Potential of APSIS-InSAR for measuring surface oscillations of tropical peatlands.

Tropical peatland across Southeast Asia is drained extensively for production of pulpwood, palm oil and other food crops. Associated increases in peat decomposition have led to widespread subsidence, deterioration of peat condition and CO2 emissions. However, quantification of subsidence and peat condition from these processes is challenging due to the scale and inaccessibility of dense tropical peat swamp forests. The development of satellite interferometric synthetic aperture radar (InSAR) has the potential to solve this problem. The Advanced Pixel System using Intermittent Baseline Subset (APSIS, formerly ISBAS) modelling technique provides improved coverage across almost all land surfaces irrespective of ground cover, enabling derivation of a time series of tropical peatland surface oscillations across whole catchments. This study aimed to establish the extent to which APSIS-InSAR can monitor seasonal patterns of tropical peat surface oscillations at North Selangor Peat Swamp Forest, Peninsular Malaysia. Results showed that C-band SAR could penetrate the forest canopy over tropical peat swamp forests intermittently and was applicable to a range of land covers. Therefore the APSIS technique has the potential for monitoring peat surface oscillations under tropical forest canopy using regularly acquired C-band Sentinel-1 InSAR data, enabling continuous monitoring of tropical peatland surface motion at a spatial resolution of 20 m.

Simulating carbon accumulation and loss in the central Congo peatlands.

Peatlands of the central Congo Basin have accumulated carbon over millennia. They currently store some 29 billion tonnes of carbon in peat. However, our understanding of the controls on peat carbon accumulation and loss and the vulnerability of this stored carbon to climate change is in its infancy. Here we present a new model of tropical peatland development, DigiBog_Congo, that we use to simulate peat carbon accumulation and loss in a rain-fed interfluvial peatland that began forming ~20,000 calendar years Before Present (cal. yr BP, where 'present' is 1950 CE). Overall, the simulated age-depth curve is in good agreement with palaeoenvironmental reconstructions derived from a peat core at the same location as our model simulation. We find two key controls on long-term peat accumulation: water at the peat surface (surface wetness) and the very slow anoxic decay of recalcitrant material. Our main simulation shows that between the Late Glacial and early Holocene there were several multidecadal periods where net peat and carbon gain alternated with net loss. Later, a climatic dry phase beginning ~5200 cal. yr BP caused the peatland to become a long-term carbon source from ~3975 to 900 cal. yr BP. Peat as old as ~7000 cal. yr BP was decomposed before the peatland's surface became wetter again, suggesting that changes in rainfall alone were sufficient to cause a catastrophic loss of peat carbon lasting thousands of years. During this time, 6.4 m of the column of peat was lost, resulting in 57% of the simulated carbon stock being released. Our study provides an approach to understanding the future impact of climate change and potential land-use change on this vulnerable store of carbon.

CH4 and N2 O emissions from smallholder agricultural systems on tropical peatlands in Southeast Asia.

There are limited data for greenhouse gas (GHG) emissions from smallholder agricultural systems in tropical peatlands, with data for non-CO2 emissions from human-influenced tropical peatlands particularly scarce. The aim of this study was to quantify soil CH4 and N2 O fluxes from smallholder agricultural systems on tropical peatlands in Southeast Asia and assess their environmental controls. The study was carried out in four regions in Malaysia and Indonesia. CH4 and N2 O fluxes and environmental parameters were measured in cropland, oil palm plantation, tree plantation and forest. Annual CH4 emissions (in kg CH4 ha-1 year-1 ) were: 70.7 ± 29.5, 2.1 ± 1.2, 2.1 ± 0.6 and 6.2 ± 1.9 at the forest, tree plantation, oil palm and cropland land-use classes, respectively. Annual N2 O emissions (in kg N2 O ha-1 year-1 ) were: 6.5 ± 2.8, 3.2 ± 1.2, 21.9 ± 11.4 and 33.6 ± 7.3 in the same order as above, respectively. Annual CH4 emissions were strongly determined by water table depth (WTD) and increased exponentially when annual WTD was above -25 cm. In contrast, annual N2 O emissions were strongly correlated with mean total dissolved nitrogen (TDN) in soil water, following a sigmoidal relationship, up to an apparent threshold of 10 mg N L-1 beyond which TDN seemingly ceased to be limiting for N2 O production. The new emissions data for CH4 and N2 O presented here should help to develop more robust country level 'emission factors' for the quantification of national GHG inventory reporting. The impact of TDN on N2 O emissions suggests that soil nutrient status strongly impacts emissions, and therefore, policies which reduce N-fertilisation inputs might contribute to emissions mitigation from agricultural peat landscapes. However, the most important policy intervention for reducing emissions is one that reduces the conversion of peat swamp forest to agriculture on peatlands in the first place.

