToptA by increasing in trees grown in warmer conditions, but the magnitude of change in ToptA differs among species from different successional groups (early- versus late succession) and climate of origin (lowland versus montane). Shifts in ToptA are largely driven by concomitant changes in thermal sensitivity parameters of underlying biochemical processes of photosynthesis (Vcmax and Jmax) with warming. We also show that, at a standard temperature of 25 °C, Vcmax is largely constant, while Jmax decreases with warming. Our findings indicate tropical montane tree species from Latin America and Africa can thermally acclimate their photosynthetic physiology, but that this thermal acclimation ability is related to species successional group and their climate of origin.

Australia is the driest inhabited continent. Annual rainfall is low and is accompanied by marked inter-annual variability, leading to multi-year droughts. n particular, ​South-East Australia​ ​has recently experienced two of the worst droughts in the historical record (2000–2009 and 2017–2020). Predicting species-level responses to drought at the landscape scale is critical to reducing uncertainty in future terrestrial carbon and water cycle projections. We embedded a stomatal optimisation model in the Community Atmosphere Biosphere Land Exchange (CABLE) land surface model and parameterised the model for 15 canopy-dominant eucalypt tree species across South-Eastern Australia (mean annual precipitation range: 344–1424 mm yr-1). We carried out three experiments: applying CABLE to the recent drought; a theoretical future drier drought (20% reduction in rainfall); and a future drier drought (20% reduction in rainfall) with a doubling of atmospheric carbon dioxide (CO2). The drought's severity was highlighted as at least 25% of their distribution ranges, and 60% of species experienced leaf water potentials beyond the water potential at which 50% of hydraulic conductivity is lost due to embolism. We identified areas of severe hydraulic stress within species’ ranges, but we also pinpointed resilience in species found in predominantly semiarid regions. The importance of the role of CO2 in ameliorating drought stress was consistent across species. Our results represent an important advance in our capacity to forecast the resilience of individual tree species, providing an evidence base for decision-making around the resilience of restoration plantings or net-zero emission strategies.

Tropical forests are expected to be highly vulnerable to climate change. Observations from the tropical montane Andean forests report a change in composition towards a greater relative abundance of warm affiliated species, i.e thermophilic species. These shifts are hypothesised to result from differential responses to warming of cold- and warm-affiliated species, with the former experiencing mortality and the latter migrating upslope. However, the drivers of these changes are poorly understood. Along a 2000m altitudinal gradient/thermosequence in the Colombian Andes, we planted 2-yr old individuals of cold- and warm-affiliated species under common soil and water conditions, exposing them to the hot and cold extremes of their thermal niches, respectively. We show that cold-affiliated species growing outside and on the hotter portion of their thermal ranges decreased their growth. Warm-affiliated species can survive but reduce their growth under the colder portion of their thermal distribution. We demonstrate that growth responses are related to species’ thermal distributions; survival probability increased as species’ distribution optima were warmer than the experimental site and decreased as species’ distribution optima were colder than the study sites. These results can be explained by the negative effects of heat stress on simulated photosynthesis. Our findings highlight the potential effects of rapid warming on the composition of highland forest species in this biodiversity hotspot.

Climate change is projected to increase the imbalance between the supply (precipitation) and atmospheric demand for water (i.e., increased potential evapotranspiration), stressing plants in water-limited environments. Plants may be able to offset increasing aridity because rising CO2 increases water use efficiency. CO2 fertilization has also been cited as one of the drivers of the widespread “greening” phenomenon. However, attributing the size of this CO2 fertilization effect is complicated, due in part to a lack of long-term vegetation monitoring and interannual- to decadal-scale climate variability. In this study we asked the question of how much CO2 has contributed towards greening. We focused our analysis on a broad aridity gradient spanning eastern Australia's woody ecosystems. Next we analyzed 38 years of satellite remote sensing estimates of vegetation greenness (normalized difference vegetation index, NDVI) to examine the role of CO2 in ameliorating climate change impacts. Multiple statistical techniques were applied to separate the CO2-attributable effects on greening from the changes in water supply and atmospheric aridity. Widespread vegetation greening occurred despite a warming climate, increases in vapor pressure deficit, and repeated record-breaking droughts and heat waves. Between 1982–2019 we found that NDVI increased (median 11.3 %) across 90.5 % of the woody regions. After masking disturbance effects (e.g., fire), we statistically estimated an 11.7 % increase in NDVI attributable to CO2, broadly consistent with a hypothesized theoretical expectation of an 8.6 % increase in water use efficiency due to rising CO2. In contrast to reports of a weakening CO2 fertilization effect, we found no consistent temporal change in the CO2 effect. We conclude rising CO2 has mitigated the effects of increasing aridity, repeated record-breaking droughts, and record-breaking heat waves in eastern Australia. However, we were unable to determine whether trees or grasses were the primary beneficiary of the CO2-induced change in water use efficiency, which has implications for projecting future ecosystem resilience. A more complete understanding of how CO2-induced changes in water use efficiency affect trees and non-tree vegetation is needed.

