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Key message: Annual stem CO2 efflux increases with stem wood production rates and are inhibited by daily moisture stress. Abstract: Tropical forests cycle a large amount of CO2 between the land and atmosphere, with a substantial portion of the return flux due tree respiratory processes. However, in situ estimates of woody tissue respiratory fluxes and carbon use efficiencies (CUEW) and their dependencies on physiological processes including stem wood production (Pw) and transpiration in tropical forests remain scarce. Here, we synthesize monthly Pw and daytime stem CO2 efflux (ES) measurements over 1 year from 80 trees with variable biomass accumulation rates in the central Amazon. On average, carbon flux to woody tissues, expressed in the same stem area normalized units as ES, averaged 0.90 ± 1.2µmolm−2s−1 for Pw, and 0.55 ± 0.33µmolm−2s−1 for daytime ES. A positive linear correlation was found between stem growth rates and stem CO2 efflux, with respiratory carbon loss equivalent to 15 ± 3% of stem carbon accrual. CUEW of stems was non-linearly correlated with growth and was as high as 77–87% for a fast-growing tree. Diurnal measurements of stem CO2 efflux for three individuals showed a daytime reduction of ES by 15–50% during periods of high sap flow and transpiration. The results demonstrate that high daytime ES fluxes are associated with high CUEW during fast tree growth, reaching higher values than previously observed in the Amazon Basin (e.g., maximum CUEW up to 77–87%, versus 30–56%). The observations are consistent with the emerging view that diurnal dynamics of stem water status influences growth processes and associated respiratory metabolism.

Windthrows (trees uprooted and broken by winds) are common across the Amazon. They range in size from single trees to large gaps that lead to changes in forest dynamics, composition, structure, and carbon balance. Yet, the current understanding of the spatial variability of windthrows is limited. By integrating remote sensing data and geospatial analysis, we present the first study to examine the occurrence, area, and direction of windthrows and the control that environmental variables exert on them across the whole Amazon. Windthrows are more frequent and larger in the northwestern Amazon (Peru and Colombia), with the central Amazon (Brazil) being another hot spot of windthrows. The predominant direction of windthrows is westward. Rainfall, surface elevation, and soil characteristics explain the variability (20%–50%) of windthrows but their effects vary regionally. A better understanding of the spatial dynamics of windthrows will improve understanding of the functioning of Amazon forests.

Increasing severity and frequency of drought is predicted for large portions of the terrestrial biosphere, with major impacts already documented in wet tropical forests. Using a 4-year rainfall exclusion experiment in the Daintree Rainforest in northeast Australia, we examined canopy tree responses to reduced precipitation and soil water availability by quantifying seasonal changes in plant hydraulic and carbon traits for 11 tree species between control and drought treatments. Even with reduced soil volumetric water content in the upper 1m of soil in the drought treatment, we found no significant difference between treatments for predawn and midday leaf water potential, photosynthesis, stomatal conductance, foliar stable carbon isotope composition, leaf mass per area, turgor loss point, xylem vessel anatomy, or leaf and stem nonstructural carbohydrates. While empirical measurements of aboveground traits revealed homeostatic maintenance of plant water status and traits in response to reduced soil moisture, modeled belowground dynamics revealed that trees in the drought treatment shifted the depth from which water was acquired to deeper soil layers. These findings reveal that belowground acclimation of tree water uptake depth may buffer tropical rainforests from more severe droughts that may arise in future with climate change.

Deep-water access is arguably the most effective, but under-studied, mechanism that plants employ to survive during drought. Vulnerability to embolism and hydraulic safety margins can predict mortality risk at given levels of dehydration, but deep-water access may delay plant dehydration. Here, we tested the role of deep-water access in enabling survival within a diverse tropical forest community in Panama using a novel data-model approach. We inversely estimated the effective rooting depth (ERD, as the average depth of water extraction), for 29 canopy species by linking diameter growth dynamics (1990-2015) to vapor pressure deficit, water potentials in the whole-soil column, and leaf hydraulic vulnerability curves. We validated ERD estimates against existing isotopic data of potential water-access depths. Across species, deeper ERD was associated with higher maximum stem hydraulic conductivity, greater vulnerability to xylem embolism, narrower safety margins, and lower mortality rates during extreme droughts over 35years (1981-2015) among evergreen species. Species exposure to water stress declined with deeper ERD indicating that trees compensate for water stress-related mortality risk through deep-water access. The role of deep-water access in mitigating mortality of hydraulically-vulnerable trees has important implications for our predictive understanding of forest dynamics under current and future climates.

Elevated atmospheric carbon dioxide (eCO2 ) is predicted to increase growth rates of forest trees. The extent to which increased growth translates to changes in biomass is dependent on the turnover time of the carbon, and thus tree mortality rates. Size- or age-dependent mortality combined with increased growth rates could result in either decreased carbon turnover from a speeding up of tree life cycles, or increased biomass from trees reaching larger sizes, respectively. However, most vegetation models currently lack any representation of size- or age-dependent mortality and the effect of eCO2 on changes in biomass and carbon turnover times is thus a major source of uncertainty in predictions of future vegetation dynamics. Using a reduced-complexity form of the vegetation demographic model the Functionally Assembled Terrestrial Ecosystem Simulator to simulate an idealised tropical forest, we find increases in biomass despite reductions in carbon turnover time in both size- and age-dependent mortality scenarios in response to a hypothetical eCO2 -driven 25% increase in woody net primary productivity (wNPP). Carbon turnover times decreased by 9.6% in size-dependent mortality scenarios due to a speeding up of tree life cycles, but also by 2.0% when mortality was age-dependent, as larger crowns led to increased light competition. Increases in aboveground biomass (AGB) were much larger when mortality was age-dependent (24.3%) compared with size-dependent (13.4%) as trees reached larger sizes before death. In simulations with a constant background mortality rate, carbon turnover time decreased by 2.1% and AGB increased by 24.0%, however, absolute values of AGB and carbon turnover were higher than in either size- or age-dependent mortality scenario. The extent to which AGB increases and carbon turnover decreases will thus depend on the mechanisms of large tree mortality: if increased size itself results in elevated mortality rates, then this could reduce by about half the increase in AGB relative to the increase in wNPP.

