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The Global Carbon Project (GCP) has published global carbon budgets annually since 2007 (Canadell et al. [2007], Proc Natl Acad Sci USA, 104, 18866–18870; Raupach et al. [2007], Proc Natl Acad Sci USA, 104, 10288–10293). There are many scientists involved, but the terrestrial fluxes that appear in the budgets are not well understood by ecologists and biogeochemists outside of that community. The purpose of this paper is to make the terrestrial fluxes of carbon in those budgets more accessible to a broader community. The GCP budget is composed of annual perturbations from pre‐industrial conditions, driven by addition of carbon to the system from combustion of fossil fuels and by transfers of carbon from land to the atmosphere as a result of land use. The budget includes a term for each of the major fluxes of carbon (fossil fuels, oceans, land) as well as the rate of carbon accumulation in the atmosphere. Land is represented by two terms: one resulting from direct anthropogenic effects (Land Use, Land‐Use Change, and Forestry or land management) and one resulting from indirect anthropogenic (e.g., CO2, climate change) and natural effects. Each of these two net terrestrial fluxes of carbon, in turn, is composed of opposing gross emissions and removals (e.g., deforestation and forest regrowth). Although the GCP budgets have focused on the two net terrestrial fluxes, they have paid little attention to the gross components, which are important for a number of reasons, including understanding the potential for land management to remove CO2 from the atmosphere and understanding the processes responsible for the sink for carbon on land. In contrast to the net fluxes of carbon, which are constrained by the global carbon budget, the gross fluxes are largely unconstrained, suggesting that there is more uncertainty than commonly believed about how terrestrial carbon emissions will respond to future fossil fuel emissions and a changing climate. [ABSTRACT FROM AUTHOR]

The terrestrial carbon cycle plays a critical role in determining levels of atmospheric CO2 that result from anthropogenic carbon emissions. Elevated atmospheric CO2 is thought to stimulate terrestrial carbon uptake, through the process of CO2 fertilization of vegetation productivity. This negative carbon cycle feedback results in reduced atmospheric CO2 growth, and has likely accounted for a substantial portion of the historical terrestrial carbon sink. However, the future strength of CO2 fertilization in response to continued carbon emissions and atmospheric CO2 rise is highly uncertain. In this paper, the ramifications of CO2 fertilization in simulations of future climate change are explored, using an intermediate complexity coupled climate–carbon model. It is shown that the absence of future CO2 fertilization results in substantially higher future CO2 levels in the atmosphere, as this removes the dominant contributor to future terrestrial carbon uptake in the model. As a result, climate changes are larger, though the radiative effect of higher CO2 on surface temperatures in the model is offset by about 30% due to reduced positive dynamic vegetation feedbacks; that is, the removal of CO2 fertilization results in less vegetation expansion in the model, which would otherwise constitute an important positive surface albedo-temperature feedback. However, the effect of larger climate changes has other important implications for the carbon cycle – notably to further weaken remaining carbon sinks in the model. As a result, positive climate–carbon cycle feedbacks are larger when CO2 fertilization is absent. This creates an interesting synergism of terrestrial carbon cycle feedbacks, whereby positive (climate–carbon cycle) feedbacks are amplified when a negative (CO2 fertilization) feedback is removed. [ABSTRACT FROM AUTHOR]

Atmospheric measurements and land-based inventories imply that terrestrial ecosystems in the northern hemisphere are taking up significant amounts of anthropogenic cabon dioxide (CO2) emissions; however, there is considerable disagreement about the causes of this uptake, and its expected future trajectory. In this paper, we use the ecosystem demography (ED) model to quantify the contributions of disturbance history, CO2 fertilization and climate variability to the past, current, and future terrestrial carbon fluxes in the Eastern United States. The simulations indicate that forest regrowth following agricultural abandonment accounts for uptake of 0.11 Pg C yr−1 in the 1980s and 0.15 Pg C yr−1 in the 1990s, and regrowth following forest harvesting accounts for an additional 0.1 Pg C yr−1 of uptake during both these decades. The addition of CO2 fertilization into the model simulations increases carbon uptake rates to 0.38 Pg C yr−1 in the 1980s and 0.47 Pg C yr−1 in the 1990s. Comparisons of predicted aboveground carbon uptake to regional-scale forest inventory measurements indicate that the model's predictions in the absence of CO2 fertilization are 14% lower than observed, while in the presence of CO2 fertilization, predicted uptake rates are 28% larger than observed. Comparable results are obtained from comparisons of predicted total Net Ecosystem Productivity to the carbon fluxes observed at the Harvard Forest flux tower site and in model simulations free-air CO2 enrichment (FACE) experiments. These results imply that disturbance history is the principal mechanism responsible for current carbon uptake in the Eastern United States, and that conventional biogeochemical formulations of plant growth overestimate the response of plants to rising CO2 levels. Model projections out to 2100 imply that the carbon uptake arising from forest regrowth will increasingly be dominated by forest regrowth following harvesting. Consequently, actual carbon storage declines to near zero by the end of the 21st century as the forest regrowth that has occurred since agricultural abandonment comes into equilibrium with the landscape's new disturbance regime. Incorporating interannual climate variability into the model simulations gives rise to large interannual variation in regional carbon fluxes, indicating that long-term measurements are necessary to detect the signature of processes that give rise to long-term uptake and storage. [ABSTRACT FROM AUTHOR]

