The impact of vegetation on soil temperatures in low Arctic and boreal regions is well documented, where dense vegetation reduces soil temperatures. However, equivalent empirical evidence for influence of sparse, low-stature high Arctic vegetation and its interaction with cloud cover is limited. Improved understanding of this relationship is vital, because soil temperatures drive important ecosystem processes, such as nutrient cycling and carbon fluxes. We investigated whether effects on soil temperature are found under smaller high Arctic vegetation in Svalbard, Norway, by modeling soil temperatures in relation to vegetation cover and height across forty plots, using July temperature recordings from 2020 to 2023. We found reduced minimum temperatures under thicker moss layers and taller forb and shrub vegetation, whereas organic layers reduced maximum temperatures. These effects were strongest under sunny conditions, whereas shrub cover showed weak interactions with cloud cover. This contrasts with lower latitudes, where shrubs exert strong shading effects and vegetation reduces maximum and raises minimum soil temperatures. This discrepancy might stem from smaller diurnal temperature fluctuations, lower vegetation height, and lower solar radiation in the high Arctic. Our results indicate that predicted vegetation shifts resulting from climate change may lead to complex soil temperature responses in the future.

Arctic ecosystems are facing increases in heavy summer rainfall events and increased year-to-year hydrological variability. However, the evidence base of impacts of heavy rainfall on Arctic vegetation is limited. The role of seasonal timing in determining heavy rainfall impacts on plant growth and the legacy impacts of such effects have not been quantified. We set up an irrigation experiment in several sites on the High Arctic archipelago of Svalbard, in which we simulated a doubling of average summer rainfall (50 mm) in individual additions of 10 mm at different timings throughout the season (early or late summer). Plant growth and phenology indicators (normalized difference vegetation index, vegetation height, specific leaf area, and senescence) of key plant species were monitored under irrigation and in the years after to capture legacy effects. Late-summer irrigation delayed end-of-summer declines in NDVI compared with control and early-summer irrigation treatments. We found subtle positive legacy effects of early as well as late-summer irrigation on NDVI in the following growing season. Irrigation only delayed senescence in Salix polaris at sites and moments where rainfall treatment compensated for low soil moisture levels prior to senescence. Other vegetation parameters did not show significant responses. Positive associations of plant growth variables with local soil moisture were strongest in late summer. Synthesis. Our findings indicate that the impact of heavy rainfall events on plant growth in the High Arctic is mediated by seasonal timing and local moisture regimes. Late-summer rainfall can extend the growing season for plants experiencing end-of-season water-limitation by delaying onset of senescence. Local soil moisture retention capacity appears to regulate this potential for growing season extension, implying that outcomes may be scalable across (micro)topographical and pedological gradients. The existence of subtle legacy effects suggests that previous years' hydrological conditions may marginally affect following years' plant growth.

Following rapid climate change, tundra plant communities are experiencing extensive compositional shifts. A conservation concern is the potential encroachment of boreal species into the tundra ('borealisation'). Tundra borealisation has been sporadically reported, but not systematically quantified. Here, we synthesised data from across 32 study areas, spanning 1137 plots and 287 vascular plant species, resurveyed between 1981 and 2023. We (i) quantified tundra borealisation as the colonisation and increase in abundance of Boreal and Boreal-Tundra species, (ii) assessed biogeographical, climatic and local borealisation drivers and (iii) identified species contributing to borealisation and their associated traits. Half of the plots experienced borealisation, although borealisation rates were not different to random expectation. Borealisation was greater in Eurasia, closer to the treeline, at higher elevations, in warmer and wetter regions, where climate change was limited, and where initial boreal abundance was lower. Boreal coloniser species were generally short-statured, and more often shrubs and graminoids. Boreal species colonised around three times less frequently than Boreal-Tundra species. Hence, our findings indicate that tundra borealisation is mainly driven by the spread of already established boreal-low Arctic tundra species. These plant community composition changes could have cascading impacts on land-atmosphere interactions, trophic dynamics and Indigenous and local livelihoods.

