Umeå University's logo

Arctic soils play an important role in Earth's climate system, as they store large amounts of carbon that, if released, could strongly increase greenhouse gas levels in our atmosphere. Most research to date has focused on how the turnover of organic matter in these soils is regulated by abiotic factors, and few studies have considered the potential role of biotic regulation. However, arctic soils are currently missing important groups of soil organisms, and here, we highlight recent empirical evidence that soil organisms' presence or absence is key to understanding and predicting future climate feedbacks from arctic soils. We propose that the arrival of soil organisms into arctic soils may introduce "novel functions", resulting in increased rates of, for example, nitrification, methanogenesis, litter fragmentation, or bioturbation, and thereby alleviate functional limitations of the current community. This alleviation can greatly enhance decomposition rates, in parity with effects predicted due to increasing temperatures. We base this argument on a series of emerging experimental evidence suggesting that the dispersal of until-then absent micro-, meso-, and macroorganisms (i.e. from bacteria to earthworms) into new regions and newly thawed soil layers can drastically affect soil functioning. These new observations make us question the current view that neglects organism-driven "alleviation effects"when predicting future feedbacks between arctic ecosystems and our planet's climate. We therefore advocate for an updated framework in which soil biota and the functions by which they influence ecosystem processes become essential when predicting the fate of soil functions in warming arctic ecosystems.

Arctic plant growth is predominantly nitrogen (N) limited. This limitation is generally attributed to slow soil microbial processes due to low temperatures. Here, we show that arctic plant-soil N cycling is also substantially constrained by the lack of larger detritivores (earthworms) able to mineralize and physically translocate litter and soil organic matter. These new functions provided by earthworms increased shrub and grass N concentration in our common garden experiment. Earthworm activity also increased either the height or number of floral shoots, while enhancing fine root production and vegetation greenness in heath and meadow communities to a level that exceeded the inherent differences between these two common arctic plant communities. Moreover, these worming effects on plant N and greening exceeded reported effects of warming, herbivory and nutrient addition, suggesting that human spreading of earthworms may lead to substantial changes in the structure and function of arctic ecosystems. Arctic plant growth is predominantly nitrogen limited, where the slow nitrogen turnover in the soil is commonly attributed to the cold arctic climate. Here the authors show that the arctic plant-soil nitrogen cycling is also constrained by the lack of larger detritivores like earthworms.

The study of ecosystem processes over multiple scales of space and time is often best achieved using comparable data from multiple sites. Yet, long-term ecological observatories have often developed their own data collection protocols. Here, we address this problem by proposing a set of ecological protocols suitable for widespread adoption by the ecological community. Scientists from the European ecological research community prioritized terrestrial ecosystem parameters that could benefit from a more consistent approach to data collection within the resources available at most long-term ecological observatories. Parameters for which standard methods are in widespread use, or for which methods are evolving rapidly, were not selected. Protocols were developed by domain experts, building on existing methods where possible, and refined through a process of field testing and training. They address above-ground plant biomass; decomposition; land use and management; leaf area index; soil mesofaunal diversity; soil C and N stocks, and greenhouse gas emissions from soils. These complement existing methods to provide a complete assessment of ecological integrity. These protocols offer integrated approaches to ecological data collection that are low cost and are starting to be used across the European Long Term Ecological Research community.

Introduction: Survival and growth of tree seedlings are key processes of regeneration in forest ecosystems. However, little is known about how climate warming modulates seedling performance either directly or in interaction with understory vegetation and post-fire successional stages.

Methods: We measured survival (over 3 years) and growth of seedlings of three tree species (Betula pubescens, Pinus sylvestris, and Picea abies) in a full-factorial field experiment with passive warming and removal of two plant functional groups (feather moss and/or ericaceous shrubs) along a post-fire chronosequence in an unmanaged boreal forest.

Results: Warming had no effect on seedling survival over time or on relative biomass growth. Meanwhile, moss removal greatly increased seedling survival overall, while shrub removal canceled this effect for B. pubescens seedlings. In addition, B. pubescens and P. sylvestris survival benefitted most from moss removal in old forests (>260 years since last fire disturbance). In contrast to survival, seedling growth was promoted by shrub removal for two out of three species, i.e., P. sylvestris and P. abies, meaning that seedling survival and growth are governed by different understory functional groups affecting seedling performance through different mechanism and modes of action.

