Pores provide important hotspots for chemical and biological processes in soils. Earthworm burrows affect the macropore structure and their actions may create new preferential pathways for water and gas flow within soils. This, in turn, indirectly affect plants, nutrient cycling, hydraulic conductivity, gas exchange, and soil organisms. While the effects of invasive earthworms on soil properties has been well-documented in temperate and boreal ecosystems, we know little how these organism may affect tundra soils. In this study, I assessed how the three-dimensional network of soil-macropores are affected by earthworm species (Aporrectodea sp. and Lumbricus sp). I hypothesized: i) that earthworms increase the frequency of macropores with a likely biological origin (biopores); ii) effects of biopores are dependent on tundra vegetation type (meadow or heath); and iii) the macropore network properties are altered by earthworms.  The hypotheses were tested using a common garden experiment with 48 mesocosms. The pore structure of each mesocosm was analyzed using X-ray CT tomography. I found that biopores increased in the tundra from on 0.05 ±0.01 % (mean ± standard deviation) in the control to about 0.59 ± 0.07 % in the earthworm treatments. However, in contrast to my second hypothesis, I found no vegetation dependent effect. Interestingly, I found that earthworms decreased the complexity and directionality of macropores. My findings strongly indicate that burrowing can severely impact the pore properties of previously uninhabited subarctic soils.

Nitrogen (N), acquired by roots and mycorrhizal fungi and supplied to plant foliage, is a growth-limiting nutrient at the subarctic treeline. Due to this limitation, interspecific competition and acquisition of N is an important control on plant community composition and distribution. The ability of trees and shrubs to access N shapes community dynamics at this ecotone undergoing species range shifts and changes in primary productivity driven by climate change. Using 15N soil labelling we investigate the fate of soil inorganic N, and spatial distances over which trees and understorey shrubs access soil N, in a treeline forest. 15N was injected into soil rooting zones in discrete 1 m2 patches and foliar samples were collected from trees between 1 and 50 m away, and understorey shrubs between 0.5 and 11 m away from labelled soil. The 15N label was found in mountain birch trees up to 5 m, and in understorey shrubs up to 2 m, away from labelled soil. We estimate that 1.27% of pulse-derived N was found in foliage of birch trees, compared to 1.16% in the understorey. However, mountain birch trees contributed only 31% of ecosystem leaf area index (LAI), thus there was a disproportionate allocation of added label to the birch canopy compared with its contribution to ecosystem LAI. The difference in root and mycorrhizal exploration distances and community N partitioning between mountain birch trees and understorey shrubs may confer competitive advantage to trees with respect to nitrogen and nutrient patches, which may alter plant community structures within these forests. This is particularly important considering predicted climate-driven tree and tall shrub expansion in subarctic regions, with likely consequences for ecosystem N and carbon (C) cycling, as well as for community composition and biodiversity.

Tundra ecosystems hold large stocks of soil organic matter (SOM), likely due to low temperatures limiting rates of microbial SOM decomposition more than those of SOM accumulation from plant primary productivity and microbial necromass inputs. Here we test the hypotheses that distinct tundra vegetation types and their carbon supply to characteristic rhizosphere microbes determine SOM cycling independent of temperature. In the subarctic Scandes, we used a three-way factorial design with paired heath and meadow vegetation at each of two elevations, and with each combination of vegetation type and elevation subjected during one growing season to either ambient light (i.e., ambient plant productivity), or 95% shading (i.e., reduced plant productivity). We assessed potential above- and belowground ecosystem linkages by uni- and multivariate analyses of variance, and structural equation modelling. We observed direct coupling between tundra vegetation type and microbial community composition and function, which underpinned the ecosystem's potential for SOM storage. Greater primary productivity at low elevation and ambient light supported higher microbial biomass and nitrogen immobilisation, with lower microbial mass-specific enzymatic activity and SOM humification. Congruently, larger SOM at lower elevation and in heath sustained fungal-dominated microbial communities, which were less substrate-limited, and invested less into enzymatic SOM mineralisation, owing to a greater carbon-use efficiency (CUE). Our results highlight the importance of tundra plant community characteristics (i.e., productivity and vegetation type), via their effects on soil microbial community size, structure and physiology, as essential drivers of SOM turnover. The here documented concerted patterns in above- and belowground ecosystem functioning is strongly supportive of using plant community characteristics as surrogates for assessing tundra carbon storage potential and its evolution under climate and vegetation changes.