Low temperatures and nutrient limitation have shaped Arctic plant communities, which are now affected by biome‐wise changes in both climate and nutrient cycling. Rising temperatures are favouring taller plant species with more resource‐acquisitive traits across the Arctic tundra. Simultaneously, declines in seabird populations may reduce subsidies of marine‐derived nutrients to terrestrial ecosystems, potentially favouring more resource‐conservative plant traits. It is crucial to understand the consequences of these concurrent changes in climate and marine‐derived nutrient inputs from seabirds for the functional composition and roles of Arctic plant communities. We use a 'space‐for‐time approach' to compare the functional composition of vascular plant communities across two elevational gradients in High Arctic Svalbard, one where climate is the major environmental driver and one influenced by nutrient input from a seabird colony. We assess changes in 13 traits related to plant size, leaf economics and nutrient cycling along the two gradients, and we also explore the relative contributions of species turnover and intraspecific variation to total trait variation across and between the gradients. Elevation per se had little impact on the plant functional composition. Instead, plants at the top of the seabird nutrient gradient, closest to the nesting sites, were taller and had resource‐acquisitive trait values, such as larger and thicker leaves and higher leaf nutrient contents. Enriched soil δ 15 N‰ signatures at these sites correlated with resource‐acquisitive values of leaf area, specific leaf area, leaf dry matter content, leaf phosphorous content and with enriched leaf δ 15 N‰ signatures. This variation in leaf economic traits and isotopes was largely driven by intraspecific variation at the nutrient gradient, whereas species turnover dominated at the reference gradient. Our results are consistent with marine‐derived nutrient subsidies from seabirds being a major driver of functional trait variation in Arctic vegetation. Ongoing declines in seabird populations may therefore affect terrestrial primary producer communities in the Arctic and beyond, with potentially important but unknown implications for biodiversity, consumer and decomposer communities, and ecosystem processes. Read the free Plain Language Summary for this article on the Journal blog.

The Arctic is warming four times faster than the global average1 and plant communities are responding through shifts in species abundance, composition and distribution2-4. However, the direction and magnitude of local changes in plant diversity in the Arctic have not been quantified. Using a compilation of 42,234 records of 490 vascular plant species from 2,174 plots across the Arctic, here we quantified temporal changes in species richness and composition through repeat surveys between 1981 and 2022. We also identified the geographical, climatic and biotic drivers behind these changes. We found greater species richness at lower latitudes and warmer sites, but no indication that, on average, species richness had changed directionally over time. However, species turnover was widespread, with 59% of plots gaining and/or losing species. Proportions of species gains and losses were greater where temperatures had increased the most. Shrub expansion, particularly of erect shrubs, was associated with greater species losses and decreasing species richness. Despite changes in plant composition, Arctic plant communities did not become more similar to each other, suggesting no biotic homogenization so far. Overall, Arctic plant communities changed in richness and composition in different directions, with temperature and plant-plant interactions emerging as the main drivers of change. Our findings demonstrate how climate and biotic drivers can act in concert to alter plant composition, which could precede future biodiversity changes that are likely to affect ecosystem function, wildlife habitats and the livelihoods of Arctic peoples5,6.

The below-ground growing season often extends beyond the above-ground growing season in tundra ecosystems and as the climate warms, shifts in growing seasons are expected. However, we do not yet know to what extent, when and where asynchrony in above- and below-ground phenology occurs and whether variation is driven by local vegetation communities or spatial variation in microclimate. Here, we combined above- and below-ground plant phenology metrics to compare the relative timings and magnitudes of leaf and fine-root growth and senescence across microclimates and plant communities at five sites across the Arctic and alpine tundra biome. We observed asynchronous growth between above- and below-ground plant tissue, with the below-ground season extending up to 74% (~56 days) beyond the onset of above-ground leaf senescence. Plant community type, rather than microclimate, was a key factor controlling the timing, productivity, and growth rates of fine roots, with graminoid roots exhibiting a distinct 'pulse' of growth later into the growing season than shrub roots. Our findings indicate the potential of vegetation change to influence below-ground carbon storage as the climate warms and roots remain active in unfrozen soils for longer. Taken together, our findings of increased root growth in soils that remain thawed later into the growing season, in combination with ongoing tundra vegetation change including increased shrub and graminoid abundance, indicate increased below-ground productivity and altered carbon cycling in the tundra biome.