Detecting tropical peatland degradation: Combining remote sensing and organic geochemistry.

Tropical peatlands are important carbon stores that are vulnerable to drainage and conversion to agriculture. Protection and restoration of peatlands are increasingly recognised as key nature based solutions that can be implemented as part of climate change mitigation. Identification of peatland areas that are important for protection and restauration with regards to the state of their carbon stocks, are therefore vital for policy makers. In this paper we combined organic geochemical analysis by Rock-Eval (6) pyrolysis of peat collected from sites with different land management history and optical remote sensing products to assess if remotely sensed data could be used to predict peat conditions and carbon storage. The study used the North Selangor Peat Swamp forest, Malaysia, as the model system. Across the sampling sites the carbon stocks in the below ground peat was ca 12 times higher than the forest (median carbon stock held in ground vegetation 114.70 Mg ha-1 and peat soil 1401.51 Mg ha-1). Peat core sub-samples and litter collected from Fire Affected, Disturbed Forest, and Managed Recovery locations (i.e. disturbed sites) had different decomposition profiles than Central Forest sites. The Rock-Eval pyrolysis of the upper peat profiles showed that surface peat layers at Fire Affected, Disturbed Forest, and Managed Recovery locations had lower immature organic matter index (I-index) values (average I-index range in upper section 0.15 to -0.06) and higher refractory organic matter index (R -index) (average R-index range in upper section 0.51 to 0.65) compared to Central Forest sites indicating enhanced decomposition of the surface peat. In the top 50 cm section of the peat profile, carbon stocks were negatively related to the normalised burns ratio (NBR) (a satellite derived parameter) (Spearman's rho = -0.664, S = 366, p-value = <0.05) while there was a positive relationship between the hydrogen index and the normalised burns ratio profile (Spearman's rho = 0.7, S = 66, p-value = <0.05) suggesting that this remotely sensed product is able to detect degradation of peat in the upper peat profile. We conclude that the NBR can be used to identify degraded peatland areas and to support identification of areas for conversation and restoration.

Hydroclimatic vulnerability of peat carbon in the central Congo Basin.

The forested swamps of the central Congo Basin store approximately 30 billion metric tonnes of carbon in peat1,2. Little is known about the vulnerability of these carbon stocks. Here we investigate this vulnerability using peat cores from a large interfluvial basin in the Republic of the Congo and palaeoenvironmental methods. We find that peat accumulation began at least at 17,500 calibrated years before present (cal. yr BP; taken as AD 1950). Our data show that the peat that accumulated between around 7,500 to around 2,000 cal. yr BP is much more decomposed compared with older and younger peat. Hydrogen isotopes of plant waxes indicate a drying trend, starting at approximately 5,000 cal. yr BP and culminating at approximately 2,000 cal. yr BP, coeval with a decline in dominant swamp forest taxa. The data imply that the drying climate probably resulted in a regional drop in the water table, which triggered peat decomposition, including the loss of peat carbon accumulated prior to the onset of the drier conditions. After approximately 2,000 cal. yr BP, our data show that the drying trend ceased, hydrologic conditions stabilized and peat accumulation resumed. This reversible accumulation-loss-accumulation pattern is consistent with other peat cores across the region, indicating that the carbon stocks of the central Congo peatlands may lie close to a climatically driven drought threshold. Further research should quantify the combination of peatland threshold behaviour and droughts driven by anthropogenic carbon emissions that may trigger this positive carbon cycle feedback in the Earth system.

Temporal changes in litterfall and potential nutrient return in cocoa agroforestry systems under organic and conventional management, Ghana.

