Tropical forests hold large stores of carbon, yet uncertainty remains regarding their quantitative contribution to the global carbon cycle. One approach to quantifying carbon biomass stores consists in inferring changes from long-term forest inventory plots. Regression models are used to convert inventory data into an estimate of aboveground biomass (AGB). We provide a critical reassessment of the quality and the robustness of these models across tropical forest types, using a large dataset of 2,410 trees >or= 5 cm diameter, directly harvested in 27 study sites across the tropics. Proportional relationships between aboveground biomass and the product of wood density, trunk cross-sectional area, and total height are constructed. We also develop a regression model involving wood density and stem diameter only. Our models were tested for secondary and old-growth forests, for dry, moist and wet forests, for lowland and montane forests, and for mangrove forests. The most important predictors of AGB of a tree were, in decreasing order of importance, its trunk diameter, wood specific gravity, total height, and forest type (dry, moist, or wet). Overestimates prevailed, giving a bias of 0.5-6.5% when errors were averaged across all stands. Our regression models can be used reliably to predict aboveground tree biomass across a broad range of tropical forests. Because they are based on an unprecedented dataset, these models should improve the quality of tropical biomass estimates, and bring consensus about the contribution of the tropical forest biome and tropical deforestation to the global carbon cycle.

Plant traits – the morphological, anatomical, physiological, biochemical and phenological characteristics of plants and their organs – determine how primary producers respond to environmental factors, affect other trophic levels, influence ecosystem processes and services and provide a link from species richness to ecosystem functional diversity. Trait data thus represent the raw material for a wide range of research from evolutionary biology, community and functional ecology to biogeography. Here we present the global database initiative named TRY, which has united a wide range of the plant trait research community worldwide and gained an unprecedented buy-in of trait data: so far 93 trait databases have been contributed. The data repository currently contains almost three million trait entries for 69 000 out of the world's 300 000 plant species, with a focus on 52 groups of traits characterizing the vegetative and regeneration stages of the plant life cycle, including growth, dispersal, establishment and persistence. A first data analysis shows that most plant traits are approximately log-normally distributed, with widely differing ranges of variation across traits. Most trait variation is between species (interspecific), but significant intraspecific variation is also documented, up to 40% of the overall variation. Plant functional types (PFTs), as commonly used in vegetation models, capture a substantial fraction of the observed variation – but for several traits most variation occurs within PFTs, up to 75% of the overall variation. In the context of vegetation models these traits would better be represented by state variables rather than fixed parameter values. The improved availability of plant trait data in the unified global database is expected to support a paradigm shift from species to trait-based ecology, offer new opportunities for synthetic plant trait research and enable a more realistic and empirically grounded representation of terrestrial vegetation in Earth system models.

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 CO 2 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 1C. 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 CO 2 balance required the introduction of substantial biome-specific closure terms. Nonclosure was taken as an indication that respiratory processes, advection, and non-CO 2 carbon fluxes are not presently being adequately accounted for. Nomenclauture:DOC 5 dissolved organic carbon; fNPP 5 foliage component of NPP; GPP 5 gross primary production (GPP40 denotes photosynthetic uptake); mNPP 5 missing component of NPP;NBP 5 net biome production (NBP40 denotes biome uptake); NECB 5 net ecosystem carbon balance (NECB40 denotes ecosystem uptake); NEE 5 net ecosystem exchange (NEE40 denotes ecosystem uptake); NEP 5 net ecosystem production (NEP40 denotes ecosystem uptake); NPP 5 net primary production (NPP40 denotes ecosystem uptake); R a 5 autotrophic respiration (R a 40 denotes respiratory losses); R e 5 ecosystem respiration (R e 40 denotes respiratory losses); R h 5 heterotrophic respiration (R h 40 denotes respiratory losses); rNPP 5 root component of NPP;R s 5 soil respiration (R s 40 denotes respiratory losses); VOC 5 volatile organic compounds; wNPP 5 wood component of NPP

There are pressing reasons for developing a better understanding of net primary production (NPP) in the world's forests. These ecosystems play a large role in the world's carbon budget, and their dynamics, which are likely to be responding to global changes in climate and atmospheric composition, have major economic implications and impacts on global biodiversity. Although there is a long history of forest NPP studies in the ecological literature, current understanding of ecosystem‐level production remains limited. Forest NPP cannot be directly measured; it must be approached by indirect methods. To date, field measurements have been largely restricted to a few aspects of NPP; methods are still lacking for field assessment of others, and past studies have involved confusion about the types of measurements needed. As a result, existing field‐based estimates of forest NPP are likely to be significant underestimates. In this paper we provide a conceptual framework to guide efforts toward improved estimates of forest NPP. We define the quantity NPP* as the summed classes of organic material that should be measured or estimated in field studies for an estimate of total NPP. We discuss the above‐ and belowground components of NPP* and the available methods for measuring them in the field. We then assess the implications of the limitations of past studies for current understanding of NPP in forest ecosystems, discuss how field NPP* measurements can be used to complement tower‐based studies of forest carbon flux, and recommend design criteria for future field studies of forest NPP.

