Decomposition rates are commonly assessed from mass loss in litterbags or teabags, but underestimation may occur if organic matter (OM) enters the bags during decomposition, e.g. through colonization by fungal hyphae. We used C4-plant (lemongrass) teabags and an isotopic mass-balance approach to quantify external OM inputs to decomposing litter during a 24-month field study in two C3-plant ecosystems, a temperate forest and a heath tundra, with contrasting climates and vegetation. After 24 months, OM inputs represented 4% of recovered mass in the tundra and 22% in the forest, corresponding to an underestimation of decomposition by 3% to 6%, with stronger biases in the forest due to accelerated litter processing and belowground activity. The use of litter- or teabags containing plant material with isotopic signatures distinct from the studied system enables the quantification of OM inputs entering the bags during litter decomposition, thereby improving the comparability of decomposition rates across ecosystems.

While litter-dwelling earthworms are widespread in northern Scandinavian forests, burrowing earthworms, i.e., species commonly considered as ecosystem engineers, are largely missing in these ecosystems. We hypothesized that their absence results not only from environmental filtering by unfavorable soil and climatic conditions but also from dispersal limitation. We examined earthworm presence-absence across Swedish forests by environmental DNA (eDNA) sequencing of 312 soil inventory samples. We then used statistical models to identify factors best explaining their occurrence among a set of variables representing soil properties, climatic factors, vegetation and distances to anthropogenic activities (e.g. roads and agricultural fields). Soil pH and texture were strong explanatory variables for the presence of burrowing earthworms, while distance to agricultural fields explained a significant part of the remaining variation. The importance of proximity to agricultural fields was especially pronounced in northern Sweden (> 60° latitude). In contrast, occurrence of litter-dwelling earthworms was only weakly linked to environmental variables. Additionally, based on random forest modelling we predicted earthworm presence for almost 7000 soil inventory sites, and created high-resolution probability maps of earthworm occurrence in Swedish forests, including a scenario without potential dispersal limitation. These maps show that occurrence of burrowing earthworms would be more likely in large parts of the northern inland forests of Sweden without dispersal limitations, suggesting that these forests may be more susceptible to earthworm establishment upon introduction than previously anticipated. Our results highlight the importance of anthropogenic activities in dispersion and distribution of burrowing earthworms in northern forests. While environmental filtering largely governs burrowing earthworm occurrence, their current distribution is also substantially influenced by dispersal limitation - an underestimated factor to date.

Alleviation of functional limitations by soil fauna is key to climate feedbacks from arctic soils 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 fauna presence or absence is key to understanding and predicting future climate feedbacks from arctic soils. We propose that the arrival of certain soil fauna into arctic soils may introduce “novel functions”, resulting in increased rates of, for example, nitrogen cycling, litter fragmentation, or bioturbation, and thereby alleviate functional limitations of the current soil organism 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 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.

Past glaciations eradicated burrowing earthworms from northern Scandinavia, and their slow natural dispersal rates have left higher latitude landscapes still mostly earthworm-free. Burrowing earthworms that are common in southern Scandinavia may expand their presence in northern ecosystems, as is suggested by occasional observations of burrowing earthworms in northern Sweden. Given the dramatic impacts earthworm colonisation can have on forests, it is important to assess the likeliness of burrowing earthworm expansion in Scandinavian forested ecosystems. To do this, we need to increase our understanding of earthworm distribution in relation to ecosystem and soil properties, human activities and climate.We have tracked earthworm distribution across a large set of archived soil samples collected by the Swedish Forest Soil Inventory. Earthworm presence at species level was identified by DNA extraction directly from soil, thus detection mostly relied on DNA traces e.g. in casts and cocoons, rather than on the sampling of earthworm individuals. Using corresponding site-specific data for vegetation, soil characteristics and climate, we identified the best predictors for earthworm presence and modelled the probability and geographic distribution of (future) earthworm occurrence in Swedish forests.

