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Abstract: Although tropical forests differ substantially in form and function, they are often represented as a single biome in global change models, hindering understanding of how different tropical forests will respond to environmental change. The response of the tropical forest biome to environmental change is strongly influenced by forest type. Forest types differ based on functional traits and forest structure, which are readily derived from high resolution airborne remotely sensed data. Whether the spatial resolution of emerging satellite-derived hyperspectral data is sufficient to identify different tropical forest types is unclear. Here, we resample airborne remotely sensed forest data at spatial resolutions relevant to satellite remote sensing (30 m) across two sites in Malaysian Borneo. Using principal component and cluster analysis, we derive and map seven forest types. We find ecologically relevant variations in forest type that correspond to substantial differences in carbon stock, growth, and mortality rate. We find leaf mass per area and canopy phosphorus are critical traits for distinguishing forest type. Our findings highlight the importance of these parameters for accurately mapping tropical forest types using space borne observations.

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.

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.

Earth system models (ESMs) have been developed to represent the role of terrestrial ecosystems on the energy, water, and carbon cycles. However, many ESMs still lack representation of within-ecosystem heterogeneity and diversity. In this paper, we present the Ecosystem Demography model version 2.2 (ED-2.2). In ED-2.2, the biophysical and physiological processes account for the horizontal and vertical heterogeneity of the ecosystem: The energy, water, and carbon cycles are solved separately for a series of vegetation cohorts (groups of individual plants of similar size and plant functional type) distributed across a series of spatially implicit patches (representing collections of microenvironments that have a similar disturbance history).We define the equations that describe the energy, water, and carbon cycles in terms of total energy, water, and carbon, which simplifies the differential equations and guarantees excellent conservation of these quantities in long-term simulation (

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.

Land conversion and changing climate are expected to significantly alter tropical forest hydrology. We used a land surface model integrated with a river routing scheme to analyze the hydrological alterations expected in the Tapajós River basin, a large portion of the Brazilian Amazon, caused by two environmental drivers: climate and land use. The model was forced with two future climate scenarios (years 2026–2045) from the Earth System Model HadGem2-ES with moderate (+4.5 W/m2 radiative forcing value in the year 2100 with respect to preindustrial levels) and severe (+8.5 W/m2) representative atmospheric carbon dioxide pathways (Representative Concentration Pathways). We tested the sensitivity of our results to the uncertainty in future climate projections by running simulations with IPSL-CM5 (wettest scenarios) and GISS-E2 (driest scenarios). Human land use effects on vegetation were evaluated using a limited and an extreme deforestation scenario. Our analysis indicates that climate change is predicted to reduce river flows across seasons (up to 20%) and bring a considerable shift in flow seasonality toward a later onset (nearly 1.5 months) and increase in interannual variability. While land use change partially counteracts the climate-driven diminishing trend in river flows, it is expected to contribute to a further increase in interannual and intraannual variability. From a water management perspective, the overall reduction of river flows and their increased variability, combined with the shift and the shortening of the wet season, could potentially affect the productivity of the large hydropower systems planned for the region and the growing demand for agricultural and transport expansion.

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.

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.

Numerous current efforts seek to improve the representation of ecosystem ecology and vegetation demographic processes within Earth System Models (ESMs). These developments are widely viewed as an important step in developing greater realism in predictions of future ecosystem states and fluxes. Increased realism, however, leads to increased model complexity, with new features raising a suite of ecological questions that require empirical constraints. Here, we review the developments that permit the representation of plant demographics in ESMs, and identify issues raised by these developments that highlight important gaps in ecological understanding. These issues inevitably translate into uncertainty in model projections but also allow models to be applied to new processes and questions concerning the dynamics of real-world ecosystems. We argue that stronger and more innovative connections to data, across the range of scales considered, are required to address these gaps in understanding. The development of first-generation land surface models as a unifying framework for ecophysiological understanding stimulated much research into plant physiological traits and gas exchange. Constraining predictions at ecologically relevant spatial and temporal scales will require a similar investment of effort and intensified inter-disciplinary communication.

Considerable uncertainty surrounds the impacts of anthropogenic climate change on the composition and structure of Amazon forests. Building upon results from two large-scale ecosystem drought experiments in the eastern Brazilian Amazon that observed increases in mortality rates among some tree species but not others, in this study we investigate the physiological traits underpinning these differential demographic responses. Xylem pressure at 50% conductivity (xylem-P50 ), leaf turgor loss point (TLP), cellular osmotic potential (πo ), and cellular bulk modulus of elasticity (ε), all traits mechanistically linked to drought tolerance, were measured on upper canopy branches and leaves of mature trees from selected species growing at the two drought experiment sites. Each species was placed a priori into one of four plant functional type (PFT) categories: drought-tolerant versus drought-intolerant based on observed mortality rates, and subdivided into early- versus late-successional based on wood density. We tested the hypotheses that the measured traits would be significantly different between the four PFTs and that they would be spatially conserved across the two experimental sites. Xylem-P50 , TLP, and πo , but not ε, occurred at significantly higher water potentials for the drought-intolerant PFT compared to the drought-tolerant PFT; however, there were no significant differences between the early- and late-successional PFTs. These results suggest that these three traits are important for determining drought tolerance, and are largely independent of wood density-a trait commonly associated with successional status. Differences in these physiological traits that occurred between the drought-tolerant and drought-intolerant PFTs were conserved between the two research sites, even though they had different soil types and dry-season lengths. This more detailed understanding of how xylem and leaf hydraulic traits vary between co-occuring drought-tolerant and drought-intolerant tropical tree species promises to facilitate a much-needed improvement in the representation of plant hydraulics within terrestrial ecosystem and biosphere models, which will enhance our ability to make robust predictions of how future changes in climate will affect tropical forests.

