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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.

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.

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

1. Alpine and Arctic treeline expansion depends on establishment of tree seedlings beyond the current treeline, which is expected to occur with climate warming. However, treelines often fail to respond to higher temperatures, and it is therefore likely that other environmental factors are important for seedling establishment.

2. We aimed to analyse our current understanding of how temperature and a range of other environmental drivers affect tree seedling establishment at the alpine and Arctic treelines world-wide and to assess the relative importance of temperature compared with other factors and how they interact.

3. We collected 366 observations from 76 experimental and observational papers for a qualitative analysis of the role of a wide range of environmental factors on tree seed germination, tree seedling growth, survival and natural occurrence. For a subset of these studies, where the experimental design allowed, we conducted formal meta-analyses to reveal if there were global drivers for different seedling life traits.

4. The analyses showed that a wide range of abiotic and biotic factors affected tree seedling establishment besides from temperature, including water, snow, nutrients, light and surrounding vegetation. The meta-analyses showed that different seedling life stages do not respond similarly to environmental factors. For example, temperature had positive effects on growth, while tree seedling survival and germination showed mixed responses to warming. Further, warming was as often as not the strongest factor controlling tree seedling establishment, when compared to with one of five other environmental factors. Moreover, warming effects often depended on other factors such as moisture or the presence of surrounding vegetation.

5. Our results suggest that population dynamics of trees at the alpine and Arctic treeline is responsive to environmental changes and show that there is a clear need for multifactorial studies if we want to fully understand and predict the interplay between warming and other environmental factors and their effect on tree seedling establishment across current treelines.

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 1year 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.

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.

Alpine and arctic tree line expansion depends on the establishment of tree seedlings above the current tree line, which is expected to occur with climate warming. However, tree lines often fail to respond to higher temperatures. Other environmental factors are therefore likely important for tree seedling establishment. Above the tree line, establishing seedlings encounter existing vegetation such as bryophytes, which often dominate in arctic and alpine tundra. Bryophytes modify their environment in various ways and may mediate climate change effects on establishing tree seedlings, and with that tree line expansion. The aim of this thesis was to understand if and how the environment, in particular bryophytes, mediates the impact of climate change on tree seedling establishment at the alpine and arctic tree line. This was explored by reviewing literature on tree seedling establishment at alpine and arctic tree lines globally. In addition, tree seedling survival and growth of Betula pubescens and Pinus sylvestris were assessed experimentally. Here, individuals were planted into mono-specific mats of different bryophytes species and exposed to warming and different precipitation regimes. The literature review revealed that besides from temperature, tree seedling establishment is affected by a wide range of abiotic and biotic factors including water, snow, nutrients, light, disturbance and surrounding vegetation. Furthermore the review revealed that for example vegetation can change tree seedling responses to climate change. The experiments showed that especially tree seedling survival was adversely affected by the presence of bryophytes and that the impacts of bryophytes were larger than those of the climate treatments. Seedling growth, on the other hand, was not hampered by the presence of bryophytes, which is in line with earlier findings that seedling survival, growth and seed germination do not respond similarly to changes in environmental conditions. Moreover, we found several indications that vegetation above the tree line, including bryophytes, mediated tree seedling responses to warming and precipitation or snow cover. This thesis shows that temperature alone should not be used to predict future tree seedling establishment above the alpine and arctic tree line and that extrapolations from climate envelope models could strongly over or under estimate tree line responses to warming. This underlines the value of multi-factorial studies for understanding the interplay between warming and other environmental factors and their effects on tree seedling establishment across current tree lines.

Nitrogen (N) fixation by N-2-fixing bacteria (diazotrophs) is the primary N input to pristine ecosystems like boreal forests and subarctic and arctic tundra. However, the contribution by the various diazotrophs to habitat N-2 fixation remains unclear. We present results from in situ assessments of N-2 fixation of five diazotroph associations (with a legume, lichen, feather moss, Sphagnum moss and free-living) incorporating the ground cover of the associations in five typical habitats in the subarctic (wet and dry heath, polygon-heath, birch forest, mire). Further, we assessed the importance of soil and air temperature, as well as moisture conditions for N-2 fixation. Across the growing season, the legume had the highest total as well as the highest fraction of N-2 fixation rates at habitat level in the heaths (> 85 % of habitat N-2 fixation), whereas the free-living diazotrophs had the highest N-2 fixation rates in the polygon heath (56 %), the lichen in the birch forest (87 %) and Sphagnum in the mire (100 %). The feather moss did not contribute more than 15 % to habitat N-2 fixation in any of the habitats despite its high ground cover. Moisture content seemed to be a major driver of N-2 fixation in the lichen, feather moss and free-living diazotrophs. Our results show that the range of N-2 fixers found in pristine habitats contribute differently to habitat N-2 fixation and that ground cover of the associates does not necessarily mirror contribution.

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 warming will induce changes in Arctic ecosystem carbon balance, but besides climate, nitrogen availability is a critical controlling factor of carbon cycling. It is therefore essential to obtain knowledge on the influence of a changing climate on nitrogen fixation, as this process is the main source of new nitrogen to arctic ecosystems. In order to gain information on future nitrogen fixation rates in a changing climate, we studied the effects of two decades of warming with passive greenhouses, shading with sackcloth, and fertilization with NPK fertilizer on nitrogen fixation rates. To expand the knowledge on species-specific responses, we measured nitrogen fixation associated with two moss species: Hylocomium splendens and Aulacomnium turgidum. Our expectations of decreased nitrogen fixation rates in the fertilizer and shading treatments were met. However, contrary to our expectation of increased nitrogen fixation in the warming treatment, we observed either no change (Hylocomium) or a decrease (Aulacomnium) in fixation in the warmed plots. We hypothesize that this could be due to moss-specific responses or to long-term induced effects of the warming. For example, we observed that the soil temperature increase induced by the warming treatment was low and insignificant as vegetation height and total vascular plant cover of the warmed plots increased, and moss cover decreased. Hence, truly long-term studies lasting more than two decades provide insights on changes in key biogeochemical processes, which differ from more transient responses to warming in the Arctic.