Tropical forests absorb large amounts of atmospheric CO2 through photosynthesis, but high surface temperatures suppress this absorption while promoting isoprene emissions. While mechanistic isoprene emission models predict a tight coupling to photosynthetic electron transport (ETR) as a function of temperature, direct field observations of this phenomenon are lacking in the tropics and are necessary to assess the impact of a warming climate on global isoprene emissions. Here we demonstrate that in the early successional species Vismia guianensis in the central Amazon, ETR rates increased with temperature in concert with isoprene emissions, even as stomatal conductance (gs ) and net photosynthetic carbon fixation (Pn ) declined. We observed the highest temperatures of continually increasing isoprene emissions yet reported (50°C). While Pn showed an optimum value of 32.6 ± 0.4°C, isoprene emissions, ETR, and the oxidation state of PSII reaction centers (qL ) increased with leaf temperature with strong linear correlations for ETR (ƿ = 0.98) and qL (ƿ = 0.99) with leaf isoprene emissions. In contrast, other photoprotective mechanisms, such as non-photochemical quenching, were not activated at elevated temperatures. Inhibition of isoprenoid biosynthesis repressed Pn at high temperatures through a mechanism that was independent of stomatal closure. While extreme warming will decrease gs and Pn in tropical species, our observations support a thermal tolerance mechanism where the maintenance of high photosynthetic capacity under extreme warming is assisted by the simultaneous stimulation of ETR and metabolic pathways that consume the direct products of ETR including photorespiration and the biosynthesis of thermoprotective isoprenoids. Our results confirm that models which link isoprene emissions to the rate of ETR hold true in tropical species and provide necessary "ground-truthing" for simulations of the large predicted increases in tropical isoprene emissions with climate warming.

Amazonian droughts are increasing in frequency and severity. However, little is known about how this may influence species-specific vulnerability to drought across different ecosystem types. We measured 16 functional traits for 16 congeneric species from six families and eight genera restricted to floodplain, swamp, white-sand or plateau forests of Central Amazonia. We investigated whether habitat distributions can be explained by species hydraulic strategies, and if habitat specialists differ in their vulnerability to embolism that would make water transport difficult during drought periods. We found strong functional differences among species. Nonflooded species had higher wood specific gravity and lower stomatal density, whereas flooded species had wider vessels, and higher leaf and xylem hydraulic conductivity. The P50 values (water potential at 50% loss of hydraulic conductivity) of nonflooded species were significantly more negative than flooded species. However, we found no differences in hydraulic safety margin among species, suggesting that all trees may be equally likely to experience hydraulic failure during severe droughts. Water availability imposes a strong selection leading to differentiation of plant hydraulic strategies among species and may underlie patterns of adaptive radiation in many tropical tree genera. Our results have important implications for modeling species distribution and resilience under future climate scenarios.

Forest mortality caused by convective storms (windthrow) is a major disturbance in the Amazon. However, the linkage between windthrows at the surface and convective storms in the atmosphere remains unclear. In addition, the current Earth system models (ESMs) lack mechanistic links between convective wind events and tree mortality. Here we find an empirical relationship that maps convective available potential energy, which is well simulated by ESMs, to the spatial pattern of large windthrow events. This relationship builds connections between strong convective storms and forest dynamics in the Amazon. Based on the relationship, our model projects a 51 ± 20% increase in the area favorable to extreme storms, and a 43 ± 17% increase in windthrow density within the Amazon by the end of this century under the high-emission scenario (SSP 585). These results indicate significant changes in tropical forest composition and carbon cycle dynamics under climate change.

Leaf respiration in the dark (Rdark) and light (Rday) is poorly characterized in diverse tropical ecosystems, and little to no information exists on the degree of light suppression in common tree species within the Amazon basin, and their dependences upon plant functional traits and position within the canopy. We quantified Rdark and apparent Rday using the Kok method and measured key leaf traits in 26 tree individuals of different species distributed in three different canopy positions: canopy, lower canopy, and understory. To explore the relationships between the leaf traits we used the standardized major axis (SMA). We found that canopy trees had significantly higher rates of Rdark and Rday than trees in the understory. The difference between Rdark and Rday (the light suppression of respiration) was greatest in the understory (68 ± 9%, 95% CI) and lower canopy (49 ± 9%, 95% CI) when compared to the canopy (37 ± 10%, 95% CI). We also found that Rday was significantly and strongly correlated with Rdark (p < 0.001) for all the canopy positions. Also, leaf mass per area (LMA) and leaf Phosphorus concentration (P) had a significant relationship with Rdark (p < 0.001; p = 0.003), respectively. In addition, a significant relationship was found for LMA in the canopy and lower canopy positions (p = 0.009; p = 0.048) while P was only significant in the canopy (p = 0.044). Finally, no significant relationship was found between Rdark and nitrogen, sugars, and starch. Our results highlight the importance of including representation of the light suppression of leaf respiration in terrestrial biosphere models and also of accounting for vertical gradients within forest canopies and connections with functional traits.

Azteca ants are widely distributed in the neotropics and have been utilized as natural insect repellent for centuries. Azteca oils provide natural defense against herbivores in mutualistic interactions between ants and their host trees. While chemical characterization of oil secretions revealed a composition dominated by iridoids and ketones, the volatile emissions from Azteca ants, and therefore the active gas-phase semiochemicals, remain uncharacterized. In this study, we determined the composition of volatile emissions from a sample of an Azteca ant nest near the Rio Negro in the central Amazon. We found Azteca emissions were composed of a blend of methyl cyclopentyl and methyl cyclopentenyl based volatiles previously identified in Azteca oil extracts as potent alarm pheromones. The ketone 6-methyl-5-hepten-2-one, which also waspreviously identified as a major component of the Azteca oil, was found to be the dominant volatile emitted. For the first time, we report emissions of the highly volatile ketones 2,3-butadione and acetoin from the Azteca nest. Our study has important implications for the better understanding of the ecology and defense strategies of Azteca ants in herbivore defense and provides a base for future commercial applications involving Azteca ant essential oils as natural insect repellents.