Litterfall is a critical link between vegetation and soils by which nutrients are returned to the soils, thus the amount and pattern of litterfall regulates nutrient cycling, soil fertility and primary productivity for most terrestrial ecosystems. We quantified, analyzed and compared macro- and micro-nutrients return through litterfall in organic and conventional cocoa agroforestry systems in Suhum, Ghana. We further assessed the contribution of shade tree species to litterfall and nutrient dynamics. The annual pattern of litterfall was affected by seasonality, with a major peak in the dry season and minor peaks during the rainy season. In terms of annual fractional litterfall, mean leaf litter from shade tree species was significantly higher (50 %) in organic systems (5.0 ± 0.5 Mg ha-1 yr-1) compared to conventional systems (3.3 ± 0.6 Mg ha-1 yr-1). Whereas cocoa leaves (45.0 %) were the predominant fraction of annual litterfall from conventional farms, both shade leaves (40.0 %) and cocoa leaves (39.4 %) dominated litterfall from organic farms. The return of primary macro-nutrients (P and K), secondary macro-nutrients (Ca, Mg and S) and micro-nutrients (Mn, B, Cu, Zn and Mo) via litterfall varied significantly with season, and annual return of nutrients were similar in organic and conventional cocoa systems. Shade tree leaf litter accounted for 30-47 % of annual macro- and micro-nutrient return (except Ni and Zn) in organic cocoa systems versus 20-35 % in conventional cocoa systems. The results emphasize the complementary role of the different shade tree species which compose organic and conventional cocoa systems in nutrient recycling. We conclude that organic management of cocoa agroforestry systems ensure nutrients return similar to those receiving synthetic fertilizer inputs, highlighting its potential to support cocoa production.

Author Correction: Greenhouse gas emissions resulting from conversion of peat swamp forest to oil palm plantation.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Greenhouse gas emissions resulting from conversion of peat swamp forest to oil palm plantation.

Conversion of tropical peat swamp forest to drainage-based agriculture alters greenhouse gas (GHG) production, but the magnitude of these changes remains highly uncertain. Current emissions factors for oil palm grown on drained peat do not account for temporal variation over the plantation cycle and only consider CO2 emissions. Here, we present direct measurements of GHGs emitted during the conversion from peat swamp forest to oil palm plantation, accounting for CH4 and N2O as well as CO2. Our results demonstrate that emissions factors for converted peat swamp forest is in the range 70-117 t CO2 eq ha-1 yr-1 (95% confidence interval, CI), with CO2 and N2O responsible for ca. 60 and ca. 40% of this value, respectively. These GHG emissions suggest that conversion of Southeast Asian peat swamp forest is contributing between 16.6 and 27.9% (95% CI) of combined total national GHG emissions from Malaysia and Indonesia or 0.44 and 0.74% (95% CI) of annual global emissions.

Methane emissions from tree stems in neotropical peatlands.

Neotropical peatlands emit large amounts of methane (CH4 ) from the soil surface, but fluxes from tree stems in these ecosystems are unknown. In this study we investigated CH4 emissions from five tree species in two forest types common to neotropical lowland peatlands in Panama. Methane from tree stems accounted for up to 30% of net ecosystem CH4 emissions. Peak CH4 fluxes were greater during the wet season when the water table was high and temperatures were lower. Emissions were greatest from the hardwood tree Campnosperma panamensis, but most species acted as emitters, with emissions declining exponentially with height along the stem for all species. Overall, species identity, stem diameter, water level, soil temperature and soil CH4 fluxes explained 54% of the variance in stem CH4 emissions from individual trees. On the landscape level, On the landscape level, the high emissions from C. panamensis forests resulted in that they emitted at 340 kg CH4  d-1 during flooded periods despite their substantially lower areal cover. We conclude that emission from tree stems is an important emission pathway for CH4 flux from Neotropical peatlands, and that these emissions vary strongly with season and forest type.

Evaluation of vegetation communities, water table, and peat composition as drivers of greenhouse gas emissions in lowland tropical peatlands.