Discussion: Our findings highlight that understory vegetation and to a lesser extent post-fire successional stage are important drivers of seedling performance while the direct effect of climate warming is not. This suggests that tree regeneration in future forests may be more responsive to changes in understory vegetation or fire regime, e.g., indirectly caused by warming, than to direct or interactive effects of rising temperatures.

As global temperatures continue to rise, a key uncertainty of climate projections is the microbial decomposition of vast organic carbon stocks in thawing permafrost soils. Decomposition rates can accelerate up to fourfold in the presence of plant roots, and this mechanism—termed the rhizosphere priming effect—may be especially relevant to thawing permafrost soils as rising temperatures also stimulate plant productivity in the Arctic. However, priming is currently not explicitly included in any model projections of future carbon losses from the permafrost area. Here, we combine high-resolution spatial and depth-resolved datasets of key plant and permafrost properties with empirical relationships of priming effects from living plants on microbial respiration. We show that rhizosphere priming amplifies overall soil respiration in permafrost-affected ecosystems by ~12%, which translates to a priming-induced absolute loss of ~40 Pg soil carbon from the northern permafrost area by 2100. Our findings highlight the need to include fine-scale ecological interactions in order to accurately predict large-scale greenhouse gas emissions, and suggest even tighter restrictions on the estimated 200 Pg anthropogenic carbon emission budget to keep global warming below 1.5 °C.

Human introductions have resulted in earthworms establishing in the Arctic, species known to cause cascading ecosystem change. However, few quantitative outdoor experiments have been performed that describe how these soil modifying earthworms are reshaping structures in tundra soils. In this study, we used three-dimensional (3-D) X-ray images of soil cores (approximately 10 cm diameter, 20 cm height, N = 48) to assess how earthworms (Aporrectodea sp. and Lumbricus sp.) affect soil structure and macropore networks in an outdoor mesocosm experiment that lasted four summers. Effects were assessed in both shrub-dominated (heath) and herb-dominated (meadow) tundra. Earthworms almost doubled the macroporosity in meadow soils and tripled macroporosity in heath. Interestingly, the fractal dimension of macropores decreased in response to earthworm burrowing in both systems, indicating that the presence of earthworms reduced the geometric complexity in comparison to other pore-generating processes active in the tundra. Observed effects on soil structure occurred along with a dramatically reduced soil moisture content, which was observed the first winter after earthworm introduction in the meadow. Our findings suggest that predictions of future changes in vegetation and soil carbon pools in the Arctic should include major impacts on soil properties that earthworms induce.

Extreme climate events are observed and predicted to increase in frequency and duration in high-latitudeecosystems as a result of global climate change. This includes extreme precipitation events, which maydirectly impact on belowground food webs and ecosystem functioning by their physical impacts and byaltering local soil moisture conditions.

We assessed responses of the Collembola community in a northern Sphagnum fuscum-dominatedombrotrophic peatland to three years of experimentally increased occurrence of extreme precipitationevents. Annual summer precipitation was doubled (an increase of 200 mm) by 16 simulated extremerain events within the three months growing season, where on each occasion 12.5 mm of rain was addedwithin a few minutes. Despite this high frequency and intensity of the rain events, no shifts in Collemboladensity, relative species abundances and community weighted means of three relevant traits (moisturepreference, vertical distribution and body size) were observed. This strongly suggests that the peatlandCollembola community is unaffected by the physical impacts of extreme precipitation and the short-termvariability in moisture conditions. The lack of response is most likely reinforced by the fact that extremeprecipitation events do not seem to alter longer-term soil moisture conditions in the peat layers inhabitedby soil fauna.

This study adds evidence to the observation that the biotic components of northern ombrotrophicpeatlands are hardly responsive to an increase in extreme summer precipitation events. Given the importance of these ecosystems for the global C balance, these findings significantly contribute to the currentknowledge of the ecological impact of future climate scenarios. (C) 2014 Elsevier B.V. All rights reserved.