Amazon forests, which store ∼ 50% of tropical forest carbon and play a vital role in global water, energy, and carbon cycling, are predicted to experience both longer and more intense dry seasons by the end of the 21st century. However, the climate sensitivity of this ecosystem remains uncertain: several studies have predicted large-scale die-back of the Amazon, whereas several more recent studies predict that the biome will remain largely intact. Combining remote-sensing and ground-based observations with a size- and age-structured terrestrial ecosystem model, we explore the sensitivity and ecological resilience of these forests to changes in climate. We demonstrate that water stress operating at the scale of individual plants, combined with spatial variation in soil texture, explains observed patterns of variation in ecosystem biomass, composition, and dynamics across the region, and strongly influences the ecosystem's resilience to changes in dry season length. Specifically, our analysis suggests that in contrast to existing predictions of either stability or catastrophic biomass loss, the Amazon forest's response to a drying regional climate is likely to be an immediate, graded, heterogeneous transition from high-biomass moist forests to transitional dry forests and woody savannah-like states. Fire, logging, and other anthropogenic disturbances may, however, exacerbate these climate change-induced ecosystem transitions.

The hemlock woolly adelgid (HWA; Adelges tsugae Annand) is an introduced insect pest that threatens to decimate eastern hemlock (Tsuga canadensis (L.) Carriere) populations. In this study, we used the ecosystem demography model in conjunction with a stochastic model of HWA spread to predict the impact of HWA infestation on the current and future forest composition, structure, and carbon (C) dynamics in the eastern United States. The spread model predicted that on average the hemlock stands south and east of the Great Lakes would he infested by 2015. southern Michigan would be reached by 2020. and northeastern Minnesota by 2030. For the period 2000-2040. the ecosystem demography model predicted a mean reduction of 0.011 Pg [C.year.sup.-1] (Pg C = [10.sup.15] g C). an 8% decrease, in the uptake of carbon from eastern United States forests as a result of HWA-caused mortality, followed by an increased uptake of 0.015 Pg [C.year.sup.-1] (a 12% increase) in the period 2040-2100. as the area recovers from the loss of hemlock. Overall, we conclude that while locally severe, HWA infestation is unlikely to have a significant impact on the regional patterns of carbon fluxes, given that, eastern emlock represents a limited fraction of the standing biomass of eastern forests and that it has relatively low productivity compared with the tree species that are likely to replace it. Resume: Le puceron lanigere de la pruche (Adelges tsugae Annand) est un insecte ravageur introduit qui menace de decimer les populations de pruche du Canada (Tsuga canadensis (L.) Carriere). Dans cette etude, nous avons utilise le modele demographique des ecosystemes conjointement avec un modele stochastique de propagation du puceron lanigere pour predire l'impact de l'infestation de cet insecte sur la composition, la structure et la dynamique du carbone (C) dans l'est des Etats-Unis. Le modele de propagation predit, qu'en moyenne, les peuplements de pruche situes au sud et a l'est des Grands Lacs seront infestes vers 2015, que le sud du Michigan sera touche vers 2020 et le nord-est du Minnesota vers 2030. Pour la periode allant de 2000 a 2040, le modele d'ecosysteme predit une reduction moyenne de 0.011 Pg [C.an.sup.-1] (Pg C = [10.sup.15 g C), soit une diminution de 8 % de 'absorption de carbone par les forets de l'est des Etats-Unis due a la mortalite causee par le puceron lanigere, suivie d'une augmentation de l'absorption de 0,015 Pg [C.an.sup.-1] (une augmentation de 12 %) au cours de la periode allant de 2040 a 2100. a mesure que la region se remet de la perte des pruches. Globalement, nous arrivons a la conclusion que. bien qu'elle soit severe localement, il est peu probable que l'infestation du puceron lanigere ait un impact significantif sur les patrons regionaux des flux de carbone etant donne que la pruche du Canada represente une faible proportion de la biomasse sur pied dans les forets de l'est et que la productivite de cette essence est relativement faible comparativement aux especes d'arbre les plus susceptibles de la remplacer. [Traduit par la Redaction], Introduction Changes in forest structure and canopy composition or wholesale losses of forestlands as a result of invasive insects, conversion to agriculture, and urbanization inevitably alter the amount of carbon [...]