Moist tropical forests in Amazonia and elsewhere are subjected to increasingly severe drought episodes through the El Niño–Southern Oscillation (ENSO) and possibly through deforestation‐driven reductions in rainfall. The effects of this trend on tropical forest canopy dynamics, emissions of greenhouse gases, and other ecological functions are potentially large but poorly understood. We established a throughfall exclusion experiment in an east‐central Amazon forest (Tapajós National Forest, Brazil) to help understand these effects. After 1‐year intercalibration period of two 1‐ha forest plots, we installed plastic panels and wooden gutters in the understory of one of the plots, thereby excluding ∼890 mm of throughfall during the exclusion period of 2000 (late January to early August) and ∼680 mm thus far in the exclusion period of 2001 (early January to late May). Average daily throughfall reaching the soil during the exclusion period in 2000 was 4.9 and 8.3 mm in the treatment and control plots and was 4.8 and 8.1 mm in 2001, respectively. During the first exclusion period, surface soil water content (0–2 m) declined by ∼100 mm, while deep soil water (2–11 m) was unaffected. During the second exclusion period, which began shortly after the dry season when soil water content was low, surface and deep soil water content declined by ∼140 and 160 mm, respectively. Although this depletion of soil water provoked no detectable increase in leaf drought stress (i.e., no reduction in predawn leaf water potential), photosynthetic capacity declined for some species, the canopy thinned (greater canopy openness and lower leaf area index) during the second exclusion period, stem radial growth of trees <15 m tall declined, and fine litterfall declined in the treatment plot, as did tree fruiting. Aboveground net primary productivity (NPP) (stemwood increment and fine litter production) declined by one fourth, from 15.1 to 11.4 Mg ha−1 yr−1, in the treatment plot and decreased slightly, from 11.9 to 11.5 Mg ha−1 yr−1, in the control plot. Stem respiration varied seasonally and was correlated with stem radial growth but showed no treatment response. The fastest response to the throughfall exclusion, and the surface soil moisture deficits that it provoked, was found in the soil itself. The treatment reduced N2O emissions and increased CH4 consumption relative to the control plot, presumably in response to the improved soil aeration that is associated with soil drying. Our hypothesis that NO emissions would increase following exclusion was not supported. The conductivity and alkalinity of water percolating through the litter layer and through the mineral soil to a depth of 200 cm was higher in the treatment plot, perhaps because of the lower volume of water that was moving through these soil layers in this plot. Decomposition of the litter showed no difference between plots. In sum, the small soil water reductions provoked during the first 2 years of partial throughfall exclusion were sufficient to lower aboveground NPP, including th...

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. Understanding how tropical forest carbon balance will respond to global change requires knowledge of individual heterotrophic and autotrophic respiratory sources, together with factors that control respiratory variability. We measured leaf, live wood, and soil respiration, along with additional environmental factors over a 1-yr period in a Central Amazon terra firme forest. Scaling these fluxes to the ecosystem, and combining our data with results from other studies, we estimated an average total ecosystem respiration (R eco ) of 7.8 mol·m Ϫ2 ·s Ϫ1. Average estimates (per unit ground area) for leaf, wood, soil, total heterotrophic, and total autotrophic respiration were 2.6, 1.1, 3.2, 5.6, and 2.2 mol·m Ϫ2 ·s Ϫ1 , respectively. Comparing autotrophic respiration with net primary production (NPP) estimates indicated that only ϳ30% of carbon assimilated in photosynthesis was used to construct new tissues, with the remaining 70% being respired back to the atmosphere as autotrophic respiration. This low ecosystem carbon use efficiency (CUE) differs considerably from the relatively constant CUE of ϳ0.5 found for temperate forests. Our R eco estimate was comparable to the above-canopy flux (F ac ) from eddy covariance during defined sustained high turbulence conditions (when presumably F ac ϭ R eco ) of 8.4 (95% CI ϭ 7.5-9.4). Multiple regression analysis demonstrated that ϳ50% of the nighttime variability in F ac was accounted for by friction velocity (u*, a measure of turbulence) variables. After accounting for u* variability, mean F ac varied significantly with seasonal and daily changes in precipitation. A seasonal increase in precipitation resulted in a decrease in F ac , similar to our soil respiration response to moisture. The effect of daily changes in precipitation was complex: precipitation after a dry period resulted in a large increase in F ac , whereas additional precipitation after a rainy period had little effect. This response was similar to that of surface litter (coarse and fine), where respiration is greatly reduced when moisture is limiting, but increases markedly and quickly saturates with an increase in moisture.

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