Predicting responses of tropical forests to climate change-type drought is challenging because of high species diversity. Detailed characterization of tropical tree hydraulic physiology is necessary to evaluate community drought vulnerability and improve model parameterization. Here, we measured xylem hydraulic conductivity (hydraulic efficiency), xylem vulnerability curves (hydraulic safety), sapwood pressure-volume curves (drought avoidance) and wood density on emergent branches of 14 common species of Eastern Amazonian canopy trees in Paracou, French Guiana across species with the densest and lightest wood in the plot. Our objectives were to evaluate relationships among hydraulic traits to identify strategies and test the ability of easy-to-measure traits as proxies for hard-to-measure hydraulic traits. Xylem efficiency was related to capacitance, sapwood water content and turgor loss point, and other drought avoidance traits, but not to xylem safety (P50 ). Wood density was correlated (r = -0.57 to -0.97) with sapwood pressure-volume traits, forming an axis of hydraulic strategy variation. In contrast to drier sites where hydraulic safety plays a greater role, tropical trees in this humid tropical site varied along an axis with low wood density, high xylem efficiency and high capacitance at one end of the spectrum, and high wood density and low turgor loss point at the other.

Systematic biases in eddy covariance measurements of net ecosystem-atmosphere carbon dioxide exchange (NEE) are ubiquitous in forests when turbulence is low at night. We propose an alternative to the conventional bias correction, the friction velocity (u*) filter, by hypothesizing that these biases have two separate, concurrent causes: (1) a subcanopy CO₂ storage pool that eludes typical storage measurements, creating a turbulence-dependent bias, and (2) advective divergence loss of CO₂, creating a turbulence-independent bias. We correct for (1) using a simple parametric model of missing storage (MS). Prior experiments have inferred (2) directly from atmospheric measurements (DRAINO). For sites at which DRAINO experiments have not been performed or are infeasible, we estimate (2) empirically using a PAR-extrapolated advective respiration loss (PEARL) approach. We compare u* filter estimates of advection and NEE to MS-PEARL estimates at one temperate forest and two tropical forest sites.We find that for tropical forests, u* filters can produce a range of extreme NEE estimates, from long-term forest carbon emission to sequestration, that diverge from independent assessments and are not physically sustainable. Our MS model eliminates the dependence of nighttime NEE on u*, consistent with findings from DRAINO studies that nighttime advective losses of CO₂ are often not dependent on the strength of turbulence. Our PEARL estimates of mean advective loss agree with available DRAINO measurements. The MS-PEARL correction to long-term NEE produces better agreement with forest inventories at all three sites. Moreover, the correction retains all nighttime eddy covariance data and is therefore more widely applicable than the u* filter approach, which rejects substantial nighttime data—up to 93% at one of the tropical sites. The full MS-PEARL NEE correction is therefore an equally defensible and more practical alternative to the u* filter, but leads to different conclusions about the resulting carbon balance. Our results therefore highlight the need to investigate which approach’s underlying hypotheses are more physically realistic.

There is mounting empirical evidence that lianas affect the carbon cycle of tropical forests. However, no single vegetation model takes into account this growth form, although such efforts could greatly improve the predictions of carbon dynamics in tropical forests. In this study, we incorporated a novel mechanistic representation of lianas in a dynamic global vegetation model (the Ecosystem Demography Model). We developed a liana-specific plant functional type and mechanisms representing liana-tree interactions (such as light competition, liana-specific allometries, and attachment to host trees) and parameterized them according to a comprehensive literature meta-analysis. We tested the model for an old-growth forest (Paracou, French Guiana) and a secondary forest (Gigante Peninsula, Panama). The resulting model simulations captured many features of the two forests characterized by different levels of liana infestation as revealed by a systematic comparison of the model outputs with empirical data, including local census data from forest inventories, eddy flux tower data, and terrestrial laser scanner-derived forest vertical structure. The inclusion of lianas in the simulations reduced the secondary forest net productivity by up to 0.46 t_C  ha^-1  year^-1 , which corresponds to a limited relative reduction of 2.6% in comparison with a reference simulation without lianas. However, this resulted in significantly reduced accumulated above-ground biomass after 70 years of regrowth by up to 20 t_C /ha (19% of the reference simulation). Ultimately, the simulated negative impact of lianas on the total biomass was almost completely cancelled out when the forest reached an old-growth successional stage. Our findings suggest that lianas negatively influence the forest potential carbon sink strength, especially for young, disturbed, liana-rich sites. In light of the critical role that lianas play in the profound changes currently experienced by tropical forests, this new model provides a robust numerical tool to forecast the impact of lianas on tropical forest carbon sinks.