Author(s): Holm, JA; Knox, RG; Zhu, Q; Fisher, RA; Koven, CD; Nogueira Lima, AJ; Riley, WJ; Longo, M; Negron-Juarez, RI; de Araujo, AC; Kueppers, LM; Moorcroft, PR; Higuchi, N; Chambers, JQ | Abstract: There is large uncertainty whether Amazon forests will remain a carbon sink as atmospheric CO2 increases. Hence, we simulated an old-growth tropical forest using six versions of four terrestrial models differing in scale of vegetation structure and representation of biogeochemical (BGC) cycling, all driven with CO2 forcing from the preindustrial period to 2100. The models were benchmarked against tree inventory and eddy covariance data from a Brazilian site for present-day predictions. All models predicted positive vegetation growth that outpaced mortality, leading to continual increases in present-day biomass accumulation. Notably, the two vegetation demographic models (VDMs) (ED2 and ELM-FATES) always predicted positive stem diameter growth in all size classes. The field data, however, indicated that a quarter of canopy trees didn't grow over the 15-year period, and while high interannual variation existed, biomass change was near neutral. With a doubling of CO2, three of the four models predicted an appreciable biomass sink (0.77 to 1.24 Mg ha−1 year−1). ELMv1-ECA, the only model used here that includes phosphorus constraints, predicted the lowest biomass sink relative to initial biomass stocks (+21%), lower than the other BGC model, CLM5 (+48%). Models projections differed primarily through variations in nutrient constraints, then carbon allocation, initial biomass, and density-dependent mortality. The VDM's performance was similar or better than the BGC models run in carbon-only mode, suggesting that nutrient competition in VDMs will improve predictions. We demonstrate that VDMs are comparable to nondemographic (i.e., “big-leaf”) models but also include finer scale demography and competition that can be evaluated against field observations.

© 2018 The US and Mexico share a common history in many areas, including language and culture. They face ecological changes due to the increased frequency and severity of droughts and rising energy demands; trends that entail economic costs for both nations and major implications for human wellbeing. We describe an ongoing effort by the Environment Working Group (EWG), created by The University of California's UC-Mexico initiative in 2015, to promote binational research, teaching, and outreach collaborations on the implications of climate change for Mexico and California. We synthesize current knowledge about the most pressing issues related to climate change in the US-Mexico border region and provide examples of cross-border discoveries and research initiatives, highlighting the need to move forward in six broad rubrics. This and similar binational cooperation efforts can lead to improved living standards, generate a collaborative mindset among participating universities, and create an international network to address urgent sustainability challenges affecting both countries.

The fate of tropical forests under climate change is unclear as a result, in part, of the uncertainty in projected changes in precipitation and in the ability of vegetation models to capture the effects of drought-induced mortality on aboveground biomass (AGB). We evaluated the ability of a terrestrial biosphere model with demography and hydrodynamics (Ecosystem Demography, ED2-hydro) to simulate AGB and mortality of four tropical tree plant functional types (PFTs) that operate along light- and water-use axes. Model predictions were compared with observations of canopy trees at Barro Colorado Island (BCI), Panama. We then assessed the implications of eight hypothetical precipitation scenarios, including increased annual precipitation, reduced inter-annual variation, El Nino-related droughts and drier wet or dry seasons, on AGB and functional diversity of the model forest. When forced with observed meteorology, ED2-hydro predictions capture multiple BCI benchmarks. ED2-hydro predicts that AGB will be sustained under lower rainfall via shifts in the functional composition of the forest, except under the drier dry-season scenario. These results support the hypothesis that inter-annual variation in mean and seasonal precipitation promotes the coexistence of functionally diverse PFTs because of the relative differences in mortality rates. If the hydroclimate becomes chronically drier or wetter, functional evenness related to drought tolerance may decline.

Plant-derived latex is widely used in rubber production and plays important roles in ecological processes in the tropics. Although latex from the commercially important tree Hevea brasiliensis regulates latex oxidation resulting in browning, little is known about latex oxidation in highly diverse tropical ecosystems. Here we show that upon physical trunk damage, yellow latex released from the canopy tree Muiratinga (Maquira sclerophylla (Ducke) C.C. Berg) is rapidly and extensively oxidized to a black resin in the presence of air within 15-30 min. In a nitrogen atmosphere, latex oxidation was inhibited, but was immediately activated upon exposure to air. The results suggest the occurrence of O2-dependent oxidative enzymes including polyphenol oxidase (PPO) within the latex of Muiratinga and supports previous findings of a key role of oxidation during latex coagulation. The findings therefore have important implications for understanding the mechanisms of latex coagulation and its ecological role(s) including wound sealing and defense against herbivores.

Metadata describe the ancillary information needed for data preservation and independent interpretation, comparison across heterogeneous datasets, and quality assessment and quality control (QA/QC). Environmental observations are vastly diverse in type and structure, can be taken across a wide range of spatiotemporal scales in a variety of measurement settings and approaches, and saved in multiple formats. Thus, well-organized, consistent metadata are required to produce usable data products from diverse environmental observations collected across field sites. However, existing metadata reporting protocols do not support the complex data synthesis and model-data integration needs of interdisciplinary earth system research. We developed a metadata reporting framework (FRAMES) to enable management and synthesis of observational data that are essential in advancing a predictive understanding of earth systems. FRAMES utilizes best practices for data and metadata organization enabling consistent data reporting and compatibility with a variety of standardized data protocols. We used an iterative scientist-centered design process to develop FRAMES, resulting in a data reporting format that incorporates existing field practices to maximize data-entry efficiency. Thus, FRAMES has a modular organization that streamlines metadata reporting and can be expanded to incorporate additional data types. With FRAMES's multi-scale measurement position hierarchy, data can be reported at observed spatial resolutions and then easily aggregated and linked across measurement types to support model-data integration. FRAMES is in early use by both data originators (persons generating data) and consumers (persons using data and metadata). In this paper, we describe FRAMES, identify lessons learned, and discuss areas of future development.

