1 Decomposition of plant litter, a key component of the global carbon budget, is hierarchically controlled by the triad: climate > litter quality > soil organisms. Given the sensitivity of decomposition to temperature, especially in cold biomes, it has been hypothesized that global warming will lead to increased litter decomposition rates, both through direct temperature effects and through indirect effects on litter quality and soil organisms. 2 A meta-analysis of experimental warming studies in cold biomes (34 site-species combinations) showed that warming resulted in slightly increased decomposition rates. However, this response was strongly dependent on the method used: open top chambers reduced decomposition rates, whereas heating lamps stimulated decomposition rates. The low responsiveness was mainly due to moisture-limited decomposition rates in the warming treatments, especially at mesic and xeric sites. This control of litter decomposition by both temperature and moisture was corroborated by natural gradient studies. 3 Interspecific differences in litter quality and decomposability are substantially larger than warming-induced phenotypic responses. Thus, the changes in the species composition and structure of plant communities that have been observed in medium-term warming studies in cold biomes will have a considerably greater impact on ecosystem litter decomposition than phenotypic responses. 4 Soil fauna communities in cold biomes are responsive to climate warming. Moreover, temperature-driven migration of the, hitherto absent, large comminuters to high-latitude sites may significantly increase decomposition rates. However, we do not know how far-reaching the consequences of changes in the species composition and structure of the soil community are for litter decomposition, as there is a lack of data on functional species redundancy and the species’ dispersal ability. 5 Global warming will lead to increased litter decomposition rates only if there is sufficient soil moisture. Hence, climate scenario and experimental studies should focus more on both factors and their interaction. As interspecific differences in potential decomposability and litter chemistry are substantially larger than phenotypic responses to warming, the focus of future research should be on the former. In addition, more light should be shed on the below-ground ‘darkness’ to evaluate the ecological significance of warming-induced soil fauna community changes for litter decomposition processes in cold biomes.

The relative importance of global versus local environmental factors for growth and thus carbon uptake of the bryophyte genusSphagnum-the main peat-former and ecosystem engineer in northern peatlands-remains unclear. We measured length growth and net primary production (NPP) of two abundantSphagnumspecies across 99 Holarctic peatlands. We tested the importance of previously proposed abiotic and biotic drivers for peatland carbon uptake (climate, N deposition, water table depth and vascular plant cover) on these two responses. Employing structural equation models (SEMs), we explored both indirect and direct effects of drivers onSphagnumgrowth. Variation in growth was large, but similar within and between peatlands. Length growth showed a stronger response to predictors than NPP. Moreover, the smaller and denserSphagnum fuscumgrowing on hummocks had weaker responses to climatic variation than the larger and looserSphagnum magellanicumgrowing in the wetter conditions. Growth decreased with increasing vascular plant cover within a site. Between sites, precipitation and temperature increased growth forS. magellanicum. The SEMs indicate that indirect effects are important. For example, vascular plant cover increased with a deeper water table, increased nitrogen deposition, precipitation and temperature. These factors also influencedSphagnumgrowth indirectly by affecting moss shoot density. Synthesis. Our results imply that in a warmer climate,S. magellanicumwill increase length growth as long as precipitation is not reduced, whileS. fuscumis more resistant to decreased precipitation, but also less able to take advantage of increased precipitation and temperature. Such species-specific sensitivity to climate may affect competitive outcomes in a changing environment, and potentially the future carbon sink function of peatlands.

1.  Climate change in northern high latitudes is predicted to be greater in winter rather than summer, yet little is known about the effects of winter climate change on northern ecosystems. Among the unknowns are the effects of an increasing frequency of acute, short‐lasting winter warming events. Such events can damage higher plants exposed to warm, then returning cold, temperatures after snow melt, and it is not known how bryophytes and lichens, which are of considerable ecological importance in high‐latitude ecosystems, are affected by such warming events. However, even physiological adaptations of these cryptogams to winter environments in general are poorly understood.

2.  Here we describe findings from a novel field experiment that uses heating from infrared lamps and soil warming cables to simulate acute mid‐winter warming events in a sub‐Arctic heath. In particular, we report the growing season responses of the dominant lichen, Peltigera aphthosa, and bryophyte, Hylocomium splendens, to warming events in three consecutive winters.

3.  While summertime photosynthetic performance of P. aphthosa was unaffected by the winter warming treatments, H. splendens showed significant reductions in net photosynthetic rates and growth rates (of up to 48% and 52%, respectively). Negative effects were evident already during the summer following the first winter warming event.

4.  While the lichen develops without going through critical phenological stages during which vulnerable organs are produced, the moss has a seasonal rhythm, which includes initiation of growth of young, freeze‐susceptible shoot apices in the early growing season; these might be damaged by breaking of dormancy during warm winter events.

5. Synthesis. Different sensitivities of the bryophyte and lichen species were unexpected, and illustrate that very little is known about the winter ecology of bryophytes and lichens from cold biomes in general. In sharp contrast to summer warming experiments that show increased vascular plant biomass and reduced lichen biomass, these results demonstrate that acute climate events in mid‐winter may be readily tolerated by lichens, in contrast to previously observed sensitivity of co‐occurring dwarf shrubs, suggesting winter climate change may compensate for (or even reverse) predicted lichen declines resulting from summer warming.