Extreme drought events in Amazon forests are expected to become more frequent and more intense with climate change, threatening ecosystem function and carbon balance. Yet large uncertainties exist on the resilience of this ecosystem to drought. A better quantification of tree hydraulics and mortality processes is needed to anticipate future drought effects on Amazon forests. Most state-of-the-art dynamic global vegetation models are relatively poor in their mechanistic description of these complex processes. Here, we implement a mechanistic plant hydraulic module within the ORCHIDEE-CAN-NHA r7236 land surface model to simulate the percentage loss of conductance (PLC) and changes in water storage among organs via a representation of the water potentials and vertical water flows along the continuum from soil to roots, stems and leaves. The model was evaluated against observed seasonal variability in stand-scale sap flow, soil moisture and productivity under both control and drought setups at the Caxiuanã throughfall exclusion field experiment in eastern Amazonia between 2001 and 2008. A relationship between PLC and tree mortality is built in the model from two empirical parameters, the cumulated drought exposure duration that triggers mortality, and the mortality fraction in each day exceeding the exposure. Our model captures the large biomass drop in the year 2005 observed four years after throughfall reduction, and produces comparable annual tree mortality rates with observation over the study period. Our hydraulic architecture module provides promising avenues for future research in assimilating experimental data to parameterize mortality due to drought-induced xylem dysfunction. We also highlight that species-based (isohydric or anisohydric) hydraulic traits should be further tested to generalize the model performance in predicting the drought risks.

Water potential is the principal driving force for the movement of water through soils and plants, and directly influences plant physiological responses. The relationships between water potential and water content in plants and soil have long been of interest, and there is increasing focus on understanding how these fundamental measures of water are linked at larger spatial and temporal scales. In this Perspective, we explore how the theory of pressure-volume relationships can be applied at ecosystem scale. We define and evaluate the concept and limitations of the ecosystem pressure-volume curve (EPV), and discuss practical ways to construct EPVs with existing data. EPVs were generated from equilibrium water potentials and water content of the above ground biomass of nine plots including tropical rainforest, savanna, temperate forest, and a long-term Amazonian rainforest drought experiment. Initial findings suggest high levels of consistency among sites where the steady-state ratio of water:biomass appears to be approximately 1:3, while ecosystem values of relative hydraulic capacitance and accessible water storage do not vary systematically with biomass. The EPV reveals useful trends across ecosystems, providing a thermodynamically consistent steady-state view of ecosystem form and function, and a biophysically robust basis for the interpretation of microwave remote sensing data.

Soil-leaf hydraulic conductance determines canopy-atmosphere coupling in vegetation models, but is typically derived from ex-situ measurements of stem segments and soil samples. Using a novel approach we derive robust in-situ estimates for whole tree conductance (ktree ), 'functional' soil conductance (ksoil ), and 'system' conductance (ksystem , water table to canopy), at two climatically different tropical rainforest sites. Hydraulic 'functional rooting depth', determined for each tree using profiles of soil water potential (Ψsoil ) and sap flux data, enabled a robust determination of ktree and ksoil . ktree was compared across species, size classes, seasons, height above nearest drainage (HAND), two field sites, and to alternative representations of ktree ; ksoil was analysed with respect to variations in site, season and HAND. Key results: i) ktree was lower and changed seasonally at the site with higher vapour pressure deficit (VPD) and rainfall, ii) ktree differed little across species but scaled with tree circumference, iii) rsoil (1/ksoil ) ranged from 0 in wet season to 10x less than rtree (1/ktree ) in the dry season. Conclusions VPD and not rainfall may influence plot-level k; leaf water potentials and sap flux can be used to determine ktree , ksoil and ksystem ; Ψsoil profiles can provide mechanistic insights into ecosystem-level water fluxes.

