Abrupt permafrost thaw drives spatially heterogeneous soil moisture and carbon dioxide fluxes in upland tundra.

Permafrost thaw causes the seasonally thawed active layer to deepen, causing the Arctic to shift toward carbon release as soil organic matter becomes susceptible to decomposition. Ground subsidence initiated by ice loss can cause these soils to collapse abruptly, rapidly shifting soil moisture as microtopography changes and also accelerating carbon and nutrient mobilization. The uncertainty of soil moisture trajectories during thaw makes it difficult to predict the role of abrupt thaw in suppressing or exacerbating carbon losses. In this study, we investigated the role of shifting soil moisture conditions on carbon dioxide fluxes during a 13-year permafrost warming experiment that exhibited abrupt thaw. Warming deepened the active layer differentially across treatments, leading to variable rates of subsidence and formation of thermokarst depressions. In turn, differential subsidence caused a gradient of moisture conditions, with some plots becoming consistently inundated with water within thermokarst depressions and others exhibiting generally dry, but more variable soil moisture conditions outside of thermokarst depressions. Experimentally induced permafrost thaw initially drove increasing rates of growing season gross primary productivity (GPP), ecosystem respiration (Reco ), and net ecosystem exchange (NEE) (higher carbon uptake), but the formation of thermokarst depressions began to reverse this trend with a high level of spatial heterogeneity. Plots that subsided at the slowest rate stayed relatively dry and supported higher CO2 fluxes throughout the 13-year experiment, while plots that subsided very rapidly into the center of a thermokarst feature became consistently wet and experienced a rapid decline in growing season GPP, Reco , and NEE (lower carbon uptake or carbon release). These findings indicate that Earth system models, which do not simulate subsidence and often predict drier active layer conditions, likely overestimate net growing season carbon uptake in abruptly thawing landscapes.

Respiratory loss during late-growing season determines the net carbon dioxide sink in northern permafrost regions.

Warming of northern high latitude regions (NHL, > 50 °N) has increased both photosynthesis and respiration which results in considerable uncertainty regarding the net carbon dioxide (CO2) balance of NHL ecosystems. Using estimates constrained from atmospheric observations from 1980 to 2017, we find that the increasing trends of net CO2 uptake in the early-growing season are of similar magnitude across the tree cover gradient in the NHL. However, the trend of respiratory CO2 loss during late-growing season increases significantly with increasing tree cover, offsetting a larger fraction of photosynthetic CO2 uptake, and thus resulting in a slower rate of increasing annual net CO2 uptake in areas with higher tree cover, especially in central and southern boreal forest regions. The magnitude of this seasonal compensation effect explains the difference in net CO2 uptake trends along the NHL vegetation- permafrost gradient. Such seasonal compensation dynamics are not captured by dynamic global vegetation models, which simulate weaker respiration control on carbon exchange during the late-growing season, and thus calls into question projections of increasing net CO2 uptake as high latitude ecosystems respond to warming climate conditions.

Permafrost carbon feedbacks threaten global climate goals.

Rapid Arctic warming has intensified northern wildfires and is thawing carbon-rich permafrost. Carbon emissions from permafrost thaw and Arctic wildfires, which are not fully accounted for in global emissions budgets, will greatly reduce the amount of greenhouse gases that humans can emit to remain below 1.5 °C or 2 °C. The Paris Agreement provides ongoing opportunities to increase ambition to reduce society's greenhouse gas emissions, which will also reduce emissions from thawing permafrost. In December 2020, more than 70 countries announced more ambitious nationally determined contributions as part of their Paris Agreement commitments; however, the carbon budgets that informed these commitments were incomplete, as they do not fully account for Arctic feedbacks. There is an urgent need to incorporate the latest science on carbon emissions from permafrost thaw and northern wildfires into international consideration of how much more aggressively societal emissions must be reduced to address the global climate crisis.

Statistical upscaling of ecosystem CO2 fluxes across the terrestrial tundra and boreal domain: Regional patterns and uncertainties.