* Despite time-lags and nonlinearity in ecological processes, the majority of our knowledge about ecosystem responses to long-term changes in climate originates from relatively short-term experiments. * We utilized the longest ongoing snow removal experiment in the world and an additional set of new plots at the same location in northern Sweden to simultaneously measure the effects of long-term (11 winters) and short-term (1 winter) absence of snow cover on boreal forest understorey plants, including the effects on root growth and phenology. * Short-term absence of snow reduced vascular plant cover in the understorey by 42%, reduced fine root biomass by 16%, reduced shoot growth by up to 53% and induced tissue damage on two common dwarf shrubs. In the long-term manipulation, more substantial effects on understorey plant cover (92% reduced) and standing fine root biomass (39% reduced) were observed, whereas other response parameters, such as tissue damage, were observed less. Fine root growth was generally reduced, and its initiation delayed by c. 3 (short-term) to 6 weeks (long-term manipulation). * Synthesis. We show that one extreme winter with a reduced snow cover can already induce ecologically significant alterations. We also show that long-term changes were smaller than suggested by an extrapolation of short-term manipulation results (using a constant proportional decline). In addition, some of those negative responses, such as frost damage and shoot growth, were even absolutely stronger in the short-term compared to the long-term manipulation. This suggests adaptation or survival of only those individuals that are able to cope with these extreme winter conditions, and that the short-term manipulation alone would overpredict long-term impacts. These results highlight both the ecological importance of snow cover in this boreal forest, and the value of combining short- and long-term experiments side by side in climate change research.

1. Transitions between major vegetation types, such as the tree line, are useful systems for monitoring the response of vegetation to climate change. Tree lines have, however, shown equivocal responses to such change. 2. Tree lines are considered to be primarily thermally controlled, although recent work has highlighted the importance of biotic factors. Dispersal limitation and the invasibility of the tundra matrix have been implicated and here we propose herbivory as an additional control at some tree lines. 3. We propose a conceptual model in which differing relative impacts of foliage consumption, availability of establishment sites, trampling, dispersal and seed predation can lead to very different tree-line responses. 4. The presence of large numbers of small trees above the current tree line at a site in northern Sweden that experiences limited reindeer (Rangifer tarandus) herbivory suggests range expansion. Other locations in the same region with higher reindeer populations have considerably fewer small trees, suggesting that range expansion is occurring much more slowly, if at all. 5. The use of tree lines as indicators of climate change is confounded by the activity of herbivores, which may either strengthen or nullify the impacts of a changed climate. Similar arguments are likely to be applicable to other ecotones.

Plant growth forms are widely used to predict the effects of environmental changes, such as climate warming and increased nitrogen deposition, on plant communities, and the consequences of species shifts for carbon and nutrient cycling. We investigated whether the relationship between growth forms and patterns in litter quality and decomposition are independent of environmental conditions and whether growth forms are as good as litter chemistry at predicting decomposability. We used a natural, latitudinal gradient in NW Europe as a spatial analogue for future increases in temperature and nitrogen availability. Our screening of 70 species typical of Sphagnum-dominated peatlands showed that leaf litters of Sphagnum mosses, evergreen and deciduous shrubs, graminoids and forbs differed significantly in litter chemistry and that the ranking of the growth forms was independent of the region for all litter chemistry variables. Differences among growth forms were usually larger than differences related to the environmental gradient. After 8 and 20 months incubation in outdoor, Sphagnum-based decomposition beds, growth forms generally differed in decomposability, but these patterns varied with latitude. Sphagnum litters decomposed slower than other litters in all regions, again explaining its high representation in organic deposits of peatlands. Forb litters generally decomposed fastest, while the differences among the other growth forms were small, particularly at higher latitudes. Multiple regression analyses showed that growth forms were better at predicting leaf litter decomposition than chemical variables in warm-temperate peatlands with a high N-load, but less so in the subarctic, low-N region. Our results indicate that environmental changes may be less important in determining ecosystem leaf litter chemistry directly than are their indirect effects through changes in the relative abundance of growth forms. However, climatic and nutritional constraints in high-latitude peatlands promote convergence towards nutrient-efficient plant traits, resulting in similar decomposition rates of vascular growth forms despite differences in litter chemistry. The usefulness of the growth-form concept in predicting plant community controls on ecosystem functioning is therefore somewhat limited.