The ecosystem pressure-volume curve (EPV) is the relationship between vegetation water content and a representative value of water potential applied on a ground-area basis. The EPV attempts to reconcile our detailed and physically rigorous understanding of small-scale field-measureable processes to the spatial scale applicable to ecosystem and climate science. Successfully bridging that gap in scale potentially allows us to use field measurements to interpret remote sensing data, and then remote sensing data to inform our understanding of vegetation-climate interactions. Here we clearly define the idea of the EPV, evaluate the limitations of applying values of water content and water potential to ecosystems on a ground area basis, and discuss practical ways to construct the EPV with existing data. We also present the first EPVs based on data from nine different plots, including tropical rainforest, savanna, temperate forest, and a long-term drought experiment in Amazonian rainforest (Caxiuanã, State of Pará, Brazil). The initial findings suggest high levels of consistency among sites. In particular, the ratio of water to biomass across ecosystems appears to be constrained to around 1:3. Seven of nine sites had closely converging ‘relative maximum water storage’ (the proportion of total stored water than can be lost before an ecosystem succumbs to physiological damage) at 9.1% +/-1.8 standard deviation. Relative ecosystem capacitance may increase with site biomass (P = 0.091), but varied little across sites with a mean of 0.068 MPa−1 +/-0.029 standard deviation. These first estimates suggest that the EPV idea may reveal useful trends across ecosystems, potentially paving the way to increasing the ecophysiological significance of remote sensing data, and enabling an alternative method for modelling long-term ecosystem-climate feedbacks based on equilibrium thermodynamics.

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. Made available in DSpace on 2021-01-12T09:05:37Z (GMT). No. of bitstreams: 1 nph.16972-2021.pdf: 1264820 bytes, checksum: 64adf8c9966954ff58dceff8df2c98cf (MD5) Previous issue date: 2021 Publicado Online em 27 set. 2020.

Climate warming could destabilise the Earth's largest terrestrial store of reactive carbon (C), by accelerating the decomposition of soil organic matter. A third of that C store resides in the tropics. The potential for tropical soils to sequester C, or to act as an additional source of CO2, will depend on the balance of C inputs and outputs, mediated by the response of soil microbial communities and their activity to perturbation. We review the impact of warming on microbial communities and C storage in humid tropical forest soils over multiple time-scales. Recent in situ experiments indicate high sensitivity of tropical forest soil C mineralisation to warming in the short term. However, whether this will translate into long-term soil C decline remains unclear. At decadal time-scales, high sensitivity of soil C mineralisation to warming is consistent with the correlation between the inter-annual variation in the tropical land surface temperature and atmospheric CO2 growth rate, and with simulations using the Carnegie-Ames-Stanford Approach biosphere model. This observed sensitivity may further contribute to climatic change over millennial time-scales, suggested by radiocarbon dating of organic matter in river basins showing a twofold acceleration in tropical soil C release during the late-glacial warming period. However, counter to this evidence, long-term stability of tropical soil C is suggested by observed steady-state soil C turnover across temperature gradients with elevation, and by the presence of C in tropical soils that pre-dates the Holocene Thermal Maximum and late-glacial warming periods. To help reconcile these recent experimental findings and long-term observations, we propose mechanisms to explain tropical soil C and microbial responses to warming across multiple time-scales. Combined in situ experimental and monitoring approaches—large-scale and cross-site—are urgently needed to resolve the interplay of these mechanisms across spatial and temporal scales, to shape a better understanding of the relationship between soil microbes and C storage in tropical soils.