The terrestrial biosphere plays an important role in the global carbon cycle. In the 1994 Intergovernmental Panel Assessment on Climate Change (IPCC), an effort was made to improve the quantification of terrestrial exchanges and potential feedbacks from climate, changing CO2, and other factors; this paper presents the key results from that assessment, together with expanded discussion. The carbon cycle is the fluxes of carbon among four main reservoirs: fossil carbon, the atmosphere, the oceans, and the terrestrial biosphere. Emissions of fossil carbon during the 1980s averaged 5.5 Gt y-1. During the same period, the atmosphere gained 3.2 Gt C y-1, and the oceans are believed to have absorbed 2.0 Gt C y-1. The regrowing forests of the Northern Hemisphere may have absorbed 0.5 Gt C y-1 during this period. Meanwhile, tropical deforestation is thought to have released an average 1.6 Gt C y-1 over the 1980s. While the fluxes among the four pools should balance, the average 1980s values lead to a 'missing sink' of 1.4 Gt C y-1 Several processes, including forest regrowth, CO2 fertilization of plant growth (c. 1.0 Gt C y-1), N deposition (c. 0.6 Gt C y-1), and their interactions, may account for the budget imbalance. However, it remains difficult to quantify the influences of these separate but interactive processes. Uncertainties in the individual numbers are large, and are themselves poorly quantified. This paper presents detail beyond the IPCC assessment on procedures used to approximate the flux uncertainties. Lack of knowledge about positive and negative feedbacks from the biosphere is a major limiting factor to credible simulations of future atmospheric CO2 concentrations. Analyses of the atmospheric gradients of CO2 and 13CO2 concentrations provide increasingly strong evidence for terrestrial sinks, potentially distributed between Northern Hemisphere and tropical regions, but conclusive detection in direct biomass and soil measurements remains elusive. Current regional-to-global terrestrial ecosystem models with coupled carbon and nitrogen cycles represent the effects of CO2 fertilization differently, but all suggest long-term responses to CO2 that are substantially smaller than potential leaf- or laboratory whole plant-level responses. Analyses of emissions and biogeochemical fluxes consistent with eventual stabilization of atmospheric CO2 concentrations are sensitive to the way in which biospheric feedbacks are modeled by c. 15%. Decisions about land use can have effects of 100s of Gt C over the next few centuries, with similarly significant effects on the atmosphere. Critical areas for future research are continued measurements and analyses of atmospheric data (CO2 and 13CO2) to serve as large-scale constraints, process studies of the scaling from the photosynthetic response to CO2 to whole-ecosystem carbon storage, and rigorous quantification of the effects of changing land use on carbon storage. [ABSTRACT FROM AUTHOR]

The effect of nutrient availability on plant growth and the terrestrial carbon sink under climate change and elevated CO2 remains one of the main uncertainties of the terrestrial carbon cycle. This is partially due to the difficulty of assessing nutrient limitation at large scales over long periods of time. Consistent declines in leaf nitrogen (N) content and leaf δ15N have been used to suggest that nitrogen limitation has increased in recent decades, most likely due to the concurrent increase in atmospheric CO2. However, such data sets are often not straightforward to interpret due to the complex factors that contribute to the spatial and temporal variation in leaf N and isotope concentration. We use the land surface model (LSM) QUINCY, which has the unique capacity to represent N isotopic processes, in conjunction with two large data sets of foliar N and N isotope content. We run the model with different scenarios to test whether foliar δ15N isotopic data can be used to infer large‐scale N limitation and if the observed trends are caused by increasing atmospheric CO2, changes in climate or changes in sources and magnitude of anthropogenic N deposition. We show that while the model can capture the observed change in leaf N content and predict widespread increases in N limitation, it does not capture the pronounced, but very spatially heterogeneous, decrease in foliar δ15N observed in the data across the globe. The addition of an observation‐based temporal trend in isotopic composition of N deposition leads to a more pronounced decrease in simulated leaf δ15N. Our results show that leaf δ15N observations cannot, on their own, be used to assess global‐scale N limitation and that using such a data set in conjunction with an LSM can reveal the drivers behind the observed patterns. [ABSTRACT FROM AUTHOR]