1. Plant species affect ecosystem carbon uptake via biomass production and carbon release via decomposition processes. Differences in their responses to climate change and effects on ecosystem carbon cycling processes may thus feedback to the atmospheric carbon balance and the climate at a global scale. Hierarchical species classifications based on plant growth forms are widely used in cold, northern biomes to generalize and predict these differences. This review investigates the usefulness of broad (vascular, non-vascular), intermediate (woody, non-woody) and narrow (evergreen shrubs, deciduous shrubs, graminoids, forbs) plant growth-form-based groups in these biomes for predicting plant responses to climate change and effects on the main processes of the full carbon cycle by looking at the similarity of species within growth-form groups and the consistency of differences among groups under changing environmental conditions. 2. Production responses to climate change differ between broad growth-form groups, but their opposite responses do not imply that the responses of non-vascular plants are consistently negative. Within vascular growth forms, production responses to climate change are not always similar among species under identical conditions, and average differences among narrow vascular growth forms are usually small. Moreover, differences in production responses among growth forms strongly depend upon the duration of the study, the region and the ecosystem type. 3. Species within narrow growth forms show a high similarity for a range of leaf litter chemistry variables and differences among narrow growth forms are often large and consistent. However, differences in leaf litter decomposability are large between broad growth-form groups, but small and environment-dependent among important narrow vascular growth forms. Litter feedback effects to plant production vary among narrow vascular growth forms, but the differences strongly depend on the study duration. Data on the climate dependence of growth-form differences regarding this aspect of the carbon cycle are currently lacking. 4. Synthesis: Overall, the usefulness of growth-form-based groups clearly differs between carbon cycling processes. Different aggregation levels are therefore needed for different processes. For most processes there is evidence that the differences among growth forms depend on environmental conditions, which hampers their use for generalizations and modelling. Future studies should therefore explicitly test for differences among growth-form groups and aim to unravel the dependence of growth-form differences on environmental conditions.

1. The stress gradient hypothesis suggests a shift from predominant competition to facilitation along gradients of increasing environmental severity. This shift is proposed to cause parallel changes from prevailing spatial segregation to aggregation among the species within a community. 2. We used 904 1-m(2) plots, each subdivided into 100 10 x 10 cm, or 25 20 x 20 cm cells, respectively, from 67 European mountain summits grouped into 18 regional altitudinal transects, to test this hypothesized correlation between fine-scale spatial patterns and environmental severity. 3. The data were analysed by first calculating standardized differences between observed and simulated random co-occurrence patterns for each plot. These standardized effect sizes were correlated to indicators of environmental severity by means of linear mixed models. In a factorial design, separate analyses were made for four different indicators of environmental severity (the mean temperature of the coldest month, the temperature sum of the growing season, the altitude above tree line, and the percentage cover of vascular plants in the whole plot), four different species groups (all species, graminoids, herbs, and all growth forms considered as pseudospecies) and at the 10 x 10 cm and 20 x 20 cm grain sizes. 4. The hypothesized trends were generally weak and could only be detected by using the mean temperature of the coldest month or the percentage cover of vascular plants as the indicator of environmental severity. The spatial arrangement of the full species set proved more responsive to changes in severity than that of herbs or graminoids. The expected trends were more pronounced at a grain size of 10 x 10 cm than at 20 x 20 cm. 5. Synthesis. In European alpine plant communities the relationships between small-scale co-occurrence patterns of vascular plants and environmental severity are weak and variable. This variation indicates that shifts in net interactions with environmental severity may differ among indicators of severity, growth forms and scales. Recognition of such variation may help to resolve some of the current debate surrounding the stress gradient hypothesis.

1. Evidence is growing that leaf litter generally decomposes faster than expected in its environment of origin, owing to specialization of litter and topsoil decomposer communities to break down litter encountered most often. Nevertheless, this home‐field advantage (HFA) in decomposition is inconsistently supported by experimental data and fails to account for situations where contrasting qualities of litter coexist within the same litter matrix.

2. In contrast to the HFA hypothesis, which expects a positive interaction between every litter species produced locally and the local decomposer communities irrespective of litter species quality, we define here an alternative substrate quality–matrix quality interaction (SMI) hypothesis that expects a continuum from positive to negative interaction between specific litters (substrates) and decomposer communities as specific litters and the ecosystem litter layer (i.e. the matrix, which drives local decomposer community composition) become increasingly dissimilar in quality.

3. To test this hypothesis, we conducted a reciprocal transplant decomposition experiment of eight leaf, six fine‐stem and nine fine‐root litter species from three neighbouring ecosystems of the subarctic biome: dry forest, riparian forest and forest‐surrounded pond; and characterized the quality (represented by lignin content and an integrated measure of carbon/nutrient economics) of each litter species and each ecosystem litter layer.

4. We found substantial overall effects of SMI on decomposition rates of leaf (20% explained variance), stem (14%) and root (15%) litters, although this effect was lower than the single effects of litter quality and microclimate (remaining explained variance). Despite being partly inconsistent across litter species, likely due to the complexity of litter quality–decomposer community relationships, the SMI hypothesis appeared more broadly applicable than the HFA hypothesis.

5.             Synthesis. We demonstrate here that plant traits, likely via their control on litter and topsoil decomposer community composition, have indirect effects on litter breakdown rates, not only at the interface between ecosystems but also within ecosystems, with likely implications for many other ecosystems world‐wide. These results suggest functional variation in decomposer communities between ecosystems with respect to their efficiency to degrade litters with contrasting qualities, such as different lignolytic and detoxification activities but also contrasting efficiencies to degrade non‐recalcitrant tissues.