Earthworms are considered ecosystem engineers due to their remarkable influence on the soil system. While creating the drilosphere in the soil, they interact with microorganisms both directly and indirectly and thereby greatly affecting soil carbon and nutrient cycling. Earthworm activity may reshape soil microbial communities in several ways. Amongst others, earthworms may cause shifts in microbial communities and activities/processes by damaging hyphal networks, selectively feeding on substrates hosting certain bacteria and fungi, and by redistributing nutrients in the litter-soil continuum of the soil. However, interactions between earthworms and soil microbes, especially fungi, are poorly understood, and the mechanisms by which earthworms affect microorganisms are challenging to study. First, given the widespread presence of earthworms, finding soils that have not been previously affected by earthworms is difficult. Second, controlled laboratory incubation experiments generally exclude certain functionally important groups of fungi such as plant-associated ectomycorrhizal (EcM) fungi. In our recently initiated project, we aim to study earthworm influence on soil fungal communities and associated soil biogeochemical processes by introducing soil-dwelling earthworms into to date yet uncolonised northern forest soils. Therefore, we established mesocosm boxes filled with soil turfs including tree saplings from northern boreal forests and placed them in an experimental forest in southern Sweden. Later on, we will introduce earthworms (Aporrectodea and Lumbricus spp) into the mesocosms and measure (depth specific) changes in microbial communities and genes using RNA and DNA sequencing. Potential microbial changes will be related to measurements of carbon and nitrogen cycling, such as carbon-dioxide flux measurements and soil mineral nitrogen content analysis in the growing season after earthworm introduction.

Vegetation changes in a warming Arctic may affect plant-associated soil microbial communities with possible consequences for the biogeochemical cycling of carbon (C) and nitrogen (N). In a sub-arctic tundra heath, we factorially removed plant species with ecto- and ericoid mycorrhizal associations. After two years, we explored how mycorrhizal type-specific plant removal influences microbial communities, soil and microbial C and N pools, and extracellular enzymatic activities. Removal of ecto- and ericoid mycorrhizal plants did not change the soil fungal or bacterial community composition or their extracellular enzyme activities. However, ericoid plant removal decreased microbial C:N ratio, suggesting a stoichiometric effect decoupled from microbial community composition. In other words, microbial communities appear to show initial plasticity in response to major changes in tundra vegetation. This highlights the importance of longer-term perspectives when investigating the effects of vegetation changes on biogeochemical processes in Arctic ecosystems.

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.

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.

Trees interact with a multitude of microbes through their roots and root symbionts such as mycorrhizal fungi and root endophytes. Here, we explore the role of fungal root symbionts as predictors of the soil and root-associated microbiomes of widespread broad-leaved trees across a European latitudinal gradient. Our results suggest that, alongside factors such as climate, soil, and vegetation properties, root colonization by ectomycorrhizal, arbuscular mycorrhizal, and dark septate endophytic fungi also shapes tree-associated microbiomes. Notably, the structure of root and soil microbiomes across our sites is more strongly and consistently associated with dark septate endophyte colonization than with mycorrhizal colonization and many abiotic factors. Root colonization by dark septate endophytes also has a consistent negative association with the relative abundance and diversity of nutrient cycling genes. Our study not only indicates that root-symbiotic interactions are an important factor structuring soil communities and functions in forest ecosystems, but also that the hitherto less studied dark septate endophytes are likely to be central players in these interactions.

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.

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.

IntroductionSurvival 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.MethodsWe 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.ResultsWarming 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.DiscussionOur 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.

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 CO 2 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 CO 2 release in the frost boils and by counterbalancing the negative effects of cryoturbation on plant CO 2 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.