The Amazon basin has experienced periodic droughts in the past, and intense and frequent droughts are predicted in the future. Landscape heterogeneity could play an important role in how tropical forests respond to drought by influencing water available to plants. Using the one-dimensional ACME Land Model and the three-dimensional ParFlow variably saturated flow model, numerical experiments were performed for a catchment in central Amazon to elucidate processes that influence water available for plant use and provide insights for improving Earth system models. Results from ParFlow show that topography has a dominant influence on groundwater table and runoff through lateral flow. Without any representations of lateral processes, ALM simulates very different seasonal variations in groundwater table and runoff compared to ParFlow even if it is able to reproduce the long-term spatial average groundwater table of ParFlow through simple parameter calibration. In the ParFlow simulations, even in the plateau with much deeper water table depth during the dry season in the drought year of 2005, plant transpiration is not water stressed as the soil saturation is still sufficient for the stomata to be fully open based on the empirical wilting formulation in the models. This finding is insensitive to uncertainty in atmospheric forcing and soil parameters, but the empirical wilting formulation is an important factor that should be addressed using observations and modeling of coupled plant hydraulics-soil hydrology processes in future studies. The results could be applicable to other catchments in the Amazon basin with similar seasonal variability and hydrologic regimes.

In mid-January 2005 a convective squall line traversed 4.5 million km2 of Amazonia from southwest to northeast. As seen in Landsat images, this atypical convective storm left blowdown imprints with diffuse geometry, unlike the fan-shaped wind disturbance of much more frequent east-to-west propagating squall lines. Previous work reported 0.2% of the forest area damaged by this one relatively rare event within one Landsat image and assumed similar disturbance across the entire traverse. We mapped convective wind damage impact to the region in 2005 by identifying large-scale (>4ha) blowdown imprints in 30 Landsat images. The diffuse-type imprints associated with this single squall line contributed up to 60-72% of total 2005 wind-disturbed area detected across the region, but damage was highly concentrated in Central Amazonia. Consequently, the distribution of large wind-damage patches in 2005 across Amazonia was very different from long-term average. Regional distribution of wind-driven tree mortality for smaller patch sizes remains unknown.

Author(s): Negron-Juarez, R; Ferreira, SJF; Mota, MC; Faybishenko, B; Monteiro, MTF; Candido, LA; Ribeiro, RP; de Oliveira, RC; de Araujo, AC; Warren, JM; Newman, BD; Gimenez, BO; Varadharajan, C; Agarwal, D; Borma, L; Tomasella, J; Higuchi, N; Chambers, JQ | Abstract: Soil moisture plays a key role in hydrological, biogeochemical, and energy budgets of terrestrial ecosystems. Accurate soil moisture measurements in remote ecosystems such as the Amazon are difficult and limited because of logistical constraints. Time domain reflectometry (TDR) sensors are widely used to monitor soil moisture and require calibration to convert the TDR's dielectric permittivity measurement (Ka) to volumetric water content (θv). In this study, our objectives were to develop a field-based calibration of TDR sensors in an old-growth upland forest in the central Amazon, to evaluate the performance of the calibration, and then to apply the calibration to determine the dynamics of soil moisture content within a 14.2-m-deep vertical soil profile. Depth-specific TDR calibration using local soils in a controlled laboratory setting yielded a novel Ka–θv third-degree polynomial calibration. The sensors were later installed to their specific calibration depth in a 14.2-m pit. The widely used Ka–θv relationship (Topp model) underestimated the site-specific θv by 22–42%, indicating significant error in the model when applied to these well-structured, clay-rich tropical forest soils. The calibrated wet- and dry-season θv data showed a variety of depth and temporal variations highlighting the importance of soil textural differentiation, root uptake depths, as well as event to seasonal precipitation effects. Data such as these are greatly needed for improving our understanding of ecohydrological processes within tropical forests and for improving models of these systems in the face of changing environmental conditions.

Tree growth and survival differ strongly between canopy trees (those directly exposed to overhead light), and understory trees. However, the structural complexity of many tropical forests makes it difficult to determine canopy positions. The integration of remote sensing and ground-based data enables this determination and measurements of how canopy and understory trees differ in structure and dynamics. Here we analyzed 2 cm resolution RGB imagery collected by a Remotely Piloted Aircraft System (RPAS), also known as drone, together with two decades of bi-annual tree censuses for 2 ha of old growth forest in the Central Amazon. We delineated all crowns visible in the imagery and linked each crown to a tagged stem through field work. Canopy trees constituted 40% of the 1244 inventoried trees with diameter at breast height (DBH) > 10 cm, and accounted for ~70% of aboveground carbon stocks and wood productivity. The probability of being in the canopy increased logistically with tree diameter, passing through 50% at 23.5 cm DBH. Diameter growth was on average twice as large in canopy trees as in understory trees. Growth rates were unrelated to diameter in canopy trees and positively related to diameter in understory trees, consistent with the idea that light availability increases with diameter in the understory but not the canopy. The whole stand size distribution was best fit by a Weibull distribution, whereas the separate size distributions of understory trees or canopy trees > 25 cm DBH were equally well fit by exponential and Weibull distributions, consistent with mechanistic forest models. The identification and field mapping of crowns seen in a high resolution orthomosaic revealed new patterns in the structure and dynamics of trees of canopy vs. understory at this site, demonstrating the value of traditional tree censuses with drone remote sensing.

Forest disturbance and regrowth are key processes in forest dynamics but detailed information of these processes is difficult to obtain in remote forests as the Amazon. We used chronosequences of Landsat satellite imagery to determine the sensitivity of surface reflectance from all spectral bands to windthrow, clearcutting, and burning and their successional pathways of forest regrowth in the Central Amazon. We also assess whether the forest demography model Functionally Assembled Terrestrial Ecosystem Simulator (FATES) implemented in the Energy Exascale Earth System Model (E3SM) Land Model (ELM), ELM-FATES, accurately represents the changes for windthrow and clearcut. The results show that all spectral bands from Landsat satellite were sensitive to the disturbances but after 3 to 6 years only the Near Infrared (NIR) band had significant changes associated with the successional pathways of forest regrowth for all the disturbances considered. In general, the NIR decreased immediately after disturbance, increased to maximum values with the establishment of pioneers and early-successional tree species, and then decreased slowly and almost linearly to pre-disturbance conditions with the dynamics of forest succession. Statistical methods predict that NIR will return to pre-disturbance values in about 39 years (consistent with observational data of biomass regrowth following windthrows), and 36 and 56 years for clearcut and burning. The NIR captured the observed successional pathways of forest regrowth after clearcut and burning that diverge through time. ELM-FATES predicted higher peaks of initial forest responses (e.g., biomass, stem density) after clearcuts than after windthrows, similar to the changes in NIR. However, ELM-FATES predicted a faster recovery of forest structure and canopy-coverage back to pre-disturbance conditions for windthrows compared to clearcuts. The similarity of ELM-FATES predictions of regrowth patterns after windthrow and clearcut to those of the NIR results suggest that the dynamics of forest regrowth for these disturbances are represented with appropriate fidelity within ELM-FATES and useful as a benchmarking tool.