Climate manipulation experiments in the field and laboratory incubations are common methods to study the impact of climate change on soils and their biota. However, both types of methods have drawbacks either on their mechanistic interpretation or ecological relevance. We propose an experimental set-up that combines the best of both methods and can be easily obtained by modifying widely available Tullgren soil fauna extractors. This set-up creates or alters temperature and moisture gradients within intact field soil cores, after which soil biota, their activity and vertical movements can be studied. We assessed the performance and demonstrated the applicability of this set-up through a case study on Collembola response to changes in microclimatic gradients in peat bogs. Warming created a vertical temperature gradient of 14 degrees C in peat cores without varying soil moisture conditions, while at a given temperature regime, precipitation and drought treatments shifted natural soil moisture gradients to 'wetter' and 'drier', respectively. This allowed for disentangling interacting warming and moisture effects on soil fauna. In our case study, Collembola communities showed peat layer-specific responses to these climate treatments. Warming decreased Collembola density and altered community composition in the shallowest layer, whereas precipitation increase affected Collembola community composition in the deepest layer. We showed that climate change can have layer-specific effects on soil organisms that are 'hidden' by not taking microclimatic vertical gradients into account. This experimental set-up facilitates studying (multitrophic) organism responses to climate changes, with only a small adjustment of equipment that is often already present in soil ecology laboratories. Moreover, this set-up can be easily customized to study many more other research questions related to wide-ranging organisms and ecosystems.

Climate change induced alterations to winter conditions may affect decomposer organisms controlling the vast carbon stores in northern soils. Soil microarthropods are particularly abundant decomposers in Arctic ecosystems. We studied whether increased snow depth affected microarthropods, and if effects were consistent over two consecutive winters. We sampled Collembola and soil mites from a snow accumulation experiment at Svalbard in early summer and used soil microclimatic data to explore to which aspects of winter climate microarthropods are most sensitive. Community densities differed substantially between years and increased snow depth had inconsistent effects. Deeper snow hardly affected microarthropods in 2015, but decreased densities and altered relative abundances of microarthropods and Collembola species after a milder winter in 2016. Although increased snow depth increased soil temperatures by 3.2 °C throughout the snow cover periods, the best microclimatic predictors of microarthropod density changes were spring soil temperature and snowmelt day. Our study shows that extrapolation of observations of decomposer responses to altered winter climate conditions to future scenarios should be avoided when communities are only sampled on a single occasion, since effects of longer-term gradual changes in winter climate may be obscured by inter-annual weather variability and natural variability in population sizes.

At high latitudes, winter warming facilitates vegetation expansion into barren frost-affected soils. The interplay of changes in winter climate and plant presence may alter soil functioning via effects on decomposers. Responses of decomposer soil fauna and microorganisms to such changes likely differ from each other, since their life histories, dispersal mechanisms and microhabitats vary greatly.

We investigated the relative impacts of short-term winter warming and increases in plant cover on bacteria and collembola community composition in cryoturbated, non-sorted circle tundra. By covering non-sorted circles with insulating gardening fibre cloth (fleeces) or using stone walls accumulating snow, we imposed two climate-change scenarios: snow accumulation increased autumn-to-late winter soil temperatures (−1 cm) by 1.4 °C, while fleeces warmed soils during that period by 1 °C and increased spring temperatures by 1.1 °C. Summer bacteria and collembola communities were sampled from within-circle locations differing in vegetation abundance and soil properties.

Two years of winter warming had no effects on either decomposer community. Instead, their community compositions were strongly determined by sampling location: communities in barren circle centres were distinct from those in vegetated outer rims, while communities in sparsely vegetated patches of circle centres were intermediate. Diversity patterns indicate that collembola communities are tightly linked to plant presence while bacteria communities correlated with soil properties.

Our results thus suggest that direct effects of short-term winter warming are likely to be minimal, but that vegetation encroachment on barren cryoturbated ground will affect decomposer community composition substantially. At decadal timescales, collembola community changes may follow relatively fast after warming-driven plant establishment into barren areas, whereas bacteria communities may take longer to respond. If shifts in decomposer community composition are indicative for changes in their activity, vegetation overgrowth will likely have much stronger effects on soil functioning in frost-affected tundra than short-term winter warming.

