Abstract: 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 intensification of the hydrological cycle, with an observed and modeled increase in drought incidence and severity, underscores the need to quantify drought effects on carbon cycling and the terrestrial sink. FLUXNET, a global network of eddy covariance towers, provides dense data streams of meteorological data, and through flux partitioning and gap filling algorithms, estimates of net ecosystem productivity (F-NEP), gross ecosystem productivity (P), and ecosystem respiration (R). We analyzed the functional relationship of these three carbon fluxes relative to evaporative fraction (EF), an index of drought and site water status, using monthly data records from 238 micrometeorological tower sites distributed globally across 11 biomes. The analysis was based on relative anomalies of both EF and carbon fluxes and focused on drought episodes by biome and climatic season. Globally P was approximate to 50% more sensitive to a drought event than R. Network-wide drought-induced decreases in carbon flux averaged -16.6 and -9.3 g C m-2 month-1 for P and R, i.e., drought events induced a net decline in the terrestrial sink. However, in evergreen forests and wetlands drought was coincident with an increase in P or R during parts of the growing season. The most robust relationships between carbon flux and EF occurred during climatic spring for F-NEP and in climatic summer for P and R. Upscaling flux sensitivities to a global map showed that spatial patterns for all three carbon fluxes were linked to the distribution of croplands. Agricultural areas exhibited the highest sensitivity whereas the tropical region had minimal sensitivity to drought. Combining gridded flux sensitivities with their uncertainties and the spatial grid of FLUXNET revealed that a more robust quantification of carbon flux response to drought requires additional towers in all biomes of Africa and Asia as well as in the cropland, shrubland, savannah, and wetland biomes globally.

Abstract: The allocation and cycling of carbon (C) within forests is an important component of the biospheric C cycle, but is particularly understudied within tropical forests. We synthesise reported and unpublished results from three lowland rainforest sites in Amazonia (in the regions of Manaus, Tapajos and Caxiuana), all major sites of the Large-Scale Biosphere-Atmosphere Programme (LBA). We attempt a comprehensive synthesis of the C stocks, nutrient status and, particularly, the allocation and internal C dynamics of all three sites. The calculated net primary productivities (NPP) are 10.1 +/- 1.4 Mg C ha(-1) yr(-1) (Manaus), 14.4 +/- 1.3 Mg C ha(-1) yr(-1) (Tapajos) and 10.0 +/- 1.2 Mg C ha(-1) yr(-1) (Caxiuana). All errors bars report standard errors. Soil and leaf nutrient analyses indicate that Tapajos has significantly more plant-available phosphorus and calcium. Autotrophic respiration at all three sites (14.9-21.4 Mg C ha yr(-1)) is more challenging to measure, with the largest component and greatest source of uncertainty being leaf dark respiration. Comparison of measured soil respiration with that predicted from C cycling measurements provides an independent constraint. It shows general good agreement at all three sites, with perhaps some evidence for measured soil respiration being less than expected. Twenty to thirty percent of fixed C is allocated belowground. Comparison of gross primary productivity (GPP), derived from ecosystem flux measurements with that derived from component studies (NPP plus autotrophic respiration) provides an additional crosscheck. The two approaches are in good agreement, giving increased confidence in both approaches to estimating GPP. The ecosystem carbon-use efficiency (CUEs), the ratio of NPP to GPP, is similar at Manaus (0.34 +/- 0.10) and Caxiuana (0.32 +/- 0.07), but may be higher at Tapajos (0.49 +/- 0.16), although the difference is not significant. Old growth or infertile tropical forests may have low CUE compared with recently disturbed and/or fertile forests.