Tropical forests are acknowledged to be the largest global source of isoprene (C5H8) and monoterpenes (C10H16) emissions, with current synthesis studies suggesting few tropical species emit isoprenoids (20-38%) and do so with highly variable emission capacities, including within the same genera. This apparent lack of a clear phylogenetic thread has created difficulties both in linking isoprenoid function with evolution and for the development of accurate biosphere-atmosphere models. Here, we present a systematic emission study of "hyperdominant" tree species in the Amazon Basin. Across 162 individuals, distributed among 25 botanical families and 113 species, isoprenoid emissions were widespread among both early and late successional species (isoprene: 61.9% of the species; monoterpenes: 15.0%; both isoprene and monoterpenes: 9.7%). The hyperdominant species (69) across the top five most abundant genera, which make up about 50% of all individuals in the Basin, had a similar abundance of isoprenoid emitters (isoprene: 63.8%; monoterpenes: 17.4%; both 11.6%). Among the abundant genera, only Pouteria had a low frequency of isoprene emitting species (15.8% of 19 species). In contrast, Protium, Licania, Inga, and Eschweilera were rich in isoprene emitting species (83.3% of 12 species, 61.1% of 18 species, 100% of 8 species, and 100% of 12 species, respectively). Light response curves of individuals in each of the five genera showed light-dependent, photosynthesis-linked emission rates of isoprene and monoterpenes. Importantly, in every genus, we observed species with light-dependent isoprene emissions together with monoterpenes including β-ocimene. These observations support the emerging view of the evolution of isoprene synthases from β-ocimene synthases. Our results have important implications for understanding isoprenoid function-evolution relationships and the development of more accurate Earth System Models.

International audience; Transpiration in humid tropical forests modulates the global water cycle and is a key driver of climate regulation. Yet, our understanding of how tropical trees regulate sap flux in response to climate variability remains elusive. With a progressively warming climate, atmospheric evaporative demand [i.e., vapor pressure deficit (VPD)] will be increasingly important for plant functioning, becoming the major control of plant water use in the twenty-first century. Using measurements in 34 tree species at seven sites across a precipitation gradient in the neotropics, we determined how the maximum sap flux velocity (vmax) and the VPD threshold at which vmax is reached (VPDmax) vary with precipitation regime [mean annual precipitation (MAP); seasonal drought intensity (PDRY)] and two functional traits related to foliar and wood economics spectra [leaf mass per area (LMA); wood specific gravity (WSG)]. We show that, even though vmax is highly variable within sites, it follows a negative trend in response to increasing MAP and PDRY across sites. LMA and WSG exerted little effect on vmax and VPDmax, suggesting that these widely used functional traits provide limited explanatory power of dynamic plant responses to environmental variation within hyper-diverse forests. This study demonstrates that long-term precipitation plays an important role in the sap flux response of humid tropical forests to VPD. Our findings suggest that under higher evaporative demand, trees growing in wetter environments in humid tropical regions may be subjected to reduced water exchange with the atmosphere relative to trees growing in drier climates.

The Camp Fire rapidly spread across a landscape in Butte County, California, toward the city of Paradise in the early morning hours of 8 November 2018. Here we provide a set of initial tools and analyses that are useful to a variety of stakeholders, including: (1) a visualization app for GOES 16 data and the surrounding landscape showing the rapid spread of the fire from 6:37-10:47 a.m. local time; (2) processed Landsat 8 images for before, during, and after the fire, along with a tool for visualizing regional impacts; (3) a timeline of fire spread from ignition over the first four hours; and (4) a description of a potential early warning app that could make use of existing data, visualization, and analysis tools, to provide additional information for effective evacuation of communities threatened by rapidly moving wildfires. Using these tools we estimate that, over the first hour, the Camp Fire was consuming ~200 ha/minute of vegetation with a linear spread rate of 14 km over the fire’s first 25 minutes, or ~33km/hr. We briefly discuss broader use of remote sensing and geospatial analysis for fire research and risk management.

International audience; The temperature response of photosynthesis is one of the key factors determining predicted responses to warming in global vegetation models (GVMs). The response may vary geographically, owing to genetic adaptation to climate, and temporally, as a result of acclimation to changes in ambient temperature. Our goal was to develop a robust quantitative global model representing acclimation and adaptation of photosynthetic temperature responses. We quantified and modelled key mechanisms responsible for photosynthetic temperature acclimation and adaptation using a global dataset of photosynthetic CO2 response curves, including data from 141 C3 species from tropical rainforest to Arctic tundra. We separated temperature acclimation and adaptation processes by considering seasonal and common‐garden datasets, respectively. The observed global variation in the temperature optimum of photosynthesis was primarily explained by biochemical limitations to photosynthesis, rather than stomatal conductance or respiration. We found acclimation to growth temperature to be a stronger driver of this variation than adaptation to temperature at climate of origin. We developed a summary model to represent photosynthetic temperature responses and showed that it predicted the observed global variation in optimal temperatures with high accuracy. This novel algorithm should enable improved prediction of the function of global ecosystems in a warming climate.

Tropical forests absorb large amounts of atmospheric CO2 through photosynthesis but elevated temperatures suppress this absorption and promote monoterpene emissions. Using 13 CO2 labeling, here we show that monoterpene emissions from tropical leaves derive from recent photosynthesis and demonstrate distinct temperature optima for five groups (Groups 1-5), potentially corresponding to different enzymatic temperature-dependent reaction mechanisms within β-ocimene synthases. As diurnal and seasonal leaf temperatures increased during the Amazonian 2015 El Nino event, leaf and landscape monoterpene emissions showed strong linear enrichments of β-ocimenes (+4.4% °C-1 ) at the expense of other monoterpene isomers. The observed inverse temperature response of α-pinene (-0.8% °C-1 ), typically assumed to be the dominant monoterpene with moderate reactivity, was not accurately simulated by current global emission models. Given that β-ocimenes are highly reactive with respect to both atmospheric and biological oxidants, the results suggest that highly reactive β-ocimenes may play important roles in the thermotolerance of photosynthesis by functioning as effective antioxidants within plants and as efficient atmospheric precursors of secondary organic aerosols. Thus, monoterpene composition may represent a new sensitive 'thermometer' of leaf oxidative stress and atmospheric reactivity, and therefore a new tool in future studies of warming impacts on tropical biosphere-atmosphere carbon-cycle feedbacks.