Svalbards permafrost is thawing as a direct consequence of climate change. In the Low Arctic, vegetation has been shown to slow down and reduce the active layer thaw, yet it is unknown whether this also applies to High Arctic regions like Svalbard where vegetation is smaller, sparser, and thus likely less able to insulate the soil. Therefore, it remains unknown which components of High Arctic vegetation impact active layer thaw and at which temporal scale this insulation could be effective. Such knowledge is necessary to predict and understand future changes in active layer in a changing Arctic. In this study we used frost tubes placed in study grids located in Svalbard with known vegetation composition, to monitor the progression of active layer thaw and analyze the relationship between vegetation composition, vegetation structure and snow conditions, and active layer thaw early in summer. We found that moss thickness, shrub and forb height, and vascular vegetation cover delayed soil thaw immediately after snow melt. These insulating effects attenuated as thaw progressed, until no effect on thaw depth was present after 8 weeks. High Arctic mosses are expected to decline due to climate change, which could lead to a loss in insulating capacity, potentially accelerating early summer active layer thaw. This may have important repercussions for a wide range of ecosystem functions such as plant phenology and decomposition processes.

The main aim of the present study has been the completion of genome size data for the diverse arctic-alpine A. ciliata species complex, with special focus on the unexplored arctic taxon A. pseudofrigida, the north-European A. norvegica, and A. gothica from Gotland (Sweden). Altogether, 46 individuals of these three Nordic taxa have been sampled from seven different regions and their genome size estimated using flow cytometry. Three other alpine taxa in the A. ciliata complex (A. multicaulis, A. ciliata subsp. ciliata, and A. ciliata subsp. bernensis) were also collected and analyzed for standardization purposes, comprising 20 individuals from six regions. A mean 2c value of 1.65 pg of DNA was recorded for A. pseudofrigida, 2.80 pg for A. norvegica, and 4.14 pg for A. gothica, as against the reconfirmed 2c value of 1.63 pg DNA for the type taxon A. ciliata subsp. ciliata. Our results presenting the first estimations of genome sizes for the newly sampled taxa, corroborate ploidy levels described in the available literature, with A. pseudofrigida being tetraploid (2n = 4x = 40), A. norvegica possessing predominantly 2n = 8x = 80, and A. gothica with 2n = 10x = 100. The present study also reconfirms genome size and ploidy level estimations published previously for the alpine members of this species complex. Reflecting a likely complex recent biogeographic history, the A. ciliata species group comprises a polyploid arctic-alpine species complex characterized by reticulate evolution, polyploidizations and hybridizations, probably associated with rapid latitudinal and altitudinal migrations in the Pleistocene–Holocene period.

The Arctic has a diverse terrestrial microarthropod fauna which overwinters in situ in soil and vegetation. This fauna is involved in key ecosystem processes, for example decomposition and pollination, and has tolerance to the Arctic's winter conditions. However, the Arctic is undergoing rapid change. Svalbard is experiencing warming rates up to four times the global average as well as alterations in precipitation (quantity and form; snow or rain) and wind direction. These will modify the conditions experienced by the overwintering fauna. Since laboratory experiments often fail to capture the diversity of environmental stressors, we employed a manipu-lation experiment using the naturally accumulating snow pack to moderate soil winter soil temperatures, combined with an extended incubator treatment, to map the duration limits of naturally induced cold tolerance. We demonstrate that the Collembola fauna can tolerate temperatures of-25 degrees C but that, in areas devoid of snow accumulation and when soil temperatures dip below-30 degrees C there is significant mortality. Furthermore, we demonstrate that exposure to a further extended 12 month period at-6 degrees C, as a simple model of a situation where snow cover is not lost during the short Arctic summer, results in additional mortality with relatively few Collembola surviving. By contrast, while oribatid mites displayed similar survival over a natural winter as the Collembola, they were highly resistant to the extended exposure at-6 degrees C, with no additional mortality occur-ring. We also documented survival amongst other invertebrate groups, including Nematocera and Brachycera larvae, Hemiptera (Acyrthosiphon svalbardicum), Coleoptera (Isochnus flagellum), and Araneae (Linyphiidae). We conclude that snow depth and winter air temperatures interact to regulate soil microarthropod populations over local scales and therefore the functioning of the Arctic soil ecosystem. Moreover, the environmental changes currently being observed in polar regions will continue to modify this fauna and its local and micro-scale distribution.