Climate change-driven increases in winter temperatures positively affect conditions for shrub growth in arctic tundra by decreasing plant frost damage and stimulation of nutrient availability. However, the extent to which shrubs may benefit from these conditions may be strongly dependent on the following spring climate. Species-specific differences in phenology and spring frost sensitivity likely affect shrub growth responses to warming. Additionally, effects of changes in winter and spring climate may differ over small spatial scales, as shrub growth may be dependent on natural variation in snow cover, shrub density and cryoturbation. We investigated the effects of winter warming and altered spring climate on growing-season performance of three common and widespread shrub species in cryoturbated non-sorted circle arctic tundra. By insulating sparsely vegetated non-sorted circles and parts of the surrounding heath with additional snow or gardening fleeces, we created two climate change scenarios: snow addition increased soil temperatures in autumn and winter and delayed snowmelt timing without increasing spring temperatures, whereas fleeces increased soil temperature similarly in autumn and winter, but created warmer spring conditions without altering snowmelt timing. Winter warming affected shrub performance, but the direction and magnitude were species-specific and dependent on spring conditions. Spring warming advanced, and later snowmelt delayed canopy green-up. The fleece treatment did not affect shoot growth and biomass in any shrub species despite decreasing leaf frost damage in Empetrum nigrum. Snow addition decreased frost damage and stimulated growth of Vaccinium vitis-idaea by c. 50%, while decreasing Betula nana growth (p < .1). All of these effects were consistent the mostly barren circles and surrounding heath. Synthesis. In cryoturbated arctic tundra, growth of Vaccinium vitis-idaea may substantially increase when a thicker snow cover delays snowmelt, whereas in longer term, warmer winters and springs may favour E. nigrum instead. This may affect shrub community composition and cover, with potentially far-reaching effects on arctic ecosystem functioning via its effects on cryoturbation, carbon cycling and trophic cascading. Our results highlight the importance of disentangling effects of winter and spring climate change timing and nature, as spring conditions are a crucial factor in determining the impact of winter warming on plant performance.

Soil mixing by earthworms can have a large impact on the fate of nutrients and pollutants and on the soil's ability to sequester carbon. Nevertheless, methods to quantify earthworm ingestion and egestion under field conditions are largely lacking. Soils of the Fennoscandian tundra offer a special possibility for such quantifications, as these soils commonly lack burrowing macrofauna and exhibit a well-defined O horizon with low bulk density on top of a mineral soil with higher density. Since ingestion-egestion mixes the two soil layers, the temporal changes in the bulk density profile of such soils may be useful for estimating field ingestion rates. In this study, we applied a model for earthworm burrowing through soil ingestion to observed changes in soil densities occurring in a mesocosm experiment carried out in the arctic during four summers with intact soil. The earthworms present in the mesocosms were Aporrectodea trapezoides, Aporrectodea tuberculata, Aporrectodea rosea, Lumbricus rubellus and Lumbricus Terrestris (fourth season only). We show that changes in soil density profiles can indeed be used to infer earthworm ingestion rates that are realistic in comparison to literature values. Although uncertainties in parameter values were sometimes large, the results from this study suggest that soil turnover rates and endogeic earthworm soil ingestion rates in tundra heath and meadow soils may be as high as those reported for temperate conditions. Such large ingestion rates can explain observed large morphological changes in arctic soils where dispersing earthworms have resulted in complete inmixing of the organic layer into the mineral soil. Our approach is applicable to soil profiles with marked vertical differences in bulk density such as the soils of the Fennoscandian tundra where earthworms are currently dispersing into new areas and to layered repacked soil samples that are incubated in the field.

Climate warming enables tree seedling establishment beyond the current alpine treeline, but to achieve this, seedlings have to establish within existing tundra vegetation. In tundra, mosses are a prominent feature, known to regulate soil temperature and moisture through their physical structure and associated water retention capacity. Moss presence and species identity might therefore modify the impact of increases in temperature and precipitation on tree seedling establishment at the arctic‐alpine treeline. We followed Betula pubescens and Pinus sylvestris seedling survival and growth during three growing seasons in the field. Tree seedlings were transplanted along a natural precipitation gradient at the subarctic‐alpine treeline in northern Sweden, into plots dominated by each of three common moss species and exposed to combinations of moss removal and experimental warming by open‐top chambers (OTCs). Independent of climate, the presence of feather moss, but not Sphagnum , strongly supressed survival of both tree species. Positive effects of warming and precipitation on survival and growth of B. pubescens seedlings occurred in the absence of mosses and as expected, this was partly dependent on moss species. P. sylvestris survival was greatest at high precipitation, and this effect was more pronounced in Sphagnum than in feather moss plots irrespective of whether the mosses had been removed or not. Moss presence did not reduce the effects of OTCs on soil temperature. Mosses therefore modified seedling response to climate through other mechanisms, such as altered competition or nutrient availability. We conclude that both moss presence and species identity pose a strong control on seedling establishment at the alpine treeline, and that in some cases mosses weaken climate‐change effects on seedling establishment. Changes in moss abundance and species composition therefore have the potential to hamper treeline expansion induced by climate warming.