Tropical peatlands are globally important source of greenhouse gases to the atmosphere, but data on carbon fluxes from these ecosystems is limited due to the logistical challenges of measuring gas fluxes in these ecosystems. Proposals to overcome the difficulties of measuring gas carbon fluxes in the tropics include remote sensing (top-down) approaches. However, these require information on the effect of vegetation communities on carbon dioxide (CO2) and methane (CH4) fluxes from the peat surface (bottom-up). Such information will help reducing the uncertainty in current carbon budgets and resolve inconsistencies between the top-down and bottom-up estimates of gas fluxes from tropical peatlands. We investigated temporal and spatial variability of CO2 and CH4 fluxes from tropical peatlands inhabited by two contrasting vegetation communities (i.e., mixed forest and palm swamp) in Panama. In addition, we explored the influence of peat chemistry and nutrient status (i.e., factorial nitrogen (N) and phosphorus (P) addition) on greenhouse gas fluxes from the peat surface. We found that: i) CO2 and CH4 fluxes were not significantly different between the two vegetation communities, but did vary temporally across an annual cycle; ii) precipitation rates and peat temperature were poor predictors of CO2 and CH4 fluxes; iii) nitrogen addition increased CH4 fluxes at the mixed forests when the water table was above the peat surface, but neither nitrogen nor phosphorus affected gas fluxes elsewhere; iv) gas fluxes varied significantly with the water table level, with CO2 flux being 80% greater at low water table, and CH4 fluxes being 81% higher with the water table above the surface. Taken together, our data suggested that water table is the most important control of greenhouse gas emissions from the peat surface in forested lowland tropical peatlands, and that neither the presence of distinct vegetation communities nor the addition of nutrients outweigh such control.

Tree diversity and its ecological importance value in organic and conventional cocoa agroforests in Ghana.

Cocoa agroforestry systems have the potential to conserve biodiversity and provide environmental or ecological benefits at various nested scales ranging from the plot to ecoregion. While integrating organic practices into cocoa agroforestry may further enhance these potentials, empirical and robust data to support this claim is lacking, and mechanisms for biodiversity conservation and the provision of environmental and ecological benefits are poorly understood. A field study was conducted in the Eastern Region of Ghana to evaluate the potential of organic cocoa agroforests to conserve native floristic diversity in comparison with conventional cocoa agroforests. Shade tree species richness, Shannon, Simpson's reciprocal and Margalef diversity indices were estimated from 84 organic and conventional cocoa agroforestry plots. Species importance value index, a measure of how dominant a species is in a given ecosystem, and conservation status were used to evaluate the conservation potential of shade trees on studied cocoa farms. Organic farms recorded higher mean shade tree species richness (5.10 ± 0.38) compared to conventional farms (3.48 ± 0.39). Similarly, mean Shannon diversity index, Simpson's reciprocal diversity index and Margalef diversity index were significantly higher on organic farms compared to conventional farms. According to the importance value index, fruit and native shade tree species were the most important on both organic and conventional farms for all the cocoa age groups but more so on organic farms. Organic farms maintained 14 native tree species facing a conservation issue compared to 10 on conventional cocoa farms. The results suggest that diversified organic cocoa farms can serve as reservoirs of native tree species, including those currently facing conservation concerns thereby providing support and contributing to the conservation of tree species in the landscape.

Are secondary forests second-rate? Comparing peatland greenhouse gas emissions, chemical and microbial community properties between primary and secondary forests in Peninsular Malaysia.

Tropical peatlands are globally important ecosystems with high C storage and are endangered by anthropogenic disturbances. Microbes in peatlands play an important role in sustaining the functions of peatlands as a C sink, yet their characteristics in these habitats are poorly understood. This research aimed to elucidate the responses of these complex ecosystems to disturbance by exploring greenhouse gas (GHG) emissions, nutrient contents, soil microbial communities and the functional interactions between these components in a primary and secondary peat swamp forest in Peninsular Malaysia. GHG measurements using closed chambers, and peat sampling were carried out in both wet and dry seasons. Microbial community phenotypes and nutrient content were determined using phospholipid fatty acid (PLFA) and inductively-coupled plasma mass spectrometry (ICP-MS) analyses respectively. CO2 emissions in the secondary peat swamp forest were > 50% higher than in the primary forest. CH4 emission rates were ca. 2 mg m-2 h-1 in the primary forest but the secondary forest was a CH4 sink, showing no seasonal variations in GHG emissions. Almost all the nutrient concentrations were significantly lower in the secondary forest, postulated to be due to nutrient leaching via drainage and higher rates of decomposition. Cu and Mo concentrations were negatively correlated with CO2 and CH4 emissions respectively. Microbial community structure was overwhelmingly dominated by bacteria in both forest types, however it was highly sensitive to land-use change and season. Gram-positive and Gram-negative relative abundance were positively correlated with CO2 and CH4 emissions respectively. Drainage related disturbances increased CO2 emissions, by reducing the nutrient content including some with known antimicrobial properties (Cu & Na) and by favouring Gram-positive bacteria over Gram-negative bacteria. These results suggest that the biogeochemistry of secondary peat swamp forest is fundamentally different from that of primary peat swamp forest, and these differences have significant functional impacts on their respective environments.