Practical lab skills are rarely directly assessed. To improve constructive alignment between the described learning outcomes of practical skills and assessment, we developed and tested a certification procedure for microscopy skills. The procedure was embedded into the ordinary learning activity, so no additional time was needed. Three slightly different protocols were developed within the framework of sociocultural learning theory and built like a skill ladder, including direct peer assessment and elements of gamified learning. The protocols varied slightly in the way students were prepared for the certification, the number of steps/levels of achievement, and the consequences of failing. We tested the protocols at three different academic institutions and within 11 courses of varying sizes and academic levels in biology or geology. Feedbacks were collected through online surveys (n = 207) or orally after sessions. One protocol provided instruction videos as preparation material. Instruction videos provided increased understanding of the task, but tactile training was most important for learning. Regardless of institution, type of preparation, and level of former experience, the certification procedure made students clearly more engaged in the exercise. The majority reported that the certification procedure increased their motivation to learn, increased their perceived learning outcome, and was appropriate for assessing practical skills. Students with no or little experience in microscopy before the exercise were more positive about the certification procedure compared to skilled students, and the level of engagement and preparation was higher when there were some consequences of failing. Most students felt comfortable being certified by peers, but some students expressed concern about peers making mistakes. The presented certification procedure can easily be adapted to assess other practical skills and, with some adjustments, be an efficient method for assessment-as-learning, merging formative- and summative assessment.

Research in global change ecology relies heavily on global climatic grids derived from estimates of air temperature in open areas at around 2 m above the ground. These climatic grids do not reflect conditions below vegetation canopies and near the ground surface, where critical ecosystem functions occur and most terrestrial species reside. Here, we provide global maps of soil temperature and bioclimatic variables at a 1‐km 2 resolution for 0–5 and 5–15 cm soil depth. These maps were created by calculating the difference (i.e. offset) between in situ soil temperature measurements, based on time series from over 1200 1‐km 2 pixels (summarized from 8519 unique temperature sensors) across all the world's major terrestrial biomes, and coarse‐grained air temperature estimates from ERA5‐Land (an atmospheric reanalysis by the European Centre for Medium‐Range Weather Forecasts). We show that mean annual soil temperature differs markedly from the corresponding gridded air temperature, by up to 10°C (mean = 3.0 ± 2.1°C), with substantial variation across biomes and seasons. Over the year, soils in cold and/or dry biomes are substantially warmer (+3.6 ± 2.3°C) than gridded air temperature, whereas soils in warm and humid environments are on average slightly cooler (−0.7 ± 2.3°C). The observed substantial and biome‐specific offsets emphasize that the projected impacts of climate and climate change on near‐surface biodiversity and ecosystem functioning are inaccurately assessed when air rather than soil temperature is used, especially in cold environments. The global soil‐related bioclimatic variables provided here are an important step forward for any application in ecology and related disciplines. Nevertheless, we highlight the need to fill remaining geographic gaps by collecting more in situ measurements of microclimate conditions to further enhance the spatiotemporal resolution of global soil temperature products for ecological applications.