Warming-induced microbial decomposition of organic matter in permafrost soils constitutes a climate-change feedback of uncertain magnitude. While physicochemical constraints on soil functioning are relatively well understood, the constraints attributable to microbial community composition remain unclear. Here we show that biogeochemical processes in permafrost can be impaired by missing functions in the microbial community-functional limitations-probably due to environmental filtering of the microbial community over millennia-long freezing. We inoculated Yedoma permafrost with a functionally diverse exogenous microbial community to test this mechanism by introducing potentially missing microbial functions. This initiated nitrification activity and increased CO2 production by 38% over 161 days. The changes in soil functioning were strongly associated with an altered microbial community composition, rather than with changes in soil chemistry or microbial biomass. The present permafrost microbial community composition thus constrains carbon and nitrogen biogeochemical processes, but microbial colonization, likely to occur upon permafrost thaw in situ, can alleviate such functional limitations. Accounting for functional limitations and their alleviation could strongly increase our estimate of the vulnerability of permafrost soil organic matter to decomposition and the resulting global climate feedback. Carbon dioxide emissions from permafrost thaw are substantially enhanced by relieving microbial functional limitations, according to incubation experiments on Yedoma permafrost.

Contrary to most soils, permafrost soils have the atypical feature of being almost entirely deprived of soil fauna. Abiotic constraints on the fate of permafrost carbon after thawing are increasingly understood, but biotic constraints remain scarcely investigated. Incubation studies, essential to estimate effects of permafrost thaw on carbon cycling, typically measure the consequences of permafrost thaw in isolation from the topsoil and thus do not account for the effects of altered biotic interactions because of e.g. colonization by soil fauna. Microarthropods facilitate the dispersal of microorganisms in soil, both on their cuticle (ectozoochory) and through their digestive tract (endozoochory), which may be particularly important in permafrost soils, considering that microbial community composition can strongly constrain permafrost biogeochemical processes.

Here we tested how a model species of microarthropod (the Collembola Folsomia candida) affected aerobic CO2 production of permafrost soil over a 25 d incubation. By using Collembola stock cultures grown on permafrost soil or on an arctic topsoil, we aimed to assess the potential for endo- and ectozoochory of soil bacteria, while cultures grown on gypsum and sprayed with soil suspensions would allow the observation of only ectozoochory.

The presence of Collembola introduced bacterial amplicon sequence variants (ASVs) absent in the no-Collembola control, regardless of their microbiome manipulation, when considering presence-absence metrics (unweighted UniFrac metrics), which resulted in increased species richness. However, these introduced ASVs did not induce changes in bacterial community composition as a whole (accounting for relative abundances, weighted UniFrac), which might only become detectable in the longer term.

CO2 production was increased by 25.85 % in the presence of Collembola, about half of which could be attributed to Collembola respiration based on respiration rates measured in the absence of soil. We argue that the rest of the CO2 being respired can be considered a priming effect of the presence of Collembola, i.e. a stimulation of permafrost CO2 production in the presence of active microarthropod decomposers. Overall, our findings underline the importance of biotic interactions in permafrost biogeochemical processes and the need to explore the additive or interactive effects of other soil food web groups of which permafrost soils are deprived.

Springtails (Collembola) inhabit soils from the Arctic to the Antarctic and comprise an estimated ~32% of all terrestrial arthropods on Earth. Here, we present a global, spatially-explicit database on springtail communities that includes 249,912 occurrences from 44,999 samples and 2,990 sites. These data are mainly raw sample-level records at the species level collected predominantly from private archives of the authors that were quality-controlled and taxonomically-standardised. Despite covering all continents, most of the sample-level data come from the European continent (82.5% of all samples) and represent four habitats: woodlands (57.4%), grasslands (14.0%), agrosystems (13.7%) and scrublands (9.0%). We included sampling by soil layers, and across seasons and years, representing temporal and spatial within-site variation in springtail communities. We also provided data use and sharing guidelines and R code to facilitate the use of the database by other researchers. This data paper describes a static version of the database at the publication date, but the database will be further expanded to include underrepresented regions and linked with trait data.