Exploring drivers of litter decomposition in a greening Arctic: results from a transplant experiment across a treeline.

Decomposition of plant litter is a key control over carbon (C) storage in the soil. The biochemistry of the litter being produced, the environment in which the decomposition is taking place, and the community composition and metabolism of the decomposer organisms exert a combined influence over decomposition rates. As deciduous shrubs and trees are expanding into tundra ecosystems as a result of regional climate warming, this change in vegetation represents a change in litter input to tundra soils and a change in the environment in which litter decomposes. To test the importance of litter biochemistry and environment in determining litter mass loss, we reciprocally transplanted litter between heath (Empetrum nigrum), shrub (Betula nana), and forest (Betula pubescens) at a sub-Arctic treeline in Sweden. As expansion of shrubs and trees promotes deeper snow, we also used a snow fence experiment in a tundra heath environment to understand the importance of snow depth, relative to other factors, in the decomposition of litter. Our results show that B. pubescens and B. nana leaf litter decomposed at faster rates than E. nigrum litter across all environments, while all litter species decomposed at faster rates in the forest and shrub environments than in the tundra heath. The effect of increased snow on decomposition was minimal, leading us to conclude that microbial activity over summer in the productive forest and shrub vegetation is driving increased mass loss compared to the heath. Using B. pubescens and E. nigrum litter, we demonstrate that degradation of carbohydrate-C is a significant driver of mass loss in the forest. This pathway was less prominent in the heath, which is consistent with observations that tundra soils typically have high concentrations of "labile" C. This experiment suggests that further expansion of shrubs and trees may stimulate the loss of undecomposed carbohydrate C in the tundra.

To what extent can zero tillage lead to a reduction in greenhouse gas emissions from temperate soils?

Soil tillage practices have a profound influence on the physical properties of soil and the greenhouse gas (GHG) balance. However there have been very few integrated studies on the emission of carbon dioxide (CO2), methane (CH4) and nitrous oxide (N2O) and soil biophysical and chemical characteristics under different soil management systems. We recorded a significantly higher net global warming potential under conventional tillage systems (26-31% higher than zero tillage systems). Crucially the 3-D soil pore network, imaged using X-ray Computed Tomography, modified by tillage played a significant role in the flux of CO2 and CH4. In contrast, N2O flux was determined mainly by microbial biomass carbon and soil moisture content. Our work indicates that zero tillage could play a significant role in minimising emissions of GHGs from soils and contribute to efforts to mitigate against climate change.

Tropical wetlands: A missing link in the global carbon cycle?

Tropical wetlands are not included in Earth system models, despite being an important source of methane (CH4) and contributing a large fraction of carbon dioxide (CO2) emissions from land use, land use change, and forestry in the tropics. This review identifies a remarkable lack of data on the carbon balance and gas fluxes from undisturbed tropical wetlands, which limits the ability of global change models to make accurate predictions about future climate. We show that the available data on in situ carbon gas fluxes in undisturbed forested tropical wetlands indicate marked spatial and temporal variability in CO2 and CH4 emissions, with exceptionally large fluxes in Southeast Asia and the Neotropics. By upscaling short-term measurements, we calculate that approximately 90 ± 77 Tg CH4 year-1 and 4540 ± 1480 Tg CO2 year-1 are released from tropical wetlands globally. CH4 fluxes are greater from mineral than organic soils, whereas CO2 fluxes do not differ between soil types. The high CO2 and CH4 emissions are mirrored by high rates of net primary productivity and litter decay. Net ecosystem productivity was estimated to be greater in peat-forming wetlands than on mineral soils, but the available data are insufficient to construct reliable carbon balances or estimate gas fluxes at regional scales. We conclude that there is an urgent need for systematic data on carbon dynamics in tropical wetlands to provide a robust understanding of how they differ from well-studied northern wetlands and allow incorporation of tropical wetlands into global climate change models.