Selective logging, fragmentation, and understory fires directly degrade forest structure and composition. However, studies addressing the effects of forest degradation on carbon, water, and energy cycles are scarce. Here, we integrate field observations and high-resolution remote sensing from airborne lidar to provide realistic initial conditions to the Ecosystem Demography Model (ED-2.2) and investigate how disturbances from forest degradation affect gross primary production (GPP), evapotranspiration (ET), and sensible heat flux (H). We used forest structural information retrieved from airborne lidar samples (13,500 ha) and calibrated with 817 inventory plots (0.25 ha) across precipitation and degradation gradients in the eastern Amazon as initial conditions to ED-2.2 model. Our results show that the magnitude and seasonality of fluxes were modulated by changes in forest structure caused by degradation. During the dry season and under typical conditions, severely degraded forests (biomass loss ≥66%) experienced water stress with declines in ET (up to 34%) and GPP (up to 35%) and increases of H (up to 43%) and daily mean ground temperatures (up to 6.5°C) relative to intact forests. In contrast, the relative impact of forest degradation on energy, water, and carbon cycles markedly diminishes under extreme, multiyear droughts, as a consequence of severe stress experienced by intact forests. Our results highlight that the water and energy cycles in the Amazon are driven by not only climate and deforestation but also the past disturbance and changes of forest structure from degradation, suggesting a much broader influence of human land use activities on the tropical ecosystems.

Despite their low contribution to forest carbon stocks, lianas (woody vines) play an important role in the carbon dynamics of tropical forests. As structural parasites, they hinder tree survival, growth and fecundity; hence, they negatively impact net ecosystem productivity and long-term carbon sequestration.Competition (for water and light) drives various forest processes and depends on the local abundance of resources over time. However, evaluating the relative role of resource availability on the interactions between lianas and trees from empirical observations is particularly challenging. Previous approaches have used labour-intensive and ecosystem-scale manipulation experiments, which are infeasible in most situations.We propose to circumvent this challenge by evaluating the uncertainty of water and light capture processes of a process-based vegetation model (ED2) including the liana growth form. We further developed the liana plant functional type in ED2 to mechanistically simulate water uptake and transport from roots to leaves, and start the model from prescribed initial conditions. We then used the PEcAn bioinformatics platform to constrain liana parameters and run uncertainty analyses.Baseline runs successfully reproduced ecosystem gas exchange fluxes (gross primary productivity and latent heat) and forest structural features (leaf area index, aboveground biomass) in two sites (Barro Colorado Island, Panama and Paracou, French Guiana) characterized by different rainfall regimes and levels of liana abundance.Model uncertainty analyses revealed that water limitation was the factor driving the competition between trees and lianas at the drier site (BCI), and during the relatively short dry season of the wetter site (Paracou). In young patches, light competition dominated in Paracou but alternated with water competition between the wet and the dry season on BCI according to the model simulations.The modelling workflow also identified key liana traits (photosynthetic quantum efficiency, stomatal regulation parameters, allometric relationships) and processes (water use, respiration, climbing) driving the model uncertainty. They should be considered as priorities for future data acquisition and model development to improve predictions of the carbon dynamics of liana-infested forests. Synthesis. Competition for water plays a larger role in the interaction between lianas and trees than previously hypothesized, as demonstrated by simulations from a process-based vegetation model.

The Ecosystem Demography model version 2.2 (ED-2.2) is a terrestrial biosphere model that simulates the biophysical, ecological, and biogeochemical dynamics of vertically and horizontally heterogeneous terrestrial ecosystems. In a companion paper (Longo et al., 2019a), we described how the model solves the energy, water, and carbon cycles, and verified the high degree of conservation of these properties in long-term simulations that include long-term (multi-decadal) vegetation dynamics. Here, we present a detailed assessment of the model's ability to represent multiple processes associated with the biophysical and biogeochemical cycles in Amazon forests. We use multiple measurements from eddy covariance towers, forest inventory plots, and regional remote-sensing products to assess the model's ability to represent biophysical, physiological, and ecological processes at multiple timescales, ranging from subdaily to century long. The ED-2.2 model accurately describes the vertical distribution of light, water fluxes, and the storage of water, energy, and carbon in the canopy air space, the regional distribution of biomass in tropical South America, and the variability of biomass as a function of environmental drivers. In addition, ED-2.2 qualitatively captures several emergent properties of the ecosystem found in observations, specifically observed relationships between aboveground biomass, mortality rates, and wood density; however, the slopes of these relationships were not accurately captured. We also identified several limitations, including the model's tendency to overestimate the magnitude and seasonality of heterotrophic respiration and to overestimate growth rates in a nutrient-poor tropical site. The evaluation presented here highlights the potential of incorporating structural and functional heterogeneity within biomes in Earth system models (ESMs) and to realistically represent their impacts on energy, water, and carbon cycles. We also identify several priorities for further model development.