1. While the importance of wood decay for the global carbon balance is widely recognized, surprisingly little is known about its long‐term dynamics and its abiotic and biotic drivers. Progress in this field is hindered by the long time‐scales inherent to the low decay rates of wood and the lack of short‐term methods to assess long‐term decomposition dynamics in standardized field conditions.

2. Here, we present such a method, which relies on the sampling and short‐term incubation of wood from several decay stages covering the entire decay process. Together these short‐term decay steps are used to model and discriminate between three potential decay dynamics (linear, exponential and sigmoid) using an iterative optimization procedure. We applied this method to analyse long‐term wood decay of six subarctic tree species (six stems and two roots) and test the hypotheses that (i) different wood species follow distinct decay dynamics and (ii) interspecific variation in wood traits controls variation in wood decay rates in a standardized environment.

3. We found interspecific variation in long‐term wood decay dynamics: decay of Alnus and Salix stems was best described by exponential models, whereas decay of Sorbus stems and Betula and Pinus roots was best fitted by linear models and Betula, Pinus and Populus stems each displayed a sigmoid decay dynamics (up to 5‐year initial lag phase). A six‐fold variation was observed between the decomposition half‐lives of all eight wood types, from 6.8 years (6.1–7.5, 95% C.I.) for Alnus stems to 41.3 years (34.5–51.8) for Pinus roots. Initial wood traits such as pH (R² = 0.92), dry matter content (R² = 0.79) and lignin (R² = 0.73) were good predictors of long‐term wood decay rates.

4. Synthesis. Our findings suggest changing decay dynamics across wood species and types that are likely to arise from changing underlying wood decay processes (i.e. varying wood functional traits/decomposer community interactions). Our new method, which combines advantages of direct observations and the chronosequence approach, allows reliable comparisons of species contributions to long‐term wood decay rates and provides future opportunities to experimentally disentangle intrinsic and external abiotic and biotic drivers of long‐term wood decay processes.

* Conceptual frameworks relating plant traits to ecosystem processes such as organic matter dynamics are progressively moving from a leaf-centred to a whole-plant perspective. Through the use of meta-analysis and global literature data, we quantified the relative roles of litters from above- and below-ground plant organs in ecosystem labile organic matter dynamics. * We found that decomposition rates of leaves, fine roots and fine stems were coordinated across species worldwide although less strongly within ecosystems. We also show that fine roots and stems had lower decomposition rates relative to leaves, with large differences between woody and herbaceous species. Further, we estimated that on average below-ground litter represents approximately 33 and 48% of annual litter inputs in grasslands and forests, respectively. * These results suggest a major role for below-ground litter as a driver of ecosystem organic matter dynamics. We also suggest that, given that fine stem and fine root litters decompose approximately 1.5 and 2.8 times slower, respectively, than leaf litter derived from the same species, cycling of labile organic matter is likely to be much slower than predicted by data from leaf litter decomposition only. * Synthesis. Our results provide evidence that within ecosystems, the relative inputs of above- versus below-ground litter strongly control the overall quality of the litter entering the decomposition system. This in turn determines soil labile organic matter dynamics and associated nutrient release in the ecosystem, which potentially feeds back to the mineral nutrition of plants and therefore plant trait values and plant community composition.

1. Phototrophic microbes, also known as micro-algae, display a high abundance in many terrestrial surface soils. They contribute to atmospheric carbon dioxide fluxes through their photosynthesis, and thus regulate climate similar to plants. However, microbial photosynthesis remains overlooked in most terrestrial ecosystems. Here, we hypothesise that phototrophic microbes significantly contribute to peatland C uptake, unless environmental conditions limit their development and their photosynthetic activity.
2. To test our hypothesis, we studied phototrophic microbial communities in five peatlands distributed along a latitudinal gradient in Europe. By means of metabarcoding, microscopy and cytometry analyses, as well as measures of photosynthesis, we investigated the diversity, absolute abundance and photosynthetic rates of the phototrophic microbial communities.
3. We identified 351 photosynthetic prokaryotic and eukaryotic operational taxonomic units (OTUs) across the five peatlands. We found that water availability and plant composition were important determinants of the composition and the structure of phototrophic microbial communities. Despite environmental shifts in community structure and composition, we showed that microbial C fixation rates remained similar along the latitudinal gradient. Our results further revealed that phototrophic microbes accounted for approximately 10% of peatland C uptake.
4. Synthesis. Our findings show that phototrophic microbes are extremely diverse and abundant in peatlands. While species turnover with environmental conditions, microbial photosynthesis similarly contributed to peatland C uptake at all latitudes. We estimate that phototrophic microbes take up around 75 MT CO2 per year in northern peatlands. This amount roughly equals the magnitude of projected peatland C loss due to climate warming and highlights the importance of phototrophic microbes for the peatland C cycle.