High descending winds generated by convective storms are a frequent and a major source of tree mortality disturbance events in the Amazon, affecting forest structure and diversity across a variety of scales, and more frequently observed in western and central portions of the basin. Soil texture in the Central Amazon also varies significantly with elevation along a topographic gradient, with decreasing clay content on plateaus, slopes and valleys respectively. In this study we investigated the critical turning moments (Mcrit - rotational force at the moment of tree failure, an indicator of tree stability or wind resistance) of 60 trees, ranging from 19.0 to 41.1 cm in diameter at breast height (DBH) and located in different topographic positions, and for different species, using a cable-winch load-cell system. Our approach used torque as a measure of tree failure to the point of snapping or uprooting. This approach provides a better understanding of the mechanical forces required to topple trees in tropical forests, and will inform models of wind throw disturbance. Across the topographic positions, size controlled variation in Mcrit was quantified for cardeiro (Scleronema mincranthum (Ducke) Ducke), mata-mata (Eschweilera spp.), and a random selection of trees from 19 other species. Our analysis of Mcrit revealed that tree resistance to failure increased with size (DBH and ABG) and differed among species. No effects of topography or failure mode were found for the species either separately or pooled. For the random species, total variance in Mcrit explained by tree size metrics increased from an R2 of 0.49 for DBH alone, to 0.68 when both DBH and stem fresh wood density (SWD) were included in a multiple regression model. This mechanistic approach allows the comparison of tree vulnerability induced by wind damage across ecosystems, and facilitates the use of forest structural information in ecosystem models that include variable resistance of trees to mortality inducing factors. Our results indicate that observed topographic differences in windthrow vulnerability are likely due to elevational differences in wind velocities, rather than by differences in soil-related factors that might effect Mcrit.

Assessing long-term effects of hurricane damage in bottomland hardwood forests is important to detect any permanent, long lasting changes to the forest. Two 75 × 75 m plots were established in a Louisiana bottomland hardwood forest in 2004 and all adult trees were measured. The plots were resurveyed in 2006 after Hurricane Katrina passed over the plots in 2005, and in 2011, to gain an understanding of mortality over time. Species composition, mortality and biomass change were assessed over the study period from 2004 to 2011. Sweetgum, water oak, and laurel oak were the most important overstory species in 2004, and American holly and American hornbeam were the most important understory species. In the more damaged plot, there was a shift in dominant species as 63 new Chinese tallow individuals recruited into the adult class (>10 cm) between 2004 and 2011. Chinese tallow is an invasive tree species that often out-competes native species in bottomland hardwood forests. Annual mortality in the bottomland hardwood forest plots between 2004 and 2011 was 6% per year, 11% per year from 2004 to 2006 (representing direct hurricane mortality), and 5% between 2006 and 2011 (delayed tree mortality). Approximately 53% of the total biomass (188,000 kg) was lost between 2004 and 2011. A plot in a cypress tupelo forest was added in 2006 and very little damage or mortality was observed. This study revealed that delayed mortality to hurricane-damaged trees is a significant factor in the long-term dynamics of bottomland hardwood forests and represents an amplification of the effects of the hurricane over time. The fact that direct and delayed mortality is different by species indicates that the measurement only of direct mortality can lead to false conclusions about which species are resistant to hurricanes. Hurricane damage opened up new habitat for invasion by Chinese tallow which grew prolifically in highly damaged, low elevation, wet areas, indicating that large disturbances are an important factor in accelerating the population expansion of this invasive species. The high mortality and low recruitment of some species into the sapling and adult layers and the corresponding expansion of Chinese tallow indicates that species composition will differ from pre-hurricane composition for some time in the future.

Widely recognized as one of the worst natural disaster in Puerto Rico's history, hurricane Maria made landfall on September 20, 2017 in southeast Puerto Rico as a high-end category 4 hurricane on the Saffir-Simpson scale causing widespread destruction, fatalities and forest disturbance. This study focused on hurricane Maria's effect on Puerto Rico's forests as well as the effect of landform and forest characteristics on observed disturbance patterns. We used Google Earth Engine (GEE) to assess the severity of forest disturbance using a disturbance metric based on Landsat 8 satellite data composites with pre and post-hurricane Maria. Forest structure, tree phenology characteristics, and landforms were obtained from satellite data products, including digital elevation model and global forest canopy height. Our analyses showed that forest structure, and characteristics such as forest age and forest type affected patterns of forest disturbance. Among forest types, highest disturbance values were found in sierra palm, transitional, and tall cloud forests; seasonal evergreen forests with coconut palm; and mangrove forests. For landforms, greatest disturbance metrics was found at high elevations, steeper slopes, and windward surfaces. As expected, high levels of disturbance were also found close to the hurricane track, with disturbance less severe as hurricane Maria moved inland. Results demonstrated that forest and landform characteristics accounted for 34% of the variation in spatial forest spectral disturbance patterns. This study demonstrated an informative regional approach, combining remote sensing with statistical analyses to investigate factors that result in variability in hurricane effects on forest ecosystems.