Predicting species-level responses to drought at the landscape scale is critical to reducing future uncertainty in terrestrial carbon and water cycle projections. We embedded a stomatal optimisation model in the Community Atmosphere Biosphere Land Exchange (CABLE) land surface model. We parameterised the model for 15 canopy dominant eucalypt tree species representative of a broad precipitation gradient across South East Australia (mean annual precipitation range: 344–1424 mm yr-1). We conducted three experiments: (i) applying CABLE to the 2017–2019 drought in South East Australia; (ii) a 20% drier drought; and (iii) a 20% drier drought with a doubling of atmospheric carbon dioxide (CO2). We identified several drought hotspots across the ranges of E.viminalis, E.obliqua, E.globulus, E.saligna, and E.grandis. By contrast, CABLE simulated drought resilience in species that are found predominately in semi-arid areas such as E.largiflorens and E.populnea. We identified several key model assumptions (e.g., the degree of stomatal control) and sensitivities (e.g., the role of CO2 in ameliorating drought) that require future research. Our results represent an important step forward in our capacity to forecast the resilience of individual tree species, providing an evidence base for decision-making around the resilience of restoration plantings or strategies associated with achieving net-zero emissions.

South-East Australia has recently been subjected to two of the worst droughts in the historical record (Millennium Drought, 2000-2009 and Big Dry, 2017-2019). Unfortunately, a lack of forest monitoring has made it difficult to determine whether widespread tree mortality has resulted from these droughts. Anecdotal observations suggest the Big Dry may have led to more significant tree mortality than the Millennium drought. Critically, to be able to robustly project future expected climate change effects on Australian vegetation, we need to assess the vulnerability of Australian trees to drought. Here we implemented a model of plant hydraulics into the Community Atmosphere Biosphere Land Exchange (CABLE) land surface model. We parameterized the drought response behaviour of five broad vegetation types, based on a common garden dry-down experiment with species originating across a rainfall gradient (188-1,125 mm/year) across South-East Australia. The new hydraulics model significantly improved (similar to 35%-45% reduction in root mean square error) CABLE's previous predictions of latent heat fluxes during periods of water stress at two eddy covariance sites in Australia. Landscape-scale predictions of the greatest percentage loss of hydraulic conductivity (PLC) of about 40%-60%, were broadly consistent with satellite estimates of regions of the greatest change in both droughts. In neither drought did CABLE predict that trees would have reached critical PLC in widespread areas (i.e. it projected a low mortality risk), although the model highlighted critical levels near the desert regions of South-East Australia where few trees live. Overall, our experimentally constrained model results imply significant resilience to drought conferred by hydraulic function, but also highlight critical data and scientific gaps. Our approach presents a promising avenue to integrate experimental data and make regional-scale predictions of potential drought-induced hydraulic failure.

Tropical soils contain one-third of the carbon stored in soils globally1, so destabilization of soil organic matter caused by the warming predicted for tropical regions this century2 could accelerate climate change by releasing additional carbon dioxide (CO2) to the atmosphere3–6. Theory predicts that warming should cause only modest carbon loss from tropical soils relative to those at higher latitudes5,7, but there have been no warming experiments in tropical forests to test this8. Here we show that in situ experimental warming of a lowland tropical forest soil on Barro Colorado Island, Panama, caused an unexpectedly large increase in soil CO2 emissions. Two years of warming of the whole soil profile by four degrees Celsius increased CO2 emissions by 55 per cent compared to soils at ambient temperature. The additional CO2 originated from heterotrophic rather than autotrophic sources, and equated to a loss of 8.2 ± 4.2 (one standard error) tonnes of carbon per hectare per year from the breakdown of soil organic matter. During this time, we detected no acclimation of respiration rates, no thermal compensation or change in the temperature sensitivity of enzyme activities, and no change in microbial carbon-use efficiency. These results demonstrate that soil carbon in tropical forests is highly sensitive to warming, creating a potentially substantial positive feedback to climate change. When tropical forest soils are warmed in situ, they release more CO2 than predicted by theory, creating a potentially substantial positive feedback to climate change.