In high-latitude ecosystems bryophytes are important drivers of ecosystem functions. Alterations in abundance of mosses due to global change may thus strongly influence carbon (C) and nitrogen (N) cycling and hence cause feedback on climate. The effects of mosses on soil microbial activity are, however, still poorly understood. Our study aims at elucidating how and by which mechanisms bryophytes influence microbial decomposition processes of soil organic matter and thus soil nutrient availability. We present results from a field experiment in a subarctic birch forest in northern Sweden, where we partly removed the moss cover and replaced it with an artificial soil cover for simulating moss effects on soil temperature and moisture. We combined this with a fertilization experiment with 15N-labelled N for analysing the effects of moss N sequestration on soil processes. Our results demonstrate the capacity of mosses to reduce soil N availability and retard N cycling. The comparison with artificial soil cover plots suggests that the effect of mosses on N cycling is linked to the thermal insulation capacity of mosses causing low average soil temperature in summer and strongly reduced soil temperature fluctuations, the latter also leading to a decreased frequency of freeze-thaw events in autumn and spring. Our results also showed, however, that the negative temperature effect of mosses on soil microbial activity was in part compensated by stimulatory effects of the moss layer, possibly linked to leaching of labile substrates from the moss. Furthermore, our results revealed that bryophytes efficiently sequester added N from wet deposition and thus prevent effects of increased atmospheric N deposition on soil N availability and soil processes.

Synthesis. Our study emphasizes the important role of mosses in carbon and nutrient cycling in high-latitude ecosystems and the potential strong impacts of reductions in moss abundance on microbial decomposition processes and nutrient availability in subarctic and boreal forests.

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.

Summary 1 Changing temperature regimes and precipitation patterns in the Subarctic will impact on vegetation composition and diversity including those of bryophyte and lichen communities, which are major drivers of high-latitude carbon and nutrient cycling and hydrology. 2 We investigated the relative importance of such impacts at different temporal, spatial and plant functional scales in subarctic Sphagnum fuscum-dominated peatlands, comprising both an in situ warming experiment and natural climatic and topographic gradients in northern Sweden and Norway. We applied multivariate analyses to investigate the relationships among cryptogam and vascular plant species composition and abiotic (temperature, moisture) and biotic (Sphagnum growth) regimes at various scales. 3 At the short-term temporal scale (4-year warming experiment), increased temperature yielded no clear effect on cryptogam or vascular plant species composition. Spatially, direct effects of temperature were decisive for overall species composition across regions (macro-scale) rather than within one region (meso-scale). Moisture and Sphagnum growth were drivers of species composition at all spatial scales, and Sphagnum growth itself depended on its position on the microtopographic gradient and on temperature. 4 Grouping of bryophytes and lichens at increasing scales of functional aggregation from species, growth form to the major higher taxon level (Sphagnum, other mosses, liverworts, lichens) revealed mostly increasing correlation with climate regimes and Sphagnum growth. Excluding liverworts from the analysis tended to reduce the correlation. 5 Abundances of lichens, liverworts, non-Sphagnum mosses and (to a lesser degree) vascular plants were negatively related to Sphagnum abundance. Few cryptogam and vascular plant species showed a positive relationship with Sphagnum abundance. Correspondingly, cryptogam species richness and Shannon Index on peatlands strongly declined as Sphagnum abundance increased, while indices for vascular plants showed no significant relationship. 6 Synthesis. Scale, be it spatial or functional, strongly determined which environmental drivers showed the clearest relationships with vegetation composition and diversity. Our findings will help to optimize predictions about long-term effects of climate on peatland vegetation composition, and subsequently its feedbacks to carbon and water cycles, at the regional scale.

1. Climate change in the subarctic is expected to influence vegetation composition, specifically bryophyte and lichen communities, thereby modifying litter decomposition rates and carbon (C) dynamics of these systems with possible feedbacks to climate.

2. In a 2‐year experiment, we investigated decomposition rates and chemical traits of 27 bryophytes, 17 lichens and 5 vascular plants in litter beds in subarctic Sweden. The majority of the sampled cryptogam species are widespread at higher northern latitudes.

3. Average 2‐year litter decomposition rates (exponential mass loss constant k) of lichen (0.44 ± 0.01) and vascular plant (0.56 ± 0.03) species were higher than that of bryophytes (0.11 ± 0.01), while within main cryptogam taxa, species identity was an important determinant of mass loss rates. At cryptogam group level, 2‐year litter mass loss of Sphagnum was significantly lower than for non‐Sphagnum mosses and liverworts. Within lichens, N₂‐fixing versus non‐N₂‐fixing lichens showed no variation in decomposability.

4. In a subset of the large species set, mass loss differed both among incubation environments (reflecting nutrient‐rich and poor birch forest and Sphagnum peatlands, respectively) and species. The pattern of mass loss across incubation environments was not consistent among cryptogam species. N₂‐fixing, in contrast to non‐N₂‐fixing lichens with lower nitrogen (N) levels displayed similar decomposition rates across incubation environments. Mass loss of non‐Sphagnum mosses was correlated with initial N irrespective of incubation environment.

5. Litter mass loss of cryptogam taxa could be predicted very well from infrared spectra of the initial chemical composition of the species, by application of Fourier transform infrared using an attenuated total reflectance probe. The initial macronutrient concentrations (N, phosphorus, C and cations) and initial litter pH correlated less well.