Recent research in the central Amazon suggests that wind is a major agent of disturbance, however, a mechanistic understanding of how wind may lead to tree mortality in Amazonian forests remains unclear. Here we estimated wind speeds necessary to topple central Amazon trees by linking both static and dynamic versions of two wind speed estimation methods (four methods total) to field data on tree failure derived from a static winching study. Static versions of these methods assumed invariant wind characteristics as more trees failed, while dynamic versions updated tree spacing, leaf area index and wind profiles progressively after each tree failure. First, we used a profile method which estimates wind force on individual trees by segments. We calculated drag on each segment and converted drag into basal turning moment, and compared the summed turning moments to the critical turning moment measured in the winching study. Estimated critical wind speeds from the static profile method varied greatly, from 10.75 m s−1 to >120.0 m s−1 with a mean of 45.70 m s−1. Critical wind speeds estimated with static approaches decreased with tree size but were not significantly different between two focal genera. Primary drivers of variation in critical wind speed were tree height and crown size. Second, we used the turning moment coefficient method of Hale, S.E., Gardiner, B., Peace, A., Nicoll, B., Taylor, P. and Pizzirani, S. 2015 Comparison and validation of three versions of a forest wind risk model. Environ. Model. Softw.68, 27–41. doi:10.1016/j.envsoft.2015.01.016.; the static version of this method yielded less-variable estimates, ranging from 18.98 to 52.01 m s−1, with a mean of 30.88 m s−1. Notably, the two static methods for estimating critical wind speeds differed in the trees they identified as having the highest and lowest critical wind speeds. Dynamic variants of the above two methods produced greatly reduced ranges in CWS estimates for our study trees, because after the early tree failures, remaining trees were subject to greater wind penetration into the stand and thus greater loading for a given above-canopy wind speed. CWS estimated with dynamic approaches differed significantly between the focal taxa. Nevertheless, both estimates suggest that wind speeds commonly observed during Amazon storms are sufficient to produce widespread tree damage and mortality.

Current climate change scenarios indicate warmer temperatures and the potential for more extreme droughts in the tropics, such that a mechanistic understanding of the water cycle from individual trees to landscapes is needed to adequately predict future changes in forest structure and function. In this study, we contrasted physiological responses of tropical trees during a normal dry season with the extreme dry season due to the 2015–2016 El Niño-Southern Oscillation (ENSO) event. We quantified high resolution temporal dynamics of sap velocity (Vs), stomatal conductance (gs) and leaf water potential (ΨL) of multiple canopy trees, and their correlations with leaf temperature (Tleaf) and environmental conditions [direct solar radiation, air temperature (Tair) and vapor pressure deficit (VPD)]. The experiment leveraged canopy access towers to measure adjacent trees at the ZF2 and Tapajós tropical forest research (near the cities of Manaus and Santarém). The temporal difference between the peak of gs (late morning) and the peak of VPD (early afternoon) is one of the major regulators of sap velocity hysteresis patterns. Sap velocity displayed species-specific diurnal hysteresis patterns reflected by changes in Tleaf. In the morning, Tleaf and sap velocity displayed a sigmoidal relationship. In the afternoon, stomatal conductance declined as Tleaf approached a daily peak, allowing ΨL to begin recovery, while sap velocity declined with an exponential relationship with Tleaf. In Manaus, hysteresis indices of the variables Tleaf-Tair and ΨL-Tleaf were calculated for different species and a significant difference (p < 0.01, α = 0.05) was observed when the 2015 dry season (ENSO period) was compared with the 2017 dry season (“control scenario”). In some days during the 2015 ENSO event, Tleaf approached 40°C for all studied species and the differences between Tleaf and Tair reached as high at 8°C (average difference: 1.65 ± 1.07°C). Generally, Tleaf was higher than Tair during the middle morning to early afternoon, and lower than Tair during the early morning, late afternoon and night. Our results support the hypothesis that partial stomatal closure allows for a recovery in ΨL during the afternoon period giving an observed counterclockwise hysteresis pattern between ΨL and Tleaf. © 2019 Gimenez, Jardine, Higuchi, Negrón-Juárez, Sampaio-Filho, Cobello, Fontes, Dawson, Varadharajan, Christianson, Spanner, Araújo, Warren, Newman, Holm, Koven, McDowell and Chambers.

Volatile terpenoid resins represent a diverse group of plant defense chemicals involved in defense against herbivory, abiotic stress, and communication. However, their composition in tropical forests remains poorly characterized. As a part of tree identification, the ‘smell’ of damaged trunks is widely used, but is highly subjective. Here, we analyzed trunk volatile monoterpene emissions from 15 species of the genus Protium in the central Amazon. By normalizing the abundances of 28 monoterpenes, 9 monoterpene ‘fingerprint’ patterns emerged, characterized by a distinct dominant monoterpene. While 4 of the ‘fingerprint’ patterns were composed of multiple species, 5 were composed of a single species. Moreover, among individuals of the same species, 6 species had a single ‘fingerprint’ pattern, while 9 species had two or more ‘fingerprint’ patterns among individuals. A comparison of ‘fingerprints’ between 2015 and 2017 from 15 individuals generally showed excellent agreement, demonstrating a strong dependence on species identity, but not time of collection. The results are consistent with a previous study that found multiple divergent copies of monoterpene synthase enzymes in Protium. We conclude that the monoterpene ‘fingerprint’ database has important implications for constraining Protium species identification and phylogenetic relationships and enhancing understanding of physiological and ecological functions of resins and their potential commercial applications. © 2019 The Authors

© 2018 Elsevier B.V. The identification of tree species in the field is often a subjective process and misidentifications cause many problems for forest management in the Amazon Forest. Near infrared spectra from dried leaves of herbarium specimens are able to distinguish species in tropical forests. However, toolsto improve species identification directly in the field are needed. In this study, we tested whether spectral reflectance of bark tissues (rhytidome and phloem) collected with a portable spectrometer in the field can be used for the discrimination of tree species. Spectral data was collected for 254 trees of 8 families, 10 genera and 11 species from terra firme forests in Central Amazon with an ASD field spectrometer. Data consisted of reflectance values within 350–2500 nm wavelengths. We compared the rate of correct species recognition for different datasets using linear discriminant models. The rate of correct species assignment using this technique was 98% when using spectra from the inner bark (phloem) and 94% with outer bark (rhytidome) spectra. We suggest that the application of this technique can improve the quality of species identification directly during field inventories, fostering better forest management practices.