The productivity of rainforests growing on highly weathered tropical soils is expected to be limited by phosphorus availability1. Yet, controlled fertilization experiments have been unable to demonstrate a dominant role for phosphorus in controlling tropical forest net primary productivity. Recent syntheses have demonstrated that responses to nitrogen addition are as large as to phosphorus2, and adaptations to low phosphorus availability appear to enable net primary productivity to be maintained across major soil phosphorus gradients3. Thus, the extent to which phosphorus availability limits tropical forest productivity is highly uncertain. The majority of the Amazonia, however, is characterized by soils that are more depleted in phosphorus than those in which most tropical fertilization experiments have taken place2. Thus, we established a phosphorus, nitrogen and base cation addition experiment in an old growth Amazon rainforest, with a low soil phosphorus content that is representative of approximately 60% of the Amazon basin. Here we show that net primary productivity increased exclusively with phosphorus addition. After 2 years, strong responses were observed in fine root (+29%) and canopy productivity (+19%), but not stem growth. The direct evidence of phosphorus limitation of net primary productivity suggests that phosphorus availability may restrict Amazon forest responses to CO2 fertilization4, with major implications for future carbon sequestration and forest resilience to climate change. The authors acknowledge funding from the UK Natural Environment Research Council (NERC), grant number NE/L007223/1. This is publication 850 in the technical series of the BDFFP. C.A.Q. acknowledges the grants from Brazilian National Council for Scientific and Technological Development (CNPq) CNPq/LBA 68/2013, CNPq/MCTI/FNDCT no. 18/2021 and his productivity grant. C.A.Q., H.F.V.C., F.D.S., I.A., L.F.L., E.O.M. and S.G. acknowledge the AmazonFACE programme for financial support in cooperation with Coordination for the Improvement of Higher Education Personnel (CAPES) and the National Institute of Amazonian Research as part of the grants CAPES-INPA/88887.154643/2017-00 and 88881.154644/2017-01. T.F.D. acknowledges funds from FundacAo de Amparo a Pesquisa do Estado de SAo Paulo (FAPESP), grant 2015/50488-5, and the Partnership for Enhanced Engagement in Research (PEER) programme grant AID-OAA-A-11-00012. L.E.O.C.A. thanks CNPq (314416/2020-0).

Droughts in a warming climate have become more common and more extreme, making understanding forest responses to water stress increasingly pressing. Analysis of water stress in trees has long focused on water potential in xylem and leaves, which influences stomatal closure and water flow through the soil-plant-atmosphere continuum. At the same time, changes of vegetation water content (VWC) are linked to a range of tree responses, including fluxes of water and carbon, mortality, flammability, and more. Unlike water potential, which requires demanding in situ measurements, VWC can be retrieved from remote sensing measurements, particularly at microwave frequencies using radar and radiometry. Here, we highlight key frontiers through which VWC has the potential to significantly increase our understanding of forest responses to water stress. To validate remote sensing observations of VWC at landscape scale and to better relate them to data assimilation model parameters, we introduce an ecosystem-scale analog of the pressure–volume curve, the non-linear relationship between average leaf or branch water potential and water content commonly used in plant hydraulics. The sources of variability in these ecosystem-scale pressure-volume curves and their relationship to forest response to water stress are discussed. We further show to what extent diel, seasonal, and decadal dynamics of VWC reflect variations in different processes relating the tree response to water stress. VWC can also be used for inferring below-ground conditions—which are difficult to impossible to observe directly. Lastly, we discuss how a dedicated geostationary spaceborne observational system for VWC, when combined with existing datasets, can capture diel and seasonal water dynamics to advance the science and applications of global forest vulnerability to future droughts.

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. Made available in DSpace on 2021-01-13T09:05:01Z (GMT). No. of bitstreams: 1 1365-2435.13689-2021.pdf: 1614823 bytes, checksum: 0ea046c5e0666a2e409e93fe849fd23e (MD5) Previous issue date: 2021 Publicado online em 29 set. 2020.