Abstract. The Ecosystem Demography Model version 2.2 (ED-2.2) is a terrestrial biosphere model that simulates the biophysical and biogeochemical cycles of dynamic ecosystems while considering the role of vertical structure of plant communities and the heterogeneity of such structures across the landscape. In a companion paper, we described in detail how the model solves the energy, water, and carbon cycles, and verified the excellent conservation of such properties in long-term simulation. Here, we present a thorough assessment of the model's ability to represent multiple processes associated with the biophysical and biogeochemical cycles, with focus on the Amazon forest. We used multiple measurements from eddy covariance towers, forest inventory plots and regional remote-sensing products to assess the model's ability to represent biophysical, physiological, and ecological processes at multiple time scales ranging from sub-daily to century-long. The ED-2.2 model accurately describes the vertical distribution of light, water fluxes and the storage of water, energy and carbon in the canopy air space, the regional distribution of biomass in tropical South America, and the variability of biomass as a function of environmental drivers. In addition, ED-2.2 also simulates emerging properties of the ecosystem found in observations, such as the relationship between biomass and mortality rates and wood density, although the relationships predicted by the model were biased. We also identified some of the model limitations, such as the model's tendency to overestimate the magnitude and seasonality of heterotrophic respiration, and to overestimate growth rates in a nutrient-poor tropical site. The evaluation presented here highlights the potential of incorporating structural and functional heterogeneity within biomes in ESMs, to realistically represent the role of forest structure and composition on energy, water, and carbon cycles, as well as the priority areas for further model development.

Tropical vegetation is a major source of global land surface evapotranspiration, and can thus play a major role in global hydrological cycles and global atmospheric circulation. Accurate prediction of tropical evapotranspiration is critical to our understanding of these processes under changing climate. We examined the controls on evapotranspiration in tropical vegetation at 21 pan-tropical eddy covariance sites, conducted a comprehensive and systematic evaluation of 13 evapotranspiration models at these sites, and assessed the ability to scale up model estimates of evapotranspiration for the test region of Amazonia. Net radiation was the strongest determinant of evapotranspiration (mean evaporative fraction was 0.72) and explained 87% of the variance in monthly evapotranspiration across the sites. Vapor pressure deficit was the strongest residual predictor (14%), followed by normalized difference vegetation index (9%), precipitation (6%) and wind speed (4%). The radiation-based evapotranspiration models performed best overall for three reasons: (1) the vegetation was largely decoupled from atmospheric turbulent transfer (calculated from Ω decoupling factor), especially at the wetter sites; (2) the resistance-based models were hindered by difficulty in consistently characterizing canopy (and stomatal) resistance in the highly diverse vegetation; (3) the temperature-based models inadequately captured the variability in tropical evapotranspiration. We evaluated the potential to predict regional evapotranspiration for one test region: Amazonia. We estimated an Amazonia-wide evapotranspiration of 1370 mm yr⁻¹, but this value is dependent on assumptions about energy balance closure for the tropical eddy covariance sites; a lower value (1096 mm yr⁻¹) is considered in discussion on the use of flux data to validate and interpolate models.

The impact of increases in drought frequency on the Amazon forest's composition, structure and functioning remain uncertain. We used a process- and individual-based ecosystem model (ED2) to quantify the forest's vulnerability to increased drought recurrence. We generated meteorologically realistic, drier-than-observed rainfall scenarios for two Amazon forest sites, Paracou (wetter) and Tapajós (drier), to evaluate the impacts of more frequent droughts on forest biomass, structure and composition. The wet site was insensitive to the tested scenarios, whereas at the dry site biomass declined when average rainfall reduction exceeded 15%, due to high mortality of large-sized evergreen trees. Biomass losses persisted when year-long drought recurrence was shorter than 2-7 yr, depending upon soil texture and leaf phenology. From the site-level scenario results, we developed regionally applicable metrics to quantify the Amazon forest's climatological proximity to rainfall regimes likely to cause biomass loss > 20% in 50 yr according to ED2 predictions. Nearly 25% (1.8 million km² ) of the Amazon forests could experience frequent droughts and biomass loss if mean annual rainfall or interannual variability changed by 2σ. At least 10% of the high-emission climate projections (CMIP5/RCP8.5 models) predict critically dry regimes over 25% of the Amazon forest area by 2100.