Uncertainties surrounding vegetation response to increased disturbance rates associated with climate change remains a major global change issue for Amazon forests. Additionally, turnover rates computed as the average of mortality and recruitment rates in the Western Amazon basin are doubled when compared to the Central Amazon, and notable gradients currently exist in specific wood density and aboveground biomass (AGB) between these two regions. This study investigates the extent to which the variation in disturbance regimes contributes to these regional gradients. To address these issues, we evaluated disturbance-recovery processes under two scenarios of increased disturbance rates in a complex Central Amazon forest using first ZELIG-TROP, a dynamic vegetation gap model which we calibrated using long-term inventory data, and second using the Community Land Model (CLM), a global land surface model that is part of the Community Earth System Model (CESM). Upon doubling the mortality rate in the Central Amazon to mirror the natural disturbance regime in the Western Amazon of ∼2% mortality, at steady-state, AGB significantly decreased by 41.9% and there was no significant difference between the modeled AGB of 104 Mg C ha−1 and empirical AGB from the western Amazon datasets of 107 Mg C ha−1. We confirm that increases in natural disturbance rates in the Central Amazon will result in terrestrial carbon loss associated with higher turnover. However, different processes were responsible for the reductions in AGB between the models and empirical datasets. We observed that with increased turnover, the subsequent decrease in wood density drives the reduction in AGB in empirical datasets. However, decrease in stand basal area was the driver of the drop in AGB in ZELIG-TROP, and decreased leaf area index (LAI) was the driver in CLM. Further comparisons found that stem density, specific wood density, and basal area growth rates differed between the two Amazonian regions. This suggests that: (1) the variability between regions cannot be entirely explained by the variability in disturbance regime, but rather potentially sensitive to intrinsic environmental factors; or (2) the models are not accurately simulating all forest characteristics in response to increased disturbances. Last, to help quantify the impacts of increased disturbances on climate and the earth system, we evaluated the fidelity of tree mortality and disturbance in a global land surface model: CLM. For a 100% increase in annual mortality rate, both ZELIG-TROP and CLM were in close agreement with each other and predicted a net carbon loss of 41.9 and 49.9%, respectively, with an insignificant effect on aboveground net primary productivity (ANPP). Likewise, a 20% increase in mortality every 50 years (i.e. periodic disturbance treatment) resulted in a reciprocal biomass loss of 18.3 and 18.7% in ZELIG-TROP and CLM, respectively.

© 2018 Elsevier B.V. Tree mortality is a critical process in forest ecosystems, as it influences floristic composition, structure, dynamics, carbon storage, and forest nutrient cycling. However, the mechanisms behind tree death in tropical regions are still poorly characterized. This lack of information is mainly because tree mortality data come from long-interval inventories and studies that measured tree death seasonally are scarce. Here we offer novel insights into the power of fine temporal scale observations and we use a natural history approach to understand the processes and mechanism of tree mortality. We monitored tree mortality every month during one year, in 10 ha of terra-firme forest. To determine the pathways of mortality, we considered the state of the tree at the start of the investigation and the pre and post-mortem characteristics. From November 2010 to October 2011, 67 out of 5808 trees died. Despite the 2010 drought, mortality was highly correlated with monthly rainfall (r = 0.85). In total, six pathways of mortality were assessed. Storms were the main cause of mortality, killing 45% of all dead trees, followed by Biotic/abiotic factors accounting for 30% of tree mortality. The high mortality registered in the rainy season was mostly (78%) due to healthy trees dying uprooted or snapped. Finally, we would benefit from studies that assess mortality on a monthly basis and in combination with quantitative long-term data, we can substantially improve our understanding of the mechanisms behind tree death in the tropics.

How plants respond physiologically to leaf warming and low water availability may determine how they will perform under future climate change. In 2015–2016, an unprecedented drought occurred across Amazonia with record-breaking high temperatures and low soil moisture, offering a unique opportunity to evaluate the performances of Amazonian trees to a severe climatic event. We quantified the responses of leaf water potential, sap velocity, whole-tree hydraulic conductance ( K wt ), turgor loss and xylem embolism, during and after the 2015–2016 El Niño for five canopy-tree species. Leaf/xylem safety margins (SMs), sap velocity and K wt showed a sharp drop during warm periods. SMs were negatively correlated with vapour pressure deficit, but had no significant relationship with soil water storage. Based on our calculations of canopy stomatal and xylem resistances, the decrease in sap velocity and K wt was due to a combination of xylem cavitation and stomatal closure. Our results suggest that warm droughts greatly amplify the degree of trees' physiological stress and can lead to mortality. Given the extreme nature of the 2015–2016 El Niño and that temperatures are predicted to increase, this work can serve as a case study of the possible impact climate warming can have on tropical trees. 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’.

No claim to original US government works New Phytologist © 2018 New Phytologist Trust Tree mortality rates appear to be increasing in moist tropical forests (MTFs) with significant carbon cycle consequences. Here, we review the state of knowledge regarding MTF tree mortality, create a conceptual framework with testable hypotheses regarding the drivers, mechanisms and interactions that may underlie increasing MTF mortality rates, and identify the next steps for improved understanding and reduced prediction. Increasing mortality rates are associated with rising temperature and vapor pressure deficit, liana abundance, drought, wind events, fire and, possibly, CO2 fertilization-induced increases in stand thinning or acceleration of trees reaching larger, more vulnerable heights. The majority of these mortality drivers may kill trees in part through carbon starvation and hydraulic failure. The relative importance of each driver is unknown. High species diversity may buffer MTFs against large-scale mortality events, but recent and expected trends in mortality drivers give reason for concern regarding increasing mortality within MTFs. Models of tropical tree mortality are advancing the representation of hydraulics, carbon and demography, but require more empirical knowledge regarding the most common drivers and their subsequent mechanisms. We outline critical datasets and model developments required to test hypotheses regarding the underlying causes of increasing MTF mortality rates, and improve prediction of future mortality under climate change.

Nonstructural carbohydrates (NSCs) are essential for maintenance of plant metabolism and may be sensitive to short- and long-term climatic variation. NSC variation in moist tropical forests has rarely been studied, so regulation of NSCs in these systems is poorly understood. We measured foliar and branch NSC content in 23 tree species at three sites located across a large precipitation gradient in Panama during the 2015-2016 El Niño to examine how short- and long-term climatic variation impact carbohydrate dynamics. There was no significant difference in total NSCs as the drought progressed (leaf P=0.32, branch P=0.30) nor across the rainfall gradient (leaf P=0.91, branch P=0.96). Foliar soluble sugars decreased while starch increased over the duration of the dry period, suggesting greater partitioning of NSCs to storage than metabolism or transport as drought progressed. There was a large variation across species at all sites, but total foliar NSCs were positively correlated with leaf mass per area, whereas branch sugars were positively related to leaf temperature and negatively correlated with daily photosynthesis and wood density. The NSC homoeostasis across a wide range of conditions suggests that NSCs are an allocation priority in moist tropical forests.