The dynamics of phosphorus (P) in the environment is important for regulating nutrient cycles in natural and managed ecosystems and an integral part in assessing biological resilience against environmental change. Organic P (P-o) compounds play key roles in biological and ecosystems function in the terrestrial environment being critical to cell function, growth and reproduction. We asked a group of experts to consider the global issues associated with P-o in the terrestrial environment, methodological strengths and weaknesses, benefits to be gained from understanding the P-o cycle, and to set priorities for P-o research. We identified seven key opportunities for P-o research including: the need for integrated, quality controlled and functionally based methodologies; assessment of stoichiometry with other elements in organic matter; understanding the dynamics of P-o in natural and managed systems; the role of microorganisms in controlling P-o cycles; the implications of nanoparticles in the environment and the need for better modelling and communication of the research. Each priority is discussed and a statement of intent for the P-o research community is made that highlights there are key contributions to be made toward understanding biogeochemical cycles, dynamics and function of natural ecosystems and the management of agricultural systems.

Motives for fishing differ among fishers, which may imply different effects of climate change on the net values of fishing. Climate change has impacts on fish population dynamics and on other factors in the fishers’ harvest decision, such as alternative sources of food or income. Here we present a bio-economic model that includes impacts of climate change on fish population and on net values of harvest by fishers with recreational or subsistence fishing motives. The conceptual analysis shows that the economic effects of climate change with simultaneous impacts on fish population growth and harvest values are inconclusive with common fishing access for both fisher types and when there are opposite simultaneous climate effects with exclusive access for one of the fisher types. Numerical results from our model of Arctic char (Salvelinus alpinus) in northern Sweden indicate that climate change, measured as temperature increases, reduces fish population growth but increases net values of fishing for both fisher types. The combined net effect of these counteracting forces is that annual net values can almost cease for the subsistence fisher in the future but increase considerably for the recreational fisher.

Soil viruses are highly abundant and have important roles in the regulation of host dynamics and soil ecology. Climate change is resulting in unprecedented changes to soil ecosystems and the life forms that reside there, including viruses. In this Review, we explore our current understanding of soil viral diversity and ecology, and we discuss how climate change (such as extended and extreme drought events or more flooding and altered precipitation patterns) is influencing soil viruses. Finally, we provide our perspective on future research needs to better understand how climate change will impact soil viral ecology.

In the nineteenth century, numerous settlements were established in the alpine region of Fennoscandia (the Scandes), an area that later became a major international scene for Arctic research. Here we raise awareness of this era and show that earthworm-driven bioturbation in “pristine” soils around contemporary Arctic research infrastructure is caused by soil fauna left behind during early land use. We use soil preserved under an alpine settlement to highlight that soils were not bioturbated when the first house was built at a site where bioturbation is now widespread. A review of archived material with unique site-specific chronology constrained the onset of bioturbation to the post-1871 era. Our results suggest that small-scale land use introduced earthworms that now thrive far beyond the realms of former cultivated fields. The legacy of soil fauna from this example of “ecological imperialism” still lingers and should be considered when studying soils of the Scandes.

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.

Over the last decade, an increasing number of studies have used soundscapes to address diverse ecological questions. Sound represents one of the few sources of information capable of providing in situ insights into processes occurring within opaque soil matrices. To date, the use of soundscapes for soil macrofauna monitoring has been experimentally tested only in controlled laboratory environments. Here we assess the validity of laboratory predictions and explore the use of soil soundscape proxies for monitoring soil macrofauna (i.e., earthworm) activities in an outdoor context. In a common garden experiment in northern Sweden, we constructed outdoor mesocosm plots (N = 36) containing two different Arctic vegetation types (meadow and heath) and introduced earthworms to half of these plots. Earthworms substantially altered the ambient soil soundscape under both vegetation types, as measured by both traditional soundscape indices and frequency band power levels, although their acoustic impacts were expressed differently in heath versus meadow soils. While these findings support the as-of-yet untapped promise of using belowground soundscape analyses to monitor soil ecosystem health, direct acoustic emissions from earthworm activities appear to be an unlikely proxy for tracking worm activities at daily timescales. Instead, earthworms indirectly altered the soil soundscape by ‘re-engineering’ the soil matrix: an effect that was dependent on vegetation type. Our findings suggest that long-term (i.e., seasonal) earthworm activities in natural soil settings can likely be monitored indirectly via their impacts on soundscape measures and acoustic indices. Analyzing soil soundscapes may enable larger-scale monitoring of high-latitude soils and is directly applicable to the specific case of earthworm invasions within Arctic soils, which has recently been identified as a potential threat to the resilience of high-latitude ecosystems. Soil soundscapes could also offer a novel means to monitor soils and soil-plant-faunal interactions in situ across diverse pedogenic, agronomic, and ecological systems.

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.