Soil life supports the functioning and biodiversity of terrestrial ecosystems. Springtails (Collembola) are among the most abundant soil arthropods regulating soil fertility and flow of energy through above- and belowground food webs. However, the global distribution of springtail diversity and density, and how these relate to energy fluxes remains unknown. Here, using a global dataset representing 2470 sites, we estimate the total soil springtail biomass at 27.5 megatons carbon, which is threefold higher than wild terrestrial vertebrates, and record peak densities up to 2 million individuals per square meter in the tundra. Despite a 20-fold biomass difference between the tundra and the tropics, springtail energy use (community metabolism) remains similar across the latitudinal gradient, owing to the changes in temperature with latitude. Neither springtail density nor community metabolism is predicted by local species richness, which is high in the tropics, but comparably high in some temperate forests and even tundra. Changes in springtail activity may emerge from latitudinal gradients in temperature, predation and resource limitation in soil communities. Contrasting relationships of biomass, diversity and activity of springtail communities with temperature suggest that climate warming will alter fundamental soil biodiversity metrics in different directions, potentially restructuring terrestrial food webs and affecting soil functioning.

The emergence of high-throughput DNA sequencing methods provides unprecedented opportunities to further unravel bacterial biodiversity and its worldwide role from human health to ecosystem functioning. However, despite the abundance of sequencing studies, combining data from multiple individual studies to address macroecological questions of bacterial diversity remains methodically challenging and plagued with biases. Here, using a machine-learning approach that accounts for differences among studies and complex interactions among taxa, we merge 30 independent bacterial data sets comprising 1,998 soil samples from 21 countries. Whereas previous meta-analysis efforts have focused on bacterial diversity measures or abundances of major taxa, we show that disparate amplicon sequence data can be combined at the taxonomy-based level to assess bacterial community structure. We find that rarer taxa are more important for structuring soil communities than abundant taxa, and that these rarer taxa are better predictors of community structure than environmental factors, which are often confounded across studies. We conclude that combining data from independent studies can be used to explore bacterial community dynamics, identify potential 'indicator' taxa with an important role in structuring communities, and propose hypotheses on the factors that shape bacterial biogeography that have been overlooked in the past.

Arctic tundra active-layer soils are at risk of soil organic carbon (SOC) depletion and degradation upon global climate warming because they are in a stage of relatively early decomposition. Non-growing season (NGS) warming is particularly pronounced, and observed increases of CO2 emissions during experimentally warmed NGSs give concern for great SOC losses to the atmosphere. Here, we used snow fences in Arctic Spitsbergen dwarf shrub tundra to simulate 1.86 degrees C NGS warming for 9 consecutive years, while growing season temperatures remained unchanged. In the snow fence treatment, the 4-11 cm thick A-horizon had a 2% lower SOC concentration and a 0.48 kg Cm-2 smaller pool size than the controls, indicating SOC pool depletion. The snow fence treatment's A-horizon's alkyl/O-alkyl ratio was also significantly increased, indicating an advance of SOC degradation. The underlying 5 cm of B/C-horizon did not show these effects. Our results support the hypothesis that SOC depletion and degradation are connected to the long-term transience of observed ecosystem respiration (ER) increases upon soil warming. We suggest that the bulk of warming induced ER increases may originate from surface and not deep active layer or permafrost horizons. The observed losses of SOC might be significant for the ecosystem in question, but are in magnitude comparatively small relative to anthropogenic greenhouse gas enrichment of the atmosphere. We conclude that a positive feedback of carbon losses from surface soils of Arctic dwarf shrub tundra to anthropogenic forcing will be minor, but not negligible.

Warming-induced increases in microbial CO2 release in northern tundra may positively feedback to climate change. However, shifts in microbial extracellular enzyme activities (EEAs) may alter the impacts of warming over the longer term. We investigated the in situ effects of 3years of winter warming in combination with the in vitro effects of a rapid warming (6days) on microbial CO2 release and EEAs in a subarctic tundra heath after snowmelt in spring. Winter warming did not change microbial CO2 release at ambient (10 degrees C) or at rapidly increased temperatures, i.e., a warm spell (18 degrees C) but induced changes (P<0.1) in the Q(10) of microbial respiration and an oxidative EEA. Thus, although warmer winters may induce legacy effects in microbial temperature acclimation, we found no evidence for changes in potential carbon mineralization after spring thaw.