6. Synthesis. We showed comprehensively that decomposition rates of bryophytes are generally lower than those of lichens and vascular plants. Among bryophyte or lichen species there is also great variation in litter decomposability which depends strongly on species‐specific chemistry. Our data will help predict changing land surface feedback to C cycles and climate in cold biomes by understanding long‐term climate effects on litter decomposability through shifting vegetation composition.

Above the alpine tree line, bryophytes cover much of the tundra soil surface in dense, often monospecific carpets. Therefore, when climate warming enables tree seedling establishment above the tree line, interaction with the bryophyte layer is inevitable. Bryophytes are known to modify their environment in various ways. However, little is known about to which extent and by which mechanisms bryophytes affect the response of tree seedlings to climate warming.

We aimed to assess and understand the importance of bryophyte species identity and traits for tree seedling performance at tree line temperatures and their response to warmer conditions. Seedlings of two common, tree line-forming tree species (Betula pubescens and Pinus sylvestris) were planted into intact cushions of eight common tundra bryophyte species and bryophyte-free soil and grown for 18 weeks at current (7·0 °C) and near-future (30–50 years; 9·2 °C) tree line average growing-season temperatures. Seedling performance (biomass increase and N-uptake) was measured and related to bryophyte species identity and traits indicative of their impact on the environment.

Tree seedlings performed equally well or better in the presence of bryophytes than in bryophyte-free soil, which contrasts to their usually negative effects in milder climates. In addition, seedling performance and their response to higher temperatures depended on bryophyte species and seedlings of both species grew largest in the pan-boreal and subarctic bryophyte Hylocomium splendens. However, B. pubescens seedlings showed much stronger responses to higher temperatures when grown in bryophytes than in bryophyte-free soil, while the opposite was true for P. sylvestris seedlings. For B. pubescens, but not for P. sylvestris, available organic nitrogen of the bryophyte species was the trait that best predicted seedling responses to higher temperatures, likely because these seedlings had increased N-demands.

Synthesis. Climatically driven changes in bryophyte species distribution may not only have knock-on effects on vascular plant establishment, but temperature effects on seedling performance are themselves moderated by bryophytes in a species-specific way. Bryophyte traits can serve as a useful tool for understanding and predicting these complex interactions.

* When tree seedlings establish beyond the current tree line due to climate warming, they encounter existing vegetation, such as bryophytes that often dominate in arctic and alpine tundra. The stress gradient hypothesis (SGH) predicts that plant interactions in tundra become increasingly negative as climate warms and conditions become less harsh. However, for seedlings, climate warming might not result in lower winter stress, if insulating snow cover is reduced. * We aimed to understand if bryophytes facilitate seedling survival in a changing winter climate and if these effects of bryophytes on tree seedlings comply with the SGH along elevational gradients under contrasting snow conditions. * In the Swedish subarctic, we transplanted intact bryophyte cores covered by each of three bryophyte species and bryophyte-free control soil from above the tree line to two field common garden sites, representing current and future tree line air temperature conditions (i.e. current tree line elevation and a lower, warmer, elevation below the tree line). We planted seedlings of Betula pubescens and Pinus sylvestris into these cores and subjected them to experimental manipulation of snow cover during one winter. * In agreement with the SGH, milder conditions caused by increased snow cover enhanced the generally negative or neutral effects of bryophytes on seedlings immediately after winter. Furthermore, survival of P. sylvestris seedlings after one full year was higher at lower elevation, especially when snow cover was thinner. However, in contrast with the SGH, impacts of bryophytes on over-winter survival of seedlings did not differ between elevations, and impacts on survival of B. pubescens seedlings after 1 year was more negative at lower elevation. Bryophyte species differed in their effect on seedling survival after winter, but these differences were not related to their insulating capacity. * Synthesis. Our study demonstrates that interactions from bryophytes can modify the impacts of winter climate change on tree seedlings, and vice versa. These responses do not always comply with SGH, but could ultimately have consequences for large-scale ecological processes such as tree line shifts. These new insights need to be taken into account in predictions of plant species responses to climate change.

Primary succession encompasses the earliest and most fundamental stages of community assembly. The relative contributions of plant dispersal and ecosystem development, the main drivers that control the rate of primary succession, remain largely unknown, in spite of their central roles in community organization in general. Understanding the contribution of dispersal is especially critical because it casts light on our ability to manage succession resulting from climate change or anthropogenic disturbance. Here, we review the literature in order to understand when and where dispersal limits primary succession. Studies from many systems (tropical, temperate, boreal, arctic, alpine, floodplain forest, desert, mines, volcanic eruption and glacier forelands) show that dispersal limitation occurs consistently in the early stages of primary succession, suggesting that this is a general pattern. On the other hand, we found strong biases in study sites towards the Northern Hemisphere, temperate regions, volcanic eruptions and glacier forelands, which suggests that there are risks in drawing general conclusions about the importance of dispersal for primary succession for other biomes. Little is known about the contribution of dispersal to the later stages of primary succession. We present a recently developed method, multiscale chronosequence comparison that compares rates of succession in similar systems at relatively small and large spatial scales. Rapid succession at small scales compared to large scales suggests that seed dispersal influences succession over time periods of centuries and also limits the development of ecosystem functions such as vegetation cover and soil carbon accumulation. Synthesis: Dispersal limitation likely matters not only in the early stages but also in the late stages of primary succession by controlling the rate at which new species arrive. Both the accumulation of comparable case studies and novel approaches, such as multiscale chronosequence comparisons and factorial experiments, in contrasting biomes, are necessary to clarify the generality or context dependency of the role of dispersal in future primary successions undergoing changing climate.