Anthropogenic climate change is expected to impact ecosystem structure, biodiversity and ecosystem services in Africa profoundly. We used the adaptive Dynamic Global Vegetation Model (aDGVM), which was originally developed and tested for Africa, to quantify sources of uncertainties in simulated African potential natural vegetation towards the end of the 21st century. We forced the aDGVM with regionally downscaled high-resolution climate scenarios based on an ensemble of six general circulation models (GCMs) under two representative concentration pathways (RCPs 4.5 and 8.5). Our study assessed the direct effects of climate change and elevated CO2 on vegetation change and its plant-physiological drivers. Total increase in carbon in aboveground biomass in Africa until the end of the century was between 18% to 43% (RCP4.5) and 37% to 61% (RCP8.5) and was associated with woody encroachment into grasslands and increased woody cover in savannas. When direct effects of CO2 on plants were omitted, woody encroachment was muted and carbon in aboveground vegetation changed between -8 to 11% (RCP 4.5) and -22 to -6% (RCP8.5). Simulated biome changes lacked consistent large-scale geographical patterns of change across scenarios. In Ethiopia and the Sahara/Sahel transition zone, the biome changes forecast by the aDGVM were consistent across GCMs and RCPs. Direct effects from elevated CO2 were associated with substantial increases in water use efficiency, primarily driven by photosynthesis enhancement, which may relieve soil moisture limitations to plant productivity. At the ecosystem level, interactions between fire and woody plant demography further promoted woody encroachment. We conclude that substantial future biome changes due to climate and CO2 changes are likely across Africa. Because of the large uncertainties in future projections, adaptation strategies must be highly flexible. Focused research on CO2 effects, and improved model representations of these effects will be necessary to reduce these uncertainties. [ABSTRACT FROM AUTHOR]

Predicted responses of transpiration to elevated atmospheric CO2 concentration ( eCO2) are highly variable amongst process-based models. To better understand and constrain this variability amongst models, we conducted an intercomparison of 11 ecosystem models applied to data from two forest free-air CO2 enrichment ( FACE) experiments at Duke University and Oak Ridge National Laboratory. We analysed model structures to identify the key underlying assumptions causing differences in model predictions of transpiration and canopy water use efficiency. We then compared the models against data to identify model assumptions that are incorrect or are large sources of uncertainty. We found that model-to-model and model-to-observations differences resulted from four key sets of assumptions, namely (i) the nature of the stomatal response to elevated CO2 (coupling between photosynthesis and stomata was supported by the data); (ii) the roles of the leaf and atmospheric boundary layer (models which assumed multiple conductance terms in series predicted more decoupled fluxes than observed at the broadleaf site); (iii) the treatment of canopy interception (large intermodel variability, 2-15%); and (iv) the impact of soil moisture stress (process uncertainty in how models limit carbon and water fluxes during moisture stress). Overall, model predictions of the CO2 effect on WUE were reasonable (intermodel μ = approximately 28% ± 10%) compared to the observations (μ = approximately 30% ± 13%) at the well-coupled coniferous site (Duke), but poor (intermodel μ = approximately 24% ± 6%; observations μ = approximately 38% ± 7%) at the broadleaf site (Oak Ridge). The study yields a framework for analysing and interpreting model predictions of transpiration responses to eCO2, and highlights key improvements to these types of models. [ABSTRACT FROM AUTHOR]

Assessing the potential future of current forest stands is a key to design conservation strategies and understanding potential future impacts to ecosystem service supplies. This is particularly true in the Mediterranean basin, where important future climatic changes are expected. Here, we assess and compare two commonly used modeling approaches (niche- and process-based models) to project the future of current stands of three forest species with contrasting distributions, using regionalized climate for continental Spain. Results highlight variability in model ability to estimate current distributions, and the inherent large uncertainty involved in making projections into the future. CO fertilization through projected increased atmospheric CO concentrations is shown to increase forest productivity in the mechanistic process-based model (despite increased drought stress) by up to three times that of the non-CO fertilization scenario by the period 2050-2080, which is in stark contrast to projections of reduced habitat suitability from the niche-based models by the same period. This highlights the importance of introducing aspects of plant biogeochemistry into current niche-based models for a realistic projection of future species distributions. We conclude that the future of current Mediterranean forest stands is highly uncertain and suggest that a new synergy between niche- and process-based models is urgently needed in order to improve our predictive ability. [ABSTRACT FROM AUTHOR]