Nitrogen fixation associated with cryptogams is potentially very important in arctic and subarctic terrestrial ecosystems, as it is a source of new nitrogen (N) into these highly N limited systems. Moss-, lichen- and legume-associated N-2 fixation was studied with high frequency (every second week) during spring, summer, autumn and early winter to uncover the seasonal variation in input of atmospheric N-2 to a subarctic heath with an altered climate. We estimated N-2 fixation from ethylene production by acetylene reduction assay in situ in a field experiment with the treatments: long- vs. short-term summer warming using plastic tents and litter addition (simulating expansion of the birch forest). N-2 fixation activity was measured from late April to mid November and 33 % of all N-2 was fixed outside the vascular plant growing season (Jun-Aug). This substantial amount underlines the importance of N-2 fixation in the cold period. Warming increased N-2 fixation two- to fivefold during late spring. However, long-term summer warming tended to decrease N-2 fixation outside the treatment (tents present) period. Litter alone did not alter N-2 fixation but in combination with warming N-2 fixation increased, probably because N-2 fixation became phosphorus limited under higher temperatures, which was alleviated by the P supply from the litter. In subarctic heath, the current N-2 fixation period extends far beyond the vascular plant growing season. Climate warming and indirect effects such as vegetation changes affect the process of N-2 fixation in different directions and thereby complicate predictions of future N cycling.

Climate change is predicted to cause milder winters and thus exacerbate soil freeze?thaw perturbations in the subarctic, recasting the environmental challenges that soil microorganisms need to endure. Historical exposure to environmental stressors can facilitate the microbial resilience to new cycles of that same stress. However, whether and how such microbial memory or stress legacy can modulate microbial responses to cycles of frost remains untested. Here, we conducted an in situ field experiment in a subarctic birch forest, where winter warming resulted in a substantial increase in the number and intensity of freeze?thaw events. After one season of winter warming, which raised mean surface and soil (?8?cm) temperatures by 2.9 and 1.4°C, respectively, we investigated whether the in situ warming-induced increase in frost cycles improved soil microbial resilience to an experimental freeze?thaw perturbation. We found that the resilience of microbial growth was enhanced in the winter warmed soil, which was associated with community differences across treatments. We also found that winter warming enhanced the resilience of bacteria more than fungi. In contrast, the respiration response to freeze?thaw was not affected by a legacy of winter warming. This translated into an enhanced microbial carbon-use efficiency in the winter warming treatments, which could promote the stabilization of soil carbon during such perturbations. Together, these findings highlight the importance of climate history in shaping current and future dynamics of soil microbial functioning to perturbations associated with climate change, with important implications for understanding the potential consequences on microbial-mediated biogeochemical cycles.

Climate change currently manifests in upward and northward shifting treelines, which encompasses changes to the carbon (C) and nitrogen (N) composition of organic inputs to soils. Whether these changed inputs will increase or decrease microbial mineralisation of native soil organic matter remains unknown, making it difficult to estimate how treeline shifts will affect the C balance. Aiming to improve mechanistic understanding of C cycling in regions experiencing treeline shifts, we quantified priming effects in soils of high altitudes (Peruvian Andes) and high latitudes (subarctic Sweden), differentiating landcover types (boreal forest, tropical forest, tundra heath, Puna grassland) and soil horizons (organic, mineral). In a controlled laboratory incubation, soils were amended with substrates of different C:N, composed of an organic C source at a constant ratio of 30% substrate-C to microbial biomass C, combined with different levels of a nutrient solution neutral in pH. Substrate additions elicited both positive and negative priming effects in both ecosystems, independent from substrate C:N. Positive priming prevailed above the treeline in high altitudes and in mineral soils in high latitudes, where consequently climate change-induced treeline shifts and deeper rooting plants may enhance SOM-mineralisation and soil C emissions. However, such C loss may be compensated by negative priming, which dominated in the other soil types and was of larger magnitude than positive priming. In line with other studies, these results indicate a consistent mechanism linking decreased SOM-mineralisation (negative priming) to increased microbial substrate utilisation, suggesting preferential substrate use as a potential tool to support soil C storage.

The microbial ecology of arctic and sub-arctic soils is an important aspect of the global carbon cycle, due to the sensitivity of the large soil carbon stocks to ongoing climate warming. These regions are characterized by strong climatic seasonality, but the emphasis of most studies on the short vegetation growing season could potentially limit our ability to predict year-round ecosystem functions. We compiled a database of studies from arctic, subarctic, and boreal environments that include sampling of microbial community and functions outside the growing season. We found that for studies comparing across seasons, in most environments, microbial biomass and community composition vary intra-annually, with the spring thaw period often identified by researchers as the most dynamic time of year. This seasonality of microbial communities will have consequences for predictions of ecosystem function under climate change if it results in: seasonality in process kinetics of microbe-mediated functions; intra-annual variation in the importance of different (a)biotic drivers; and/or potential temporal asynchrony between climate change-related perturbations and their corresponding effects. Future research should focus on (i) sampling throughout the entire year; (ii) linking these multi-season measures of microbial community composition with corresponding functional or physiological measurements to elucidate the temporal dynamics of the links between them; and (iii) identifying dominant biotic and abiotic drivers of intra-annual variation in different ecological contexts.