1. Ecological processes in mixed‐species assemblages are not always an additive function of those in monocultures. In areas with high ground cover of bryophytes, renowned for their considerable water retention capacity, non‐additive interactions in mixed‐species cushions could play a key role in the ecosystem water economy.

2. We investigated mixture effects on external water loss in natural cushions with different species pairs of mosses and liverworts and the underlying mechanism of any non‐additivity through shoot characteristics and canopy traits.

3. Species mixtures in bryophyte cushions had both additive and non‐additive effects on the water economy, and these interactions were dependent on the composition of species assemblages and on plant tissue mass. Non‐additivity of species mixtures was positive, resulting in the improvement of water retention.

4. Our results suggest advantages for bryophyte species to grow smaller and denser when in mixtures. They appear to alter the surface exchange area to converge in size with their neighbours, thus controlling boundary‐layer properties and evaporation to reduce water loss.

5. Synthesis. A shift in bryophyte assemblages thus may influence ecohydrological processes of various ecosystems that cannot be simply predicted from the water economy of the component species when in monospecific cushions. In contrast to vascular plants, bryophytes do not compete for water but share it, and trait plasticity amongst bryophyte mixtures acts as a critical physical strategy in the community water economy.

1 Nutrient-rich litter of hemiparasites (and N-fixers) has the potential to influence ecosystem-level decomposition and nutrient cycling, which may be important in ecosystems where productivity is nutrient limited. This influence may be both direct, and indirect via impacts on the decomposition of other species’ litters in mixed-species litter layers. 2 We investigate the importance of these direct and indirect effects of sub-arctic hemiparasite litter in a litterbed study and a field litterbag experiment, and compare these data with those from a microcosm study also involving litter mixtures. 3 Field-placed litterbags demonstrate for the first time the direct importance of hemiparasite litter on decomposition and nutrient cycling in a field setting: litter of the hemiparasite Bartsia alpina lost 18 times more N over 2 years than Betula nana litter. 4 The majority of this N was released during the growing season, strongly supporting the proposal that hemiparasites can have a substantial direct impact on nutrient availability to other plants. The short spring and autumn periods were important for litter mass and N dynamics, with consequences for predicting effects of environmental change. 5 However, hemiparasite litter did not indirectly affect the decomposition rate or nutrient release from litter mixtures in field-placed litterbags. Furthermore, the direction and magnitude of litter mixing effects on mass loss varied strongly between methods; interactions were absent or negative in the litterbed, non-significant in field litterbags and positive in microcosms. 6 We thereby demonstrate the importance of the decomposition environment, rather than species-specific litter quality, in determining litter-mixing interactions, with implications for evaluating the impact of particular species on ecosystem carbon and nutrient fluxes via litter mixing 7 We conclude that hemiparasites have a particular functional role in sub-arctic heath systems: in effect they ‘short circuit’ generally slow nutrient cycles, with potential impacts on resource patchiness and local scale biodiversity.

1. Arctic landscapes are characterized by extreme vegetation patchiness, often with sharply defined borders between very different vegetation types. This patchiness makes it difficult to predict landscape-level C balance and its change in response to environment. 2. Here we develop a model of net CO2 flux by arctic landscapes that is independent of vegetation composition, using instead a measure of leaf area derived from NDVI (normalized-difference vegetation index). 3. Using the light response of CO2 flux (net ecosystem exchange, NEE) measured in a wide range of vegetation in arctic Alaska and Sweden, we exercise the model using various data subsets for parameter estimation and tests of predictions. 4. Overall, the model consistently explains similar to 80% of the variance in NEE knowing only the estimated leaf area index (LAI), photosynthetically active photon flux density (PPFD) and air temperature. 5. Model parameters derived from measurements made in one site or vegetation type can be used to predict NEE in other sites or vegetation types with acceptable accuracy and precision. Further improvements in model prediction may come from incorporating an estimate of moss area in addition to LAI, and from using vegetation-specific estimates of LAI. 6. The success of this model at predicting NEE independent of any information on species composition indicates a high level of convergence in canopy structure and function in the arctic landscape.

* Roots account for > 50% of net primary production in many ecosystems and are widely accepted as playing a fundamental role in ecosystem carbon and nutrient cycling. Despite this, the timing and controls of root production and the relationships between root and leaf phenology are still poorly understood in many plant communities, making it challenging to elucidate broad scale patterns and to predict ecosystem responses to a changing climate. Here, we investigate and compare the extent of synchrony of community

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.