The regional variability in tundra and boreal carbon dioxide (CO2 ) fluxes can be high, complicating efforts to quantify sink-source patterns across the entire region. Statistical models are increasingly used to predict (i.e., upscale) CO2 fluxes across large spatial domains, but the reliability of different modeling techniques, each with different specifications and assumptions, has not been assessed in detail. Here, we compile eddy covariance and chamber measurements of annual and growing season CO2 fluxes of gross primary productivity (GPP), ecosystem respiration (ER), and net ecosystem exchange (NEE) during 1990-2015 from 148 terrestrial high-latitude (i.e., tundra and boreal) sites to analyze the spatial patterns and drivers of CO2 fluxes and test the accuracy and uncertainty of different statistical models. CO2 fluxes were upscaled at relatively high spatial resolution (1 km2 ) across the high-latitude region using five commonly used statistical models and their ensemble, that is, the median of all five models, using climatic, vegetation, and soil predictors. We found the performance of machine learning and ensemble predictions to outperform traditional regression methods. We also found the predictive performance of NEE-focused models to be low, relative to models predicting GPP and ER. Our data compilation and ensemble predictions showed that CO2 sink strength was larger in the boreal biome (observed and predicted average annual NEE -46 and -29 g C m-2  yr-1 , respectively) compared to tundra (average annual NEE +10 and -2 g C m-2  yr-1 ). This pattern was associated with large spatial variability, reflecting local heterogeneity in soil organic carbon stocks, climate, and vegetation productivity. The terrestrial ecosystem CO2 budget, estimated using the annual NEE ensemble prediction, suggests the high-latitude region was on average an annual CO2 sink during 1990-2015, although uncertainty remains high.

Author Correction: Warming shortens flowering seasons of tundra plant communities.

In the version of this Article originally published, the following sentence was missing from the Acknowledgements: "This work was supported by the Norwegian Research Council SnoEco project, grant number 230970". This text has now been added.

Drainage enhances modern soil carbon contribution but reduces old soil carbon contribution to ecosystem respiration in tundra ecosystems.

Warming temperatures are likely to accelerate permafrost thaw in the Arctic, potentially leading to the release of old carbon previously stored in deep frozen soil layers. Deeper thaw depths in combination with geomorphological changes due to the loss of ice structures in permafrost, may modify soil water distribution, creating wetter or drier soil conditions. Previous studies revealed higher ecosystem respiration rates under drier conditions, and this study investigated the cause of the increased ecosystem respiration rates using radiocarbon signatures of respired CO2 from two drying manipulation experiments: one in moist and the other in wet tundra. We demonstrate that higher contributions of CO2 from shallow soil layers (0-15 cm; modern soil carbon) drive the increased ecosystem respiration rates, while contributions from deeper soil (below 15 cm from surface and down to the permafrost table; old soil carbon) decreased. These changes can be attributed to more aerobic conditions in shallow soil layers, but also the soil temperature increases in shallow layers but decreases in deep layers, due to the altered thermal properties of organic soils. Decreased abundance of aerenchymatous plant species following drainage in wet tundra reduced old carbon release but increased aboveground plant biomass elevated contributions of autotrophic respiration to ecosystem respiration. The results of this study suggest that drier soils following drainage may accelerate decomposition of modern soil carbon in shallow layers but slow down decomposition of old soil carbon in deep layers, which may offset some of the old soil carbon loss from thawing permafrost.

Large loss of CO2 in winter observed across the northern permafrost region.

Recent warming in the Arctic, which has been amplified during the winter1-3, greatly enhances microbial decomposition of soil organic matter and subsequent release of carbon dioxide (CO2)4. However, the amount of CO2 released in winter is highly uncertain and has not been well represented by ecosystem models or by empirically-based estimates5,6. Here we synthesize regional in situ observations of CO2 flux from arctic and boreal soils to assess current and future winter carbon losses from the northern permafrost domain. We estimate a contemporary loss of 1662 Tg C yr-1 from the permafrost region during the winter season (October through April). This loss is greater than the average growing season carbon uptake for this region estimated from process models (-1032 Tg C yr-1). Extending model predictions to warmer conditions in 2100 indicates that winter CO2 emissions will increase 17% under a moderate mitigation scenario-Representative Concentration Pathway (RCP) 4.5-and 41% under business-as-usual emissions scenario-RCP 8.5. Our results provide a new baseline for winter CO2 emissions from northern terrestrial regions and indicate that enhanced soil CO2 loss due to winter warming may offset growing season carbon uptake under future climatic conditions.

Strengthened scientific support for the Endangerment Finding for atmospheric greenhouse gases.