Biological decomposition and wildfire are two predominant and alternative processes that can mineralize organic C in forest litter. Currently, the relationships between decomposition and fire are still poorly understood. We provide an empirical test of the hypothesized decoupling of surface litter bed decomposability and flammability, and the underlying traits and trait spectra. We employed a 41‐species set of gymnosperms of very broad evolutionary and geographic spread, because of the wide range of (absent to frequent) fire regimes they are associated with. We found that the interspecific pattern of mass loss proportions in a “common garden” decomposition experiment was not correlated with any of the flammability parameters and an RDA analysis also showed that the decomposability and flammability of leaf litter in litter layers were decoupled across species. This decoupling originates from the former depending mostly on size and shape spectrum traits and the latter on PES traits and those trait spectra being virtually uncorrelated. Synthesis : Our results show that, indeed, leaf litter decomposability and flammability parameters are decoupled across species, and this decoupling can be explained by their different drivers in terms of trait spectra: chemical traits for decomposability and size‐shape traits for litter layer flammability.

Targeted removal experiments are a powerful tool to assess the effects of plant species or (functional) groups on ecosystem functions. However, removing plant biomass in itself can bias the observed responses. This bias is commonly addressed by waiting until ecosystem recovery, but this is inherently based on unverified proxies or anecdotal evidence. Statistical control methods are efficient, but restricted in scope by underlying assumptions.We propose accounting for such biases within the experimental design, using a gradient of biomass removal controls. We demonstrate the relevance of this design by presenting (1) conceptual examples of suspected biases and (2) how to observe and control for these biases.Using data from a mycorrhizal association-based removal experiment, we show that ignoring biomass removal biases (including by assuming ecosystem recovery) can lead to incorrect, or even contrary conclusions (e.g. false positive and false negative). Our gradient design can prevent such incorrect interpretations, regardless of whether aboveground biomass has fully recovered.Our approach provides more objective and quantitative insights, independently assessed for each variable, than using a proxy to assume ecosystem recovery. Our approach circumvents the strict statistical assumptions of, for example, ANCOVA and thus offers greater flexibility in data analysis.

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.

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.

This archive contains the processed sequencing data, processed CO2 flux data and the script used to generate figures and tables presented in "Dispersal of bacteria and stimulation of permafrost decomposition by Collembola" by Monteux S., Mariën J., and Krab E.J., Biogeosciences 2022, 19, 4089-4105, https://doi.org/10.5194/bg-19-4089-2022 . A more exhaustive repository including bioinformatics processing is found at https://git.bolin.su.se/bolin/monteux-2022-collembola and the raw DNA data is deposited at ENA under accession number PRJEB51992

The supply of recent photosynthate from plants to soils is thought to be a critical mechanism regulating the activity and diversity of soil biota. In the Arctic, large-scale vegetation transitions are underway in response to warming, and there is an urgent need to understand how these changes affect soil biodiversity and function. We investigated how abundance and diversity of soil fungi and invertebrates responded to a reduction in fresh below-ground photosynthate supply in treeline birch and willow, achieved using stem girdling. We hypothesised that birch forest would support greater abundance of ectomycorrhizal (ECM) fungal species and fauna than willow shrubs, and that girdling would result in a rapid switch from ECM fungi to saprotrophs as canopy supply of C was cut, with a concomitant decline in soil fauna. Birch forest had greater fungal and faunal abundance with a large contribution of root-associated ascomycetes (ericoid mycorrhizal fungi and root endophytes) compared to willow shrub plots, which had a higher proportion of saprotrophs and, contrary to our expectations, ECM fungi. Broad-scale soil fungal and faunal functional group composition was not significantly changed by girdling, even in the third year of treatment. Within the ECM community, there were some changes, with genera that are believed to be particularly C-demanding declining in girdled plots. However, it was notable how most ECM fungi remained present after 3 years of isolation of the below-ground compartment from contemporary photosynthate supply. Synthesis. In a treeline/tundra ecosystem, distinct soil communities existed in contrasting vegetation patches within the landscape, but the structure of these communities was resistant to canopy disturbance and concomitant reduction of autotrophic C inputs.

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.