1. Plant–soil feedback (PSF) results from the influence of plants on the composition and abundance of various taxa and functional groups of soil micro-organisms, and their reciprocal effects on the plants. However, little is understood about the importance of fine root traits and root economic strategies in moderating microbial-driven PSF.
2. We examined the relationships between PSF and 11 chemical and morphological root traits from 18 sub-arctic meadow plant species, as well as the soil microbial community composition which we characterized using phospholipid fatty acids (PLFAs) and high-throughput sequencing. We also investigated the importance of the root economics spectrum in influencing PSF, because it indicates plant below-ground economic strategies via trade-offs between resource acquisition and conservation.
3. When we considered the entire root economics spectrum, we found that PSFs were more negative when root trait values were more acquisitive across the 18 species. In addition, PSF was more negative when values of root nitrogen content and root forks per root length were higher, and more positive when root dry matter content was higher. We additionally identified two fungal orders that were negatively related to PSF. However, we found no evidence that root traits influenced PSF through its relationship with these fungal orders.
4. Synthesis. Our results provide evidence that for some fine root traits, the root economics spectrum and some fungal orders have an important role in influencing PSF. By investigating the roles of soil micro-organisms and fine root traits in driving PSF, this study enables us to better understand root trait–microbial linkages across species and therefore offers new insights about the mechanisms that underpin PSFs and ultimately plant community assembly.

Plant community assembly processes shape the composition and abundances of species, and encompass functional traits and resource acquisition strategy of species, biotic interactions and abiotic filtering. Hence, an understanding of these complex processes requires disentangling the effects of multiple factors influencing plant community assembly. In this thesis, I investigated fine root trait associations with soil microorganisms, the resulting feedback effects from those interactions (i.e., plant-soil feedbacks), plant-plant interactions under warming, and the effects of temperature on fine root traits of plant communities in the Swedish sub-arctic tundra.

Here, the chemical root economics spectrum (i.e., tradeoff between acquisitive and conservative strategies) predicted the abundance of broad microbial groups, whereas individual fine root traits were associated with the relative abundances of fungal taxa. It also explained plant-soil feedback, with acquisitive trait values resulting in negative feedbacks. In addition, plant-plant interactions were altered under warming, but this was not related to resource-acquisition strategy. Further, community-level root trait responses to temperature were not necessarily related to root resource investment strategy. 

Taken together, this thesis shows the importance of fine root traits for understanding plant community responses to global change. This has implications for plant community assembly, as well as carbon and nutrient cycling in a future warmer sub-arctic tundra.

This study explores soil nutrient cycling processes and microbial properties for two contrasting vegetation types along an elevational gradient in subarctic tundra to improve our understanding of how temperature influences nutrient availability in an ecosystem predicted to be sensitive to global warming. We measured total amino acid (Amino-N), mineral nitrogen (N) and phosphorus (P) concentrations, in situ net N and P mineralization, net Amino-N consumption, and microbial biomass C, N and P in both heath and meadow soils across an elevational gradient near Abisko, Sweden. For the meadow, NH4 (+) concentrations and net N mineralization were highest at high elevations and microbial properties showed variable responses; these variables were largely unresponsive to elevation for the heath. Amino-N concentrations sometimes showed a tendency to increase with elevation and net Amino-N consumption was often unresponsive to elevation. Overall, PO4-P concentrations decreased with elevation and net P immobilization mostly occurred at lower elevations; these effects were strongest for the heath. Our results reveal that elevation-associated changes in temperature can have contrasting effects on the cycling of N and P in subarctic soils, and that the strength and direction of these effects depend strongly on dominant vegetation type.

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.

Considerable progress has been made in ecological and evolutionary genetics with studies demonstrating how genes underlying plant and microbial traits can influence adaptation and even 'extend' to influence community structure and ecosystem level processes. Progress in this area is limited to model systems with deep genetic and genomic resources that often have negligible ecological impact or interest. Thus, important linkages between genetic adaptations and their consequences at organismal and ecological scales are often lacking. Here we introduce the Sphagnome Project, which incorporates genomics into a long-running history of Sphagnum research that has documented unparalleled contributions to peatland ecology, carbon sequestration, biogeochemistry, microbiome research, niche construction, and ecosystem engineering. The Sphagnome Project encompasses a genus-level sequencing effort that represents a new type of model system driven not only by genetic tractability, but by ecologically relevant questions and hypotheses.