Abstract: Terrestrial ecosystems sequester 2.1 Pg of atmospheric carbon annually. A large amount of the terrestrial sink is realized by forests. However, considerable uncertainties remain regarding the fate of this carbon over both short and long timescales. Relevant data to address these uncertainties are being collected at many sites around the world, but syntheses of these data are still sparse. To facilitate future synthesis activities, we have assembled a comprehensive global database for forest ecosystems, which includes carbon budget variables (fluxes and stocks), ecosystem traits (e.g. leaf area index, age), as well as ancillary site information such as management regime, climate, and soil characteristics. This publicly available database can be used to quantify global, regional or biome-specific carbon budgets; to re-examine established relationships; to test emerging hypotheses about ecosystem functioning [e.g. a constant net ecosystem production (NEP) to gross primary production (GPP) ratio]; and as benchmarks for model evaluations. In this paper, we present the first analysis of this database. We discuss the climatic influences on GPP, net primary production (NPP) and NEP and present the CO2 balances for boreal, temperate, and tropical forest biomes based on micrometeorological, ecophysiological, and biometric flux and inventory estimates. Globally, GPP of forests benefited from higher temperatures and precipitation whereas NPP saturated above either a threshold of 1500 mm precipitation or a mean annual temperature of 10 degrees C. The global pattern in NEP was insensitive to climate and is hypothesized to be mainly determined by nonclimatic conditions such as successional stage, management, site history, and site disturbance. In all biomes, closing the CO2 balance required the introduction of substantial biome-specific closure terms. Nonclosure was taken as an indication that respiratory processes, advection, and non-CO2 carbon fluxes are not presently being adequately accounted for.

Abstract: Comparisons were made among Douglas-fir forest, aspen (broad leaf deciduous) forest and wheatgrass (C-3) grassland for ecosystem-level water-use efficiency (WUE). WUE was defined as the ratio of photosynthetic CO2 assimilation rate and evapotranspiration (ET) rate. The ET data measured by eddy covariance were screened so that they overwhelmingly represented transpiration. The three sites used in this comparison spanned a range of vegetation (plant functional) types and environmental conditions within western Canada. When compared in the relative order Douglas-fir (located on Vancouver Island, BC), aspen (northern Saskatchewan), grassland (southern Alberta), the sites demonstrated a progressive decline in precipitation and a general increase in maximum air temperature and atmospheric saturation deficit (D-max) during the mid-summer. The average (+/- SD) WUE at the grassland site was 2.6 +/- 0.7 mmol mol(-1), which was much lower than the average values observed for the two other sites (aspen: 5.4 +/- 2.3, Douglas-fir: 8.1 +/- 2.4). The differences in WUE among sites were primarily because of variation in ET. The highest maximum ET rates were approximately 5, 3.2 and 2.7 mm day(-1) for the grassland, aspen and Douglas-fir sites, respectively. There was a strong negative correlation between WUE and D-max for all sites. We also made seasonal measurements of the carbon isotope ratio of ecosystem respired CO2 (delta(R)) in order to test for the expected correlation between shifts in environmental conditions and changes to the ecosystem-integrated ratio of leaf intercellular to ambient CO2 concentration (c(i)/c(a)). There was a consistent increase in delta(R) values in the grassland, aspen forest and Douglas-fir forest associated with a seasonal reduction in soil moisture. Comparisons were made between WUE measured using eddy covariance with that calculated based on D and delta(R) measurements. There was excellent agreement between WUE values calculated using the two techniques. Our delta(R) measurements indicated that c(i)/c(a) values were quite similar among the Douglas-fir, aspen and grassland sites, despite large variation in environmental conditions among sites. This implied that the shorter-lived grass species had relatively high c(i)/c(a) values for the D of their habitat. By contrast, the longer-lived Douglas-fir trees were more conservative in water-use with lower c(i)/c(a) values relative to their habitat D. This illustrates the interaction between biological and environmental characteristics influencing ecosystem-level WUE. The strong correlation we observed between the two independent measurements of WUE, indicates that the stable isotope composition of respired CO2 is a useful ecosystem-scale tool to help study constraints to photosynthesis and acclimation of ecosystems to environmental stress.