Environmental controls of temporal and spatial variability in CO2 and CH4 fluxes in a neotropical peatland.

Tropical peatlands play an important role in the global storage and cycling of carbon (C) but information on carbon dioxide (CO2) and methane (CH4) fluxes from these systems is sparse, particularly in the Neotropics. We quantified short and long-term temporal and small scale spatial variation in CO2 and CH4 fluxes from three contrasting vegetation communities in a domed ombrotrophic peatland in Panama. There was significant variation in CO2 fluxes among vegetation communities in the order Campnosperma panamensis > Raphia taedigera > Cyperus. There was no consistent variation among sites and no discernible seasonal pattern of CH4 flux despite the considerable range of values recorded (e.g. -1.0 to 12.6 mg m(-2) h(-1) in 2007). CO2 fluxes varied seasonally in 2007, being greatest in drier periods (300-400 mg m(-2) h(-1)) and lowest during the wet period (60-132 mg m(-2) h(-1)) while very high emissions were found during the 2009 wet period, suggesting that peak CO2 fluxes may occur following both low and high rainfall. In contrast, only weak relationships between CH4 flux and rainfall (positive at the C. panamensis site) and solar radiation (negative at the C. panamensis and Cyperus sites) was found. CO2 fluxes showed a diurnal pattern across sites and at the Cyperus sp. site CO2 and CH4 fluxes were positively correlated. The amount of dissolved carbon and nutrients were strong predictors of small scale within-site variability in gas release but the effect was site-specific. We conclude that (i) temporal variability in CO2 was greater than variation among vegetation communities; (ii) rainfall may be a good predictor of CO2 emissions from tropical peatlands but temporal variation in CH4 does not follow seasonal rainfall patterns; and (iii) diurnal variation in CO2 fluxes across different vegetation communities can be described by a Fourier model.

Potential impact of CO2 leakage from carbon capture and storage systems on field bean (Vicia faba).

Capture and geological storage of carbon dioxide (CO(2)) has been suggested to be essential to reduce emissions to the atmosphere and aid mitigation of global climate change. However, leakage from transport pipelines or carbon capture and storage (CCS) reservoirs may pose risks to vegetation and contribute to rising atmospheric concentrations [CO(2)]. This study examined effects on seedling emergence and growth when field bean plants (Vicia faba cv. Wizard) grown under field conditions were subjected to elevated soil [CO(2)] for 39 days after planting. The strong negative correlation between soil [CO(2)] and [O(2)] during the injection period created a hypoxic soil environment under conditions of elevated soil [CO(2)]. The damaging impact of this treatment became apparent early in the experiment as no seeds exposed to soil [CO(2)] >50% emerged, even after injection was discontinued. Some seeds exposed to soil [CO(2)] <50% produced seedlings, but many did not survive. Seedling emergence and survival in the gassed plots was greatest at [CO(2)] of 5-20%, but root and shoot growth was reduced relative to control plants. Seedling emergence and growth were negatively related to soil [CO(2)] and positively related to [O(2)], although it is not known which was more important in inducing the observed effects. These findings suggest that leakage of CO(2) from transport pipelines or CCS sites may greatly reduce seedling emergence and crop establishment in the vicinity of such infrastructures.

Recovery of ecosystem carbon fluxes and storage from herbivory.

The carbon (C) sink strength of arctic tundra is under pressure from increasing populations of arctic breeding geese. In this study we examined how CO2 and CH4 fluxes, plant biomass and soil C responded to the removal of vertebrate herbivores in a high arctic wet moss meadow that has been intensively used by barnacle geese (Branta leucopsis) for ca. 20 years. We used 4 and 9 years old grazing exclosures to investigate the potential for recovery of ecosystem function during the growing season (July 2007). The results show greater above- and below-ground vascular plant biomass within the grazing exclosures with graminoid biomass being most responsive to the removal of herbivory whilst moss biomass remained unchanged. The changes in biomass switched the system from net emission to net uptake of CO2 (0.47 and -0.77 µmol m-2 s-1 in grazed and exclosure plots, respectively) during the growing season and doubled the C storage in live biomass. In contrast, the treatment had no impact on the CH4 fluxes, the total litter C pool or the soil C concentration. The rapid recovery of the above ground biomass and CO2 fluxes demonstrates the plasticity of this high arctic ecosystem in terms of response to changing herbivore pressure.