Climate change is projected to increase the imbalance between the supply (precipitation) and atmospheric demand for water (i.e. increased potential evapotranspiration), stressing plants in water-limited environments. Plants may be able to offset increasing aridity because rising CO2 increases water-use-efficiency. CO2 fertilization has also been cited as one of the drivers of the widespread ‘greening’ phenomenon. However, attributing the size of this CO2 fertilization effect is complicated, due in part to a lack of long-term vegetation monitoring and interannual to decadal-scale climate variability. In this study we asked the question, how much has CO2 contributed towards greening? We focused our analysis on a broad aridity gradient spanning eastern Australia’s woody ecosystems. Next we analysed 38-years of satellite remote sensing estimates of vegetation greenness (normalized difference vegetation index, NDVI) to examine the role of CO2 in ameliorating climate change impacts. Multiple statistical techniques were applied to separate the CO2-attributable effects on greening from the changes in water supply and atmospheric aridity. Widespread vegetation greening occurred despite a warming climate, increases in vapor pressure deficit, and repeated record-breaking droughts and heatwaves. Between 1982–2019 we found that NDVI increased (median 11.3 %) across 90.5 % of the woody regions. After masking disturbance effects (e.g. fire), we statistically estimated an 11.7 % increase in NDVI attributable to CO2, broadly consistent with a hypothesized theoretical expectation of an 8.6 % increase in water-use-efficiency due to rising CO2. In contrast to reports of a weakening CO2 fertilization effect, we found no consistent temporal change in the CO2 effect. We conclude rising CO2 has mitigated the effects of increasing aridity, repeated record-breaking droughts, and record-breaking heat waves in eastern Australia. However, we were unable to determine whether trees or grasses were the primary beneficiary of the CO2 induced change in water-use-efficiency, which has implications for projecting future ecosystem resilience. A more complete understanding of how CO2 induced changes in water-use-efficiency affect trees and non-tree vegetation is needed.

Fine roots constitute a significant component of the net primary productivity (NPP) of forest ecosystems but are much less studied than aboveground NPP. Comparisons across sites and regions are also hampered by inconsistent methodologies, especially in tropical areas. Here, we present a novel dataset of fine root biomass, productivity, residence time, and allocation in tropical old-growth rainforest sites worldwide, measured using consistent methods, and examine how these variables are related to consistently determined soil and climatic characteristics. Our pantropical dataset spans intensive monitoring plots in lowland (wet, semi-deciduous, and deciduous) and montane tropical forests in South America, Africa, and Southeast Asia (n = 47). Large spatial variation in fine root dynamics was observed across montane and lowland forest types. In lowland forests, we found a strong positive linear relationship between fine root productivity and sand content, this relationship was even stronger when we considered the fractional allocation of total NPP to fine roots, demonstrating that understanding allocation adds explanatory power to understanding fine root productivity and total NPP. Fine root residence time was a function of multiple factors: soil sand content, soil pH, and maximum water deficit, with longest residence times in acidic, sandy, and water-stressed soils. In tropical montane forests, on the other hand, a different set of relationships prevailed, highlighting the very different nature of montane and lowland forest biomes. Root productivity was a strong positive linear function of mean annual temperature, root residence time was a strong positive function of soil nitrogen content in montane forests, and lastly decreasing soil P content increased allocation of productivity to fine roots. In contrast to the lowlands, environmental conditions were a better predictor for fine root productivity than for fractional allocation of total NPP to fine roots, suggesting that root productivity is a particularly strong driver of NPP allocation in tropical mountain regions. © 2021 The Authors. Global Change Biology published by John Wiley & Sons Ltd. This study is a product of the Global Ecosystem Monitoring network (GEM), Andes Biodiversity and Ecosystem Research Group (ABERG), Amazon Forest Inventory Network (RAINFOR), Stability of Altered Forest Ecosystem (SAFE), and Instituto de Investigaciones de la Amazonia Peruana (IIAP). WHH was funded by Peruvian FONDECYT/CONCYTEC (grant contract number 213-2015-FONDECYT). The GEM network was supported by a European Research Council Advanced Investigator Grant to YM (GEM-TRAITS: 321131) under the European Union's Seventh Framework Programme (FP7/2007-2013). The field data collection was funded NERC Grants NE/D014174/1 and NE/J022616/1 for in Peru, BALI (NE/K016369/1) for work in Malaysia, the Royal Society-Leverhulme Africa Capacity Building Programme for work in Ghana and Gabon and ESPA-ECOLIMITS (NE/1014705/1) in Ghana and Ethiopia. Plot inventories in South America were supported by funding from the US National Science Foundation Long-Term Research in Environmental Biology program (LTREB; DEB 1754647) and the Gordon and Betty Moore Foundation Andes-Amazon Program. GEM data in Gabon were collected under authorization to YM and supported by the Gabon National Parks Agency. Y.M. is supported by the Jackson Foundation. We would like to acknowledge the GEM team across the tropical regions and countries of Bolivia, Brazil, Ghana, Gabon, Ethiopia, Malaysia, and Peru.