As atmospheric CO2 levels rise, temperate and boreal forests in the Northern Hemisphere are gaining importance as carbon sinks. Quantification of that role, however, has been difficult due to the confounding effects of climate change. Recent large-scale experiments with quaking aspen ( Populus tremuloides), a dominant species in many northern forest ecosystems, indicate that elevated CO2 levels can enhance net primary production. Field studies also reveal that droughts contribute to extensive aspen mortality. To complement this work, we analyzed how the growth of wild aspen clones in Wisconsin has responded to historical shifts in CO2 and climate, accounting for age, genotype (microsatellite heterozygosity), and other factors. Aspen growth has increased an average of 53% over the past five decades, primarily in response to the 19.2% rise in ambient CO2 levels. CO2-induced growth is particularly enhanced during periods of high moisture availability. The analysis accounts for the highly nonlinear changes in growth rate with age, and is unaffected by sex or location sampled. Growth also increases with individual heterozygosity, but this heterozygote advantage has not changed with rising levels of CO2 or moisture. Thus, increases in future growth predicted from previous large-scale, common-garden work are already evident in this abundant and ecologically important tree species. Owing to aspen's role as a foundation species in many North American forest ecosystems, CO2-stimulated growth is likely to have repercussions for numerous associated species and ecosystem processes. [ABSTRACT FROM AUTHOR]

Increased atmospheric [CO2] could theoretically lead to increased forest productivity (‘CO2 fertilization’). This mechanism was hypothesized as a possible explanation for biomass increases reported from tropical forests in the last 30+ years. We used unique long-term records of annually measured stands (eighteen 0.5 ha plots, 10 years) and focal tree species (six species, 24 years) to assess the effects of rainfall, temperature, and atmospheric [CO2] on annual wood production in a neotropical rain forest. Our study area was a meso-scale section (600 ha) of old-growth Tropical Wet Forest in NE Costa Rica. Using the repeated remeasurements we directly assessed the relative effects of interannual climatic variation and increasing atmospheric [CO2] on wood production. A remarkably simple two-factor model explained 91% of the interannual variance in stand-level tree growth; the statistically independent factors were total dry season rainfall (positive effect, r2=0.85) and night-time temperature (negative effect, r2=0.42). Stand-level tree mortality increased significantly with night-time temperature. After accounting for dry season rainfall and night-time temperature, there was no effect of annual [CO2] on tree growth in either the stand or focal species data. Tree growth in this Tropical Wet Forest was surprisingly sensitive to the current range of dry season conditions and to variations in mean annual night-time temperature of 1–2°. Our results suggest that wood production in the lowland rainforests of NE Costa Rica (and by extension in other tropical regions) may be severely reduced in future climates that are only slightly drier and/or warmer. [ABSTRACT FROM AUTHOR]

Stem radial growth responds to environmental conditions, and has been widely used as a proxy to study long-term patterns of tree growth and to assess the impact of environmental changes on growth patterns. In this study, we use a tree ring dataset from the Catalan Ecological and Forest Inventory to study the temporal variability of Scots pine ( Pinus sylvestris L.) stem growth during the 20th century across a relatively large region (Catalonia, NE Spain) close to the southern limit of the distribution of the species. Basal area increment (BAI) was modelled as a function of tree size and environmental variables by means of mixed effects models. Our results showed an overall increase of 84% in Scots pine BAI during the 20th century, consistent with most previous studies for temperate forests. This trend was associated with increased atmospheric CO2 concentrations and, possibly, with a general increase in nutrient availability, and we interpreted it as a fertilization effect. Over the same time period, there was also a marked increase in temperature across the study region (0.19 °C per decade on average). This warming had a negative impact on radial growth, particularly at the drier sites, but its magnitude was not enough to counteract the fertilization effect. In fact, the substantial warming observed during the 20th century in the study area did not result in a clear pattern of increased summer drought stress because of the large variability in precipitation, which did not show any clear time trend. But the situation may change in the future if temperatures continue to rise and/or precipitation becomes scarcer. Such a change could potentially reverse the temporal trend in growth, particularly at the driest sites, and is suggested in our data by the relative constancy of radial growth after ca. 1975, coinciding with the warmer period. If this situation is representative of other relatively dry, temperate forests, the implications for the regional carbon balance would be substantial. [ABSTRACT FROM AUTHOR]

Results from free-air CO2 enrichment (FACE) experiments in temperate climates indicate that the response of forest net primary productivity (NPP) to elevated CO2 might be highly conserved across a broad range of productivities. In this study, we show that the LPJ-GUESS dynamic vegetation model reproduces the magnitude of the NPP enhancement at temperate forest FACE experiments. A global application of the model suggests that the response found in the experiments might also be representative of the average response of forests globally. However, the predicted NPP enhancement in tropical forests is more than twice as high as in boreal forests, suggesting that currently available FACE results are not applicable to these ecosystems. The modeled geographic pattern is to a large extent driven by the temperature dependence of the relative affinities of the primary assimilation enzyme (Rubisco) for CO2 and O2. [ABSTRACT FROM AUTHOR]