We assess scientific evidence that has emerged since the U.S. Environmental Protection Agency's 2009 Endangerment Finding for six well-mixed greenhouse gases and find that this new evidence lends increased support to the conclusion that these gases pose a danger to public health and welfare. Newly available evidence about a wide range of observed and projected impacts strengthens the association between the risk of some of these impacts and anthropogenic climate change, indicates that some impacts or combinations of impacts have the potential to be more severe than previously understood, and identifies substantial risk of additional impacts through processes and pathways not considered in the Endangerment Finding.

Warming shortens flowering seasons of tundra plant communities.

Advancing phenology is one of the most visible effects of climate change on plant communities, and has been especially pronounced in temperature-limited tundra ecosystems. However, phenological responses have been shown to differ greatly between species, with some species shifting phenology more than others. We analysed a database of 42,689 tundra plant phenological observations to show that warmer temperatures are leading to a contraction of community-level flowering seasons in tundra ecosystems due to a greater advancement in the flowering times of late-flowering species than early-flowering species. Shorter flowering seasons with a changing climate have the potential to alter trophic interactions in tundra ecosystems. Interestingly, these findings differ from those of warmer ecosystems, where early-flowering species have been found to be more sensitive to temperature change, suggesting that community-level phenological responses to warming can vary greatly between biomes.

Fire severity effects on soil carbon and nutrients and microbial processes in a Siberian larch forest.

Fire frequency and severity are increasing in tundra and boreal regions as climate warms, which can directly affect climate feedbacks by increasing carbon (C) emissions from combustion of the large soil C pool and indirectly via changes in vegetation, permafrost thaw, hydrology, and nutrient availability. To better understand the direct and indirect effects of changing fire regimes in northern ecosystems, we examined how differences in soil burn severity (i.e., extent of soil organic matter combustion) affect soil C, nitrogen (N), and phosphorus (P) availability and microbial processes over time. We created experimental burns of three fire severities (low, moderate, and high) in a larch forest in the northeastern Siberian Arctic and analyzed soils at 1, 8 days, and 1 year postfire. Labile dissolved C and N increased with increasing soil burn severity immediately (1 day) postfire by up to an order of magnitude, but declined significantly 1 week later; both variables were comparable or lower than unburned soils by 1 year postfire. Soil burn severity had no effect on P in the organic layer, but P increased with increasing severity in mineral soil horizons. Most extracellular enzyme activities decreased by up to 70% with increasing soil burn severity. Increasing soil burn severity reduced soil respiration 1 year postfire by 50%. However, increasing soil burn severity increased net N mineralization rates 1 year postfire, which were 10-fold higher in the highest burn severity. While fires of high severity consumed approximately five times more soil C than those of low severity, soil C pools will also be driven by indirect effects of fire on soil processes. Our data suggest that despite an initial increase in labile C and nutrients with soil burn severity, soil respiration and extracellular activities related to the turnover of organic matter were greatly reduced, which may mitigate future C losses following fire.

Biotic responses buffer warming-induced soil organic carbon loss in Arctic tundra.

Climate warming can result in both abiotic (e.g., permafrost thaw) and biotic (e.g., microbial functional genes) changes in Arctic tundra. Recent research has incorporated dynamic permafrost thaw in Earth system models (ESMs) and indicates that Arctic tundra could be a significant future carbon (C) source due to the enhanced decomposition of thawed deep soil C. However, warming-induced biotic changes may influence biologically related parameters and the consequent projections in ESMs. How model parameters associated with biotic responses will change under warming and to what extent these changes affect projected C budgets have not been carefully examined. In this study, we synthesized six data sets over 5 years from a soil warming experiment at the Eight Mile Lake, Alaska, into the Terrestrial ECOsystem (TECO) model with a probabilistic inversion approach. The TECO model used multiple soil layers to track dynamics of thawed soil under different treatments. Our results show that warming increased light use efficiency of vegetation photosynthesis but decreased baseline (i.e., environment-corrected) turnover rates of SOC in both the fast and slow pools in comparison with those under control. Moreover, the parameter changes generally amplified over time, suggesting processes of gradual physiological acclimation and functional gene shifts of both plants and microbes. The TECO model predicted that field warming from 2009 to 2013 resulted in cumulative C losses of 224 or 87 g/m2 , respectively, without or with changes in those parameters. Thus, warming-induced parameter changes reduced predicted soil C loss by 61%. Our study suggests that it is critical to incorporate biotic changes in ESMs to improve the model performance in predicting C dynamics in permafrost regions.

