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.
Models of vegetation function are widely used to predict the effects of climate change on carbon, water and nutrient cycles of terrestrial ecosystems, and their feedbacks to climate. Stomatal conductance, the process that governs plant water use and carbon uptake, is fundamental to such models. In this paper, we reconcile two long-standing theories of stomatal conductance. The empirical approach, which is most commonly used in vegetation models, is phenomenological, based on experimental observations of stomatal behaviour in response to environmental conditions. The optimal approach is based on the theoretical argument that stomata should act to minimize the amount of water used per unit carbon gained. We reconcile these two approaches by showing that the theory of optimal stomatal conductance can be used to derive a model of stomatal conductance that is closely analogous to the empirical models. Consequently, we obtain a unified stomatal model which has a similar form to existing empirical models, but which now provides a theoretical interpretation for model parameter values. The key model parameter, g 1 , is predicted to increase with growth temperature and with the marginal water cost of carbon gain. The new model is fitted to a range of datasets ranging from tropical to boreal trees. The parameter g 1 is shown to vary with growth temperature, as predicted, and also with plant functional type. The model is shown to correctly capture responses of stomatal conductance to changing atmospheric CO 2 , and thus can be used to test for stomatal acclimation to elevated CO 2 . The reconciliation of the optimal and empirical approaches to modelling stomatal conductance is important for global change biology because it provides a simple theoretical framework for analyzing, and simulating, the coupling between carbon and water cycles under environmental change.
The FLUXNET2015 dataset provides ecosystem-scale data on CO 2 , water, and energy exchange between the biosphere and the atmosphere, and other meteorological and biological measurements, from 212 sites around the globe (over 1500 site-years, up to and including year 2014). These sites, independently managed and operated, voluntarily contributed their data to create global datasets. Data were quality controlled and processed using uniform methods, to improve consistency and intercomparability across sites. The dataset is already being used in a number of applications, including ecophysiology studies, remote sensing studies, and development of ecosystem and Earth system models. FLUXNET2015 includes derived-data products, such as gap-filled time series, ecosystem respiration and photosynthetic uptake estimates, estimation of uncertainties, and metadata about the measurements, presented for the first time in this paper. In addition, 206 of these sites are for the first time distributed under a Creative Commons (CC-BY 4.0) license. This paper details this enhanced dataset and the processing methods, now made available as open-source codes, making the dataset more accessible, transparent, and reproducible.
Climate change is predicted to increase both drought frequency and duration, and when coupled with substantial warming, will establish a new hydroclimatological model for many regions. Large-scale, warm droughts have recently occurred in North America, Africa, Europe, Amazonia and Australia, resulting in major effects on terrestrial ecosystems, carbon balance and food security. Here we compare the functional response of above-ground net primary production to contrasting hydroclimatic periods in the late twentieth century (1975-1998), and drier, warmer conditions in the early twenty-first century (2000-2009) in the Northern and Southern Hemispheres. We find a common ecosystem water-use efficiency (WUE(e): above-ground net primary production/evapotranspiration) across biomes ranging from grassland to forest that indicates an intrinsic system sensitivity to water availability across rainfall regimes, regardless of hydroclimatic conditions. We found higher WUE(e) in drier years that increased significantly with drought to a maximum WUE(e) across all biomes; and a minimum native state in wetter years that was common across hydroclimatic periods. This indicates biome-scale resilience to the interannual variability associated with the early twenty-first century drought--that is, the capacity to tolerate low, annual precipitation and to respond to subsequent periods of favourable water balance. These findings provide a conceptual model of ecosystem properties at the decadal scale applicable to the widespread altered hydroclimatic conditions that are predicted for later this century. Understanding the hydroclimatic threshold that will break down ecosystem resilience and alter maximum WUE(e) may allow us to predict land-surface consequences as large regions become more arid, starting with water-limited, low-productivity grasslands.
Leaf phenology was monitored for 49 woody species (trees and tall shrubs) each month over a 2.5-year period in a humid, wet-dry tropical eucalypt savanna at Solar Village, near Darwin, Australia. In the 10 most common species, which spanned the range of phenological types, phenology was monitored every two weeks. To investigate the relationships between leaf phenology and plant water status, pre-dawn leaf water potential was monitored in eight common species every 4-6 weeks. Four main phenological types were described: (1) evergreen species, which retained full canopy throughout the year; (2) brevi-or partly deciduous species, in which the amount of canopy fell significantly, but briefly, during at least one dry season during the study period, but to levels not below 50% of full canopy; (3) semideciduous species in which canopy fell to below 50% of full canopy in each of the dry seasons; and (4) fully deciduous species, which lost all leaves during the early-mid dry season, and remained leafless for at least one month. Of these 49 species, 24% were evergreen, 20% were brevideciduous, 29% were semideciduous, and 27% were fully deciduous. Leaf fall occurred 1-2 months earlier in the dry season for the fully deciduous species than for the semideciduous species. Leaf fall in all species was coincident with the attainment of seasonal minima in leaf water potential, which were, on average, about Ϫ1.5 to Ϫ2.0 MPa in the evergreen and semideciduous species, compared with about Ϫ0.5 to Ϫ1.0 MPa in the fully deciduous species. Leaf flushing occurred throughout the dry season in the two evergreen species, with a major peak in the late dry season. In five semideciduous species and one of the fully deciduous species, leaf flushing commenced in the late dry season prior to the occurrence of any rain. In two fully deciduous species, leaf flushing occurred only after the first storms of the early wet season. There was variation in the timing of flushing, both between species within years and between years for some species. However, all species commenced leaf flushing after water potentials rose, following the attainment of seasonal minima in pre-dawn leaf water potential. Soil moisture at 1 m did not fall below permanent wilting point during the dry season; hence, reserves of soil water at the end of the dry season were sufficient to support the whole-plant rehydration that preceded leaf flushing in the absence of rain. These results are consistent with hypotheses, developed in the neotropics, that leaf phenology in trees from the wet-dry tropics is largely controlled by endogenous mechanisms.
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