Cryogenic land surface processes (CLSPs), such as cryoturbation, are currently active in landscapes covering 25% of our planet where they dictate key functions, such as carbon (C) cycling, and maintain patterned landscape features. While CLSPs are expected to diminish in the near future due to milder winters especially in the southern parts of the Arctic, the shifts in C cycling in these landscapes may be more complex, since climate change can affect C cycling directly but also indirectly via CLSPs. Here, we study the effects of changing winter and spring climate on CLSPs and C cycling in non-sorted circles consisting of barren frost boils and their vegetated rims. We do this by measuring cryoturbation and ecosystem CO2 fluxes repeatedly in alpine subarctic tundra where temperatures during naturally snow covered period have been experimentally increased with snow-trapping fences and temperatures during winter and spring period after snowmelt have been increased with insulating fleeces. Opposite to our hypothesis, warming treatments did not decrease cryoturbation. However, winter warming via deeper snow increased ecosystem C sink during summer by decreasing ecosystem CO2 release in the frost boils and by counterbalancing the negative effects of cryoturbation on plant CO2 uptake in the vegetated rims. Our results suggest that short-term changes in winter and spring climate may not alter cryoturbation and jeopardize the tundra C sink.

Frost-patterned grounds, such as mostly barren frost boils surrounded by denser vegetation, are typical habitat mosaics in tundra. Plant and microbial processes in these habitats may be susceptible to short-term warming outside the growing season, while the areal cover of barren frost boils has decreased during the past decades due to climate warming-induced shrub expansion. The relative importance of such short-term and long-term climate impacts on carbon (C) dynamics remains unknown. We measured ecosystem CO2 uptake and release (in the field), microbial respiration (in the laboratory), as well as microbial biomass N and soil extractable N in frost boils and the directly adjacent heath in late spring and late summer. These habitats had been experimentally warmed with insulating fleeces from late September until late May for three consecutive years, which allowed us to investigate the direct short-term effects of warming and longer-term, indirect climate effects via vegetation establishment into frost boils. Non-growing season warming increased C uptake at the frost boils in late spring and decreased it in late summer, while the timing and direction of responses was opposite for the heath. Experimental warming had no effects on microbial or ecosystem C release or soil N at either of the habitats. However, C cycling was manifold higher at the heath compared to the frost boils, likely because of a higher SOM stock in the soil. Short-term climate change can thus directly alter ecosystem C uptake at frost-patterned grounds but will most likely not affect microbial C release. We conclude that the C dynamics at frost-patterned grounds under a changing climate depend most strongly on the potential of vegetation to encroach into frost boils in the long-term.

Environmental changes feedback to climate through their impact on soil functions such as carbon (C) and nutrient sequestration. Abiotic conditions and the interactions between above- and belowground biota drive soil responses to environmental change but these (a)biotic interactions are challenging to study. Nonetheless, better understanding of these interactions would improve predictions of future soil functioning and the soil-climate feedback and, in this context, permafrost soils are of particular interest due to their vast soil C-stores. We need new tools to isolate abiotic (microclimate, chemistry) and biotic (roots, fauna, microorganisms) components and to identify their respective roles in soil processes. We developed a new experimental setup, in which we mimic thermokarst (permafrost thaw-induced soil subsidence) by fitting thawed permafrost and vegetated active layer sods side by side into mesocosms deployed in a subarctic tundra over two growing seasons. In each mesocosm, the two sods were separated from each other by barriers with different mesh sizes to allow varying degrees of physical connection and, consequently, (a)biotic exchange between active layer and permafrost. We demonstrate that our mesh-approach succeeded in controlling 1) lateral exchange of solutes between the two soil types, 2) colonization of permafrost by microbes but not by soil fauna, and 3) ingrowth of roots into permafrost. In particular, experimental thermokarst induced a similar to 60% decline in permafrost nitrogen (N) content, a shift in soil bacteria and a rapid buildup of root biomass (+33.2 g roots m(-2) soil). This indicates that cascading plant-soil-microbe linkages are at the heart of biogeochemical cycling in thermokarst events. We propose that this novel setup can be used to explore the effects of (a)biotic ecosystem components on focal biogeochemical processes in permafrost soils and beyond.