It is easy to measure annual growth of a tree stem. It is hard to measure its daily growth. The reason for this difficulty is the microscopic scale and the need to separate processes that simultaneously result in reversible and irreversible stem expansion. Here we present a model that separates reversible from irreversible cell expansion. Our model is novel, because it explains reversible expansion as consequence of longitudinally and, importantly, radially transmitted changes of hydraulic and osmotic pressures in xylem and bark. To capture and quantify these changes, we manipulated daily stem growth by applying a phloem girdle to stems of 9-m tall trees. The model was informed by measurements of radial movement in stem tissues and sap flow before and after and positions below and above the girdle. Additional measurements of whole-crown fluxes of H2O and CO2, leaf water potentials, non-structural carbohydrates and respiration were used to document the physiological impacts of girdling. This work sheds new light on the role of radial transport processes underpinning daily growth of tree stems. The model helps explain diel patterns of stem growth in trees.

21 Pág. Instituto de Ciencias Forestales (ICIFOR), Ecological theory is built on trade-offs, where trait differences among species evolved as adaptations to different environments. Trade-offs are often assumed to be bidirectional, where opposite ends of a gradient in trait values confer advantages in different environments. However, unidirectional benefits could be widespread if extreme trait values confer advantages at one end of an environmental gradient, whereas a wide range of trait values are equally beneficial at the other end. Here, we show that root traits explain species occurrences along broad gradients of temperature and water availability, but model predictions only resembled trade-offs in two out of 24 models. Forest species with low specific root length and high root tissue density (RTD) were more likely to occur in warm climates but species with high specific root length and low RTD were more likely to occur in cold climates. Unidirectional benefits were more prevalent than trade-offs: for example, species with large-diameter roots and high RTD were more commonly associated with dry climates, but species with the opposite trait values were not associated with wet climates. Directional selection for traits consistently occurred in cold or dry climates, whereas a diversity of root trait values were equally viable in warm or wet climates. Explicit integration of unidirectional benefits into ecological theory is needed to advance our understanding of the consequences of trait variation on species responses to environmental change., sPlot was initiated by sDiv and funded by the German Research Foundation (FZT 118) and is now a platform of iDiv. The sRoot workshops and L.M. were also supported by NWO-Vidi grant 864.14.006. C.M.I. and the Fine-Root Ecology Database were supported by the Biological and Environmental Research program in the US Department of Energy’s Office of Science. J.B. was supported by Deutsche Forschungsgemeinschaft (DFG) project 432975993. N.R.G.-R. thanks the Dorothea Schlözer Postdoctoral Programme of the Georg-August-Universität.

Tropical soils contain huge carbon stocks, which climate warming is projected to reduce by stimulating organic matter decomposition, creating a positive feedback that will promote further warming. Models predict that the loss of carbon from warming soils will be mediated by microbial physiology, but no empirical data are available on the response of soil carbon and microbial physiology to warming in tropical forests, which dominate the terrestrial carbon cycle. Here we show that warming caused a considerable loss of soil carbon that was enhanced by associated changes in microbial physiology. By translocating soils across a 3000 m elevation gradient in tropical forest, equivalent to a temperature change of ± 15 °C, we found that soil carbon declined over 5 years by 4% in response to each 1 °C increase in temperature. The total loss of carbon was related to its original quantity and lability, and was enhanced by changes in microbial physiology including increased microbial carbon‐use‐efficiency, shifts in community composition towards microbial taxa associated with warmer temperatures, and increased activity of hydrolytic enzymes. These findings suggest that microbial feedbacks will cause considerable loss of carbon from tropical forest soils in response to predicted climatic warming this century.