1 The arctic environment is highly heterogeneous in terms of plant distribution and productivity. If we are to make regional scale predictions of carbon exchange it is necessary to find robust relationships that can simplify this variability. One such potential relationship is that of leaf area to photosynthetic CO2 flux at the canopy scale. 2 In this paper we assess the effectiveness of canopy leaf area in explaining variation in gross primary productivity (GPP): (i) across different vegetation types; (ii) at various stages of leaf development; and (iii) under enhanced nutrient availability. To do this we measure net CO2 flux light response curves with a 1 x 1 m chamber, and calculate GPP at a photosynthetic photon flux density (PPFD) of 600 mu mol m(-2) s(-1). 3 At a subarctic site in Sweden, we report 10-fold variation in GPP among natural vegetation types with leaf area index (LAI) values of 0.05-2.31 m(2) m(-2). At a site of similar latitude in Alaska we document substantially elevated rates of GPP in fertilized vegetation. 4 We can explain 80% of the observed variation in GPP in natural vegetation (including vegetation measured before deciduous leaf bud burst) by leaf area alone, when leaf area is predicted from measurements of normalized difference vegetation index (NDVI). 5 In fertilized vegetation the relative increase in leaf area between control and fertilized treatments exceeds the relative increase in GPP. This suggests that higher leaf area causes increased self-shading, or that lower leaf nitrogen per unit leaf area causes a reduction in the rate of photosynthesis. 6 The results of this study indicate that canopy leaf area is an excellent predictor of GPP in diverse low arctic tundra, across a wide range of plant functional types.

Sphagnum mosses are keystone species in northern peatlands. Notably, they play an important role in peatland carbon (C) cycling by regulating the composition and activity of microbial communities. However, it remains unclear whether information on Sphagnum phylogeny and/or traits-based composition (i.e. anatomical and morphological traits and metabolites) can be used to predict the structure of microbial communities and their functioning. Here we evaluated whether Sphagnum phylogeny and traits predict additional variation in peatland microbial community composition and functioning beyond what would be predicted from environmental characteristics (i.e. climatic and edaphic conditions). We collected Sphagnum and microbial data from five European peatlands distributed along a latitudinal gradient from northern Sweden to southern France. These data allowed us to assess Sphagnum anatomical and morphological traits and metabolites at different sites along changing environmental conditions. Using structural equation modelling (SEM) and phylogenetic distance analyses, we investigated the role of Sphagnum traits in shaping microbial community composition and functioning along with environmental conditions. We show that microbial community composition and traits varied independently from both Sphagnum phylogeny and the latitudinal gradient. Specifically, the addition of Sphagnum traits to climatic and edaphic variables to the SEM allowed it to explain a larger proportion of the explained variance (R2). This observation was most apparent for the biomass of decomposers (+42%) and phototrophs (+19%), as well as for growth yield microbial traits (+10%). As such, that Sphagnum metabolites were important drivers for microbial community structure and traits, while Sphagnum anatomical and morphological traits were poor predictors.

Synthesis. Our results highlight that Sphagnum metabolites are more likely to influence peatland microbial food web structure and functioning than Sphagnum anatomical and morphological traits. We provide further evidence that measurements of the plant metabolome, when combined with classical functional traits, improve our understanding of how the plants interact with their associated microbiomes.

Observed climate change in northern high latitudes is strongest in winter, but still relatively little is known about the effects of winter climate change on tundra ecosystems. Ongoing changes in winter climate and snow cover will change the intensity, duration and frequency of frost events. Bryophytes form a major component of northern ecosystems but their responses to winter climate changes are largely unknown.Here, we studied how changes in overall winter climate and snow regime affect frost damage in three common bryophyte taxa that differ in desiccation tolerance in a subarctic tundra ecosystem. We used a snow manipulation experiment where bryophyte cores were transplanted from just above the tree line to similar elevation (i.e. current cold climate) and lower elevation (i.e. near-future warmer climate scenario) in Abisko, Sweden. Here, we measured frost damage in shoots of Ptilidium ciliare, Hylocomium splendens and Sphagnum fuscum with the relative electrolyte leakage (REL) method, during late winter and spring in two consecutive years. We hypothesized that frost damage would be lower in a milder climate (low site) and higher under reduced snow cover and that taxa from moister habitats with assumed low desiccation tolerance would be more sensitive to lower temperature and thinner snow cover than those from drier and more exposed habitats.Contrary to our expectations, frost damage was highest at low elevation, while the effect of snow treatment differed across sites and taxa. At the high site, frost damage was reduced under snow addition in the taxon with the assumed lowest desiccation tolerance, S. fuscum. Surprisingly, frost damage increased with mean temperature in the bryophyte core of the preceding 14 days leading up to REL measurements and decreased with higher frost degree sums, that is, was highest in the milder climate at the low site.Synthesis Our results imply that climate warming in late winter and spring increases frost damage in bryophytes. Given the high abundance of bryophytes in tundra ecosystems, higher frost damage could alter the appearance and functioning of the tundra landscape, although the short and long-term effects on bryophyte fitness remain to be studied.