Understory vegetation mediates permafrost active layer dynamics and carbon dioxide fluxes in open-canopy larch forests of northeastern Siberia.

Arctic ecosystems are characterized by a broad range of plant functional types that are highly heterogeneous at small (~1-2 m) spatial scales. Climatic changes can impact vegetation distribution directly, and also indirectly via impacts on disturbance regimes. Consequent changes in vegetation structure and function have implications for surface energy dynamics that may alter permafrost thermal dynamics, and are therefore of interest in the context of permafrost related climate feedbacks. In this study we examine small-scale heterogeneity in soil thermal properties and ecosystem carbon and water fluxes associated with varying understory vegetation in open-canopy larch forests in northeastern Siberia. We found that lichen mats comprise 16% of understory vegetation cover on average in open canopy larch forests, and lichen abundance was inversely related to canopy cover. Relative to adjacent areas dominated by shrubs and moss, lichen mats had 2-3 times deeper permafrost thaw depths and surface soils warmer by 1-2°C in summer and less than 1°C in autumn. Despite deeper thaw depths, ecosystem respiration did not differ across vegetation types, indicating that autotrophic respiration likely dominates areas with shrubs and moss. Summertime net ecosystem exchange of CO2 was negative (i.e. net uptake) in areas with high shrub cover, while positive (i.e. net loss) in lichen mats and areas with less shrub cover. Our results highlight relationships between vegetation and soil thermal dynamics in permafrost ecosystems, and underscore the necessity of considering both vegetation and permafrost dynamics in shaping carbon cycling in permafrost ecosystems.

Warming effects on permafrost ecosystem carbon fluxes associated with plant nutrients.

Large uncertainties exist in carbon (C)-climate feedback in permafrost regions, partly due to an insufficient understanding of warming effects on nutrient availabilities and their subsequent impacts on vegetation C sequestration. Although a warming climate may promote a substantial release of soil C to the atmosphere, a warming-induced increase in soil nutrient availability may enhance plant productivity, thus offsetting C loss from microbial respiration. Here, we present evidence that the positive temperature effect on carbon dioxide (CO2 ) fluxes may be weakened by reduced plant nitrogen (N) and phosphorous (P) concentrations in a Tibetan permafrost ecosystem. Although experimental warming initially enhanced ecosystem CO2 uptake, the increased rate disappeared after the period of peak plant growth during the early growing season, even though soil moisture was not a limiting factor in this swamp meadow ecosystem. We observed that warming did not significantly affect soil extractable N or P during the period of peak growth, but decreased both N and P concentrations in the leaves of dominant plant species, likely caused by accelerated plant senescence in the warmed plots. The attenuated warming effect on CO2 assimilation during the late growing season was associated with lowered leaf N and P concentrations. These findings suggest that warming-mediated nutrient changes may not always benefit ecosystem C uptake in permafrost regions, making our ability to predict the C balance in these warming-sensitive ecosystems more challenging than previously thought.

Nonlinear CO2 flux response to 7 years of experimentally induced permafrost thaw.

Rapid Arctic warming is expected to increase global greenhouse gas concentrations as permafrost thaw exposes immense stores of frozen carbon (C) to microbial decomposition. Permafrost thaw also stimulates plant growth, which could offset C loss. Using data from 7 years of experimental Air and Soil warming in moist acidic tundra, we show that Soil warming had a much stronger effect on CO2 flux than Air warming. Soil warming caused rapid permafrost thaw and increased ecosystem respiration (Reco ), gross primary productivity (GPP), and net summer CO2 storage (NEE). Over 7 years Reco , GPP, and NEE also increased in Control (i.e., ambient plots), but this change could be explained by slow thaw in Control areas. In the initial stages of thaw, Reco , GPP, and NEE increased linearly with thaw across all treatments, despite different rates of thaw. As thaw in Soil warming continued to increase linearly, ground surface subsidence created saturated microsites and suppressed Reco , GPP, and NEE. However Reco and GPP remained high in areas with large Eriophorum vaginatum biomass. In general NEE increased with thaw, but was more strongly correlated with plant biomass than thaw, indicating that higher Reco in deeply thawed areas during summer months was balanced by GPP. Summer CO2 flux across treatments fit a single quadratic relationship that captured the functional response of CO2 flux to thaw, water table depth, and plant biomass. These results demonstrate the importance of indirect thaw effects on CO2 flux: plant growth and water table dynamics. Nonsummer Reco models estimated that the area was an annual CO2 source during all years of observation. Nonsummer CO2 loss in warmer, more deeply thawed soils exceeded the increases in summer GPP, and thawed tundra was a net annual CO2 source.