Mangroves often allocate a relatively large proportion of their total biomass production to their roots, and the belowground biomass of these forests contributes towards globally significant carbon sinks. However, little information is available on root production in mangroves due to the difficulties in carrying out measurements of belowground processes, particularly if there is regular flooding. In this study, we examined fine and coarse root production in the east coast of the Malaysian Peninsula. Ingrowth cores were used over the course of 17 months. In September 2014, twenty cores were randomly placed in each of five plots. Three cores were collected from each plot (fifteen cores in total), once every three months. Each core was divided into five 10 cm layers and root dry mass was recorded. Standing root biomass was also measured at the time of final collection using an additional 15 cores. There was a seasonal pattern in root production, which peaked in March and December 2015, after and during the monsoon season. Root biomass in the cores peaked at 33.23 ± 6.3 t ha−1 and 21.46 ± 7.3 t ha−1 in March and December respectively. Standing root biomass in February 2016 in the forest was 20.81 ± 2.8 t ha−1. After 17 months, the final root biomass in the cores was 14% less than the standing root biomass. These data suggest surprisingly rapid growth rates and turnover for mangrove roots. Total root biomass significantly increased with root depth and 78% of the roots, in all soil layers, consisted of fine roots (

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 (

Australia is the driest inhabited continent. Annual rainfall is low and is accompanied by marked inter-annual variability, leading to multi-year droughts. Climate change is expected to alter the frequency, magnitude, and intensity of future droughts, with potentially major environmental and socio-economic consequences for Australia. However, Australian vegetation is well adapted to extended dry periods, thus, the likelihood of drought-induced mortality in the future depends both on the severity of future drought events and inherent vegetation resilience. Here, we used the Community Atmosphere Biosphere Land Exchange (CABLE) land surface model, coupled with a stomatal optimisation scheme, to examine the projected impact of future drought for 24 Eucalyptus species. We forced CABLE with future climate from four global climate models (MIROC, ECHAM, CCCMA, and CSIRO) dynamically downscaled by three regional climate models. We separated the impact of climate change (e.g. increasing VPD, precipitation variability) from rising CO2 (increasing water use-efficiency) to provide the first assessment of future drought risk to Australian trees.

Tropical forests, while only occupying 12% to 15% of the Earth's surface, contain about 25% of the world's carbon biomass, with soils representing the second largest reservoir. Yet, recent studies have suggested that, in response to changing environmental conditions, in future decades tropical forests can switch from carbon sinks to carbon sources, with profound implications for the global carbon cycle. Most of these conclusions result from studies in lowland humid forests. However, other tropical forests, such as those occurring in the Andes are also important determinants of regional-to-global biogeochemical functioning, and their sensitivity to future warming has been less studied than in lowland forests. In this study, we explore intra and interspecific thermal sensitivity of soil respiration and its components (autotrophic and heterotrophic) in 15 dominant tree species in the tropical Andes, through an experimental thermosecuence in the Colombian Andes that uses elevation as a proxy for warming. In this thermosequence, a common garden experiment was set up and individuals from 15 dominant species were planted in three sites that represent a temperature gradient: the higher elevation site (2452 masl) corresponds to the base condition; the mid-elevation site (1326 masl) represents a warming of 8°C; and the lower site (575 masl) and it represents a warming of 12°C. Our results indicate consistently higher respiration values with increased temperature both within and between tree species. We used 𝑸10 values (the factor by which soil respiration increases for every 10-degree rise in temperature) to determine the temperature sensitivity of soil respiration. More specifically, for a warming of 5°C there is a temperature coefficient of 𝑸10 = 2 and for a warming of 9°C and there is a temperature coefficient 𝑸10 = 3, this means that for the greater increase temperature the soil respiration can increase faster. Notably, our results show that not all species respond equally to augmented temperatures, highlighting the potential for differential effects of increased temperature and more generally, of environmental change in the compositions and function of these strategic ecosystems. Collectively, our results are relevant for the management and adaptation of ecosystems, particularly tropical Andean forest, and for the refinement of ecological models that support projections of global environmental change and carbon cycle.