Greater temperature sensitivity of plant phenology at colder sites: implications for convergence across northern latitudes.

Warmer temperatures are accelerating the phenology of organisms around the world. Temperature sensitivity of phenology might be greater in colder, higher latitude sites than in warmer regions, in part because small changes in temperature constitute greater relative changes in thermal balance at colder sites. To test this hypothesis, we examined up to 20 years of phenology data for 47 tundra plant species at 18 high-latitude sites along a climatic gradient. Across all species, the timing of leaf emergence and flowering was more sensitive to a given increase in summer temperature at colder than warmer high-latitude locations. A similar pattern was seen over time for the flowering phenology of a widespread species, Cassiope tetragona. These are among the first results highlighting differential phenological responses of plants across a climatic gradient and suggest the possibility of convergence in flowering times and therefore an increase in gene flow across latitudes as the climate warms.

Nitrogen availability increases in a tundra ecosystem during five years of experimental permafrost thaw.

Perennially frozen soil in high latitude ecosystems (permafrost) currently stores 1330-1580 Pg of carbon (C). As these ecosystems warm, the thaw and decomposition of permafrost is expected to release large amounts of C to the atmosphere. Fortunately, losses from the permafrost C pool will be partially offset by increased plant productivity. The degree to which plants are able to sequester C, however, will be determined by changing nitrogen (N) availability in these thawing soil profiles. N availability currently limits plant productivity in tundra ecosystems but plant access to N is expected improve as decomposition increases in speed and extends to deeper soil horizons. To evaluate the relationship between permafrost thaw and N availability, we monitored N cycling during 5 years of experimentally induced permafrost thaw at the Carbon in Permafrost Experimental Heating Research (CiPEHR) project. Inorganic N availability increased significantly in response to deeper thaw and greater soil moisture induced by Soil warming. This treatment also prompted a 23% increase in aboveground biomass and a 49% increase in foliar N pools. The sedge Eriophorum vaginatum responded most strongly to warming: this species explained 91% of the change in aboveground biomass during the 5 year period. Air warming had little impact when applied alone, but when applied in combination with Soil warming, growing season soil inorganic N availability was significantly reduced. These results demonstrate that there is a strong positive relationship between the depth of permafrost thaw and N availability in tundra ecosystems but that this relationship can be diminished by interactions between increased thaw, warmer air temperatures, and higher levels of soil moisture. Within 5 years of permafrost thaw, plants actively incorporate newly available N into biomass but C storage in live vascular plant biomass is unlikely to be greater than losses from deep soil C pools.

Decadal warming causes a consistent and persistent shift from heterotrophic to autotrophic respiration in contrasting permafrost ecosystems.

Soil carbon in permafrost ecosystems has the potential to become a major positive feedback to climate change if permafrost thaw increases heterotrophic decomposition. However, warming can also stimulate autotrophic production leading to increased ecosystem carbon storage-a negative climate change feedback. Few studies partitioning ecosystem respiration examine decadal warming effects or compare responses among ecosystems. Here, we first examined how 11 years of warming during different seasons affected autotrophic and heterotrophic respiration in a bryophyte-dominated peatland in Abisko, Sweden. We used natural abundance radiocarbon to partition ecosystem respiration into autotrophic respiration, associated with production, and heterotrophic decomposition. Summertime warming decreased the age of carbon respired by the ecosystem due to increased proportional contributions from autotrophic and young soil respiration and decreased proportional contributions from old soil. Summertime warming's large effect was due to not only warmer air temperatures during the growing season, but also to warmer deep soils year-round. Second, we compared ecosystem respiration responses between two contrasting ecosystems, the Abisko peatland and a tussock-dominated tundra in Healy, Alaska. Each ecosystem had two different timescales of warming (<5 years and over a decade). Despite the Abisko peatland having greater ecosystem respiration and larger contributions from heterotrophic respiration than the Healy tundra, both systems responded consistently to short- and long-term warming with increased respiration, increased autotrophic contributions to ecosystem respiration, and increased ratios of autotrophic to heterotrophic respiration. We did not detect an increase in old soil carbon losses with warming at either site. If increased autotrophic respiration is balanced by increased primary production, as is the case in the Healy tundra, warming will not cause these ecosystems to become growing season carbon sources. Warming instead causes a persistent shift from heterotrophic to more autotrophic control of the growing season carbon cycle in these carbon-rich permafrost ecosystems.

A pan-Arctic synthesis of CH4 and CO2 production from anoxic soil incubations.

Permafrost thaw can alter the soil environment through changes in soil moisture, frequently resulting in soil saturation, a shift to anaerobic decomposition, and changes in the plant community. These changes, along with thawing of previously frozen organic material, can alter the form and magnitude of greenhouse gas production from permafrost ecosystems. We synthesized existing methane (CH4 ) and carbon dioxide (CO2 ) production measurements from anaerobic incubations of boreal and tundra soils from the geographic permafrost region to evaluate large-scale controls of anaerobic CO2 and CH4 production and compare the relative importance of landscape-level factors (e.g., vegetation type and landscape position), soil properties (e.g., pH, depth, and soil type), and soil environmental conditions (e.g., temperature and relative water table position). We found fivefold higher maximum CH4 production per gram soil carbon from organic soils than mineral soils. Maximum CH4 production from soils in the active layer (ground that thaws and refreezes annually) was nearly four times that of permafrost per gram soil carbon, and CH4 production per gram soil carbon was two times greater from sites without permafrost than sites with permafrost. Maximum CH4 and median anaerobic CO2 production decreased with depth, while CO2 :CH4 production increased with depth. Maximum CH4 production was highest in soils with herbaceous vegetation and soils that were either consistently or periodically inundated. This synthesis identifies the need to consider biome, landscape position, and vascular/moss vegetation types when modeling CH4 production in permafrost ecosystems and suggests the need for longer-term anaerobic incubations to fully capture CH4 dynamics. Our results demonstrate that as climate warms in arctic and boreal regions, rates of anaerobic CO2 and CH4 production will increase, not only as a result of increased temperature, but also from shifts in vegetation and increased ground saturation that will accompany permafrost thaw.

Permafrost degradation stimulates carbon loss from experimentally warmed tundra.

A large pool of organic carbon (C) has been accumulating in the Arctic for thousands of years because cold and waterlogged conditions have protected soil organic material from microbial decomposition. As the climate warms this vast and frozen C pool is at risk of being thawed, decomposed, and released to the atmosphere as greenhouse gasses. At the same time, some C losses may be offset by warming-mediated increases in plant productivity. Plant and microbial responses to warming ultimately determine net C exchange from ecosystems, but the timing and magnitude of these responses remain uncertain. Here we show that experimental warming and permafrost (ground that remains below 0 degrees C for two or more consecutive years) degradation led to a two-fold increase in net ecosystem C uptake during the growing season. However, warming also enhanced winter respiration, which entirely offset growing-season C gains. Winter C losses may be even higher in response to actual climate warming than to our experimental manipulations, and, in that scenario, could be expected to more than double overall net C losses from tundra to the atmosphere. Our results highlight the importance of winter processes in determining whether tundra acts as a C source or sink, and demonstrate the potential magnitude of C release from the permafrost zone that might be expected in a warmer climate.

Plant-soil distribution of potentially toxic elements in response to elevated atmospheric CO2.

The distribution of contaminant elements within ecosystems is an environmental concern because of these elements' potential toxicity to animals and plants and their ability to hinder microbial ecosystem services. As with nutrients, contaminants are cycled within and through ecosystems. Elevated atmospheric CO2 generally increases plant productivity and alters nutrient element cycling, but whether CO2 causes similar effects on the cycling of contaminant elements is unknown. Here we show that 11 years of experimental CO2 enrichment in a sandy soil with low organic matter content causes plants to accumulate contaminants in plant biomass, with declines in the extractable contaminant element pools in surface soils. These results indicate that CO2 alters the distribution of contaminant elements in ecosystems, with plant element accumulation and declining soil availability both likely explained by the CO2 stimulation of plant biomass. Our results highlight the interdependence of element cycles and the importance of taking a broad view of the periodic table when the effects of global environmental change on ecosystem biogeochemistry are considered.