The relative contribution of bryophytes to plant diversity, primary productivity, and ecosystem functioning increases towards colder climates. Bryophytes respond to environmental changes at the species level, but because bryophyte species are relatively difficult to identify, they are often lumped into one functional group. Consequently, bryophyte function remains poorly resolved. Here, we explore how higher resolution of bryophyte functional diversity can be encouraged and implemented in tundra ecological studies. We briefly review previous bryophyte functional classifications and the roles of bryophytes in tundra ecosystems and their susceptibility to environmental change. Based on shoot morphology and colony organization, we then propose twelve easily distinguishable bryophyte functional groups. To illustrate how bryophyte functional groups can help elucidate variation in bryophyte effects and responses, we compiled existing data on water holding capacity, a key bryophyte trait. Although plant functional groups can mask potentially high interspecific and intraspecific variability, we found better separation of bryophyte functional group means compared with previous grouping systems regarding water holding capacity. This suggests that our bryophyte functional groups truly represent variation in the functional roles of bryophytes in tundra ecosystems. Lastly, we provide recommendations to improve the monitoring of bryophyte community changes in tundra study sites.

Question Cold environments are stressful for vascular plants, and stress-tolerant non-vascular photoautotrophs, e.g. bryophytes and lichens, become relatively more important as competition from vascular plants decreases towards higher elevations. Under increasingly stressful climatic conditions, species assembly of vascular plants is commonly driven more by environmental filtering, and abiotic constraints may lead to increased similarity between species and thus low functional diversity. Because bryophytes and lichens are less constrained by harsh environments, environmental filtering may be less strong. Instead, reduced competition from vascular plants can potentially free up niche space for non-vascular vegetation. Therefore, we hypothesized that functional diversity of vascular plants, bryophytes and lichens are likely to show contrasting responses to elevation. Location Finse Alpine Research Centre, Southern Norway. Methods We utilized measurements of species abundance and functional traits of the three groups along a 500-m elevational gradient in alpine southern Norway and calculated multi-trait and single-trait functional dispersion. Results Functional diversity of vascular plants declined with elevation, indicating increased environmental filtering. By contrast, functional diversity of lichens and bryophytes increased along the same gradient, suggesting they are less exposed to environmental filtering, in line with our hypothesis. Instead, they likely benefit from the lower abundance of vascular plants at higher elevation. Conclusions Our findings suggest that different photoautotroph groups vary in how they respond to the same environmental gradient, which may contribute to contrasting community assembly processes across groups. These divergent responses likely occur because non-vascular vegetation differs from vascular plants in terms of nutrient acquisition and water economy strategies, meaning that they respond differently to the same factors. This highlights the need to explicitly consider bryophytes and lichens in community-level studies whenever these groups are abundant.

Through changes in climate and other environmental factors, alpine tundra ecosystems are subject to increased cover of erect shrubs, reduced predictability of rodent dynamics and changes in wild and domesticated herbivore densities. To predict the dynamics of these ecosystems, we need to understand how these simultaneous changes affect alpine vegetation. In the long term, vegetation dynamics may depend critically on seedling recruitment. To study drivers of alpine plant seedling recruitment, we set up a field experiment where we manipulated the opportunity for plant–plant interactions through vegetation removal and introduction of willow transplants, the occurrence of herbivory through caging of plots, and then sowed 14 species into the plots. We replicated the experiment in three common alpine vegetation types (heath, meadow and Salix shrubland) and recorded seedling emergence and survival over five years. Strong effects of vegetation removal and substantial differences in recruitment among dominant vegetation types suggested important effects of local vegetation on the recruitment success of vascular‐plant seedlings. Similarly, herbivore exclusion had strong positive effects on recruitment success. This effect arose primarily via reduced seedling mortality in plots from which herbivores had been experimentally excluded and became noticeably stronger over time. In contrast, we detected no consistent effects of experimental willow shrub introduction on seedling recruitment. These results demonstrate that large and small herbivores can affect alpine plant seedling recruitment negatively by trampling and feeding on seedlings. Importantly, the effects became stronger over time, suggesting that effects of herbivory on seedling recruitment accumulates over time and may relate to recruitment phases beyond initial seedling emergence.

Both vascular plants and lichens have evolved a high variety of secondary compounds, as defence against herbivores (e.g. Coley et al., 1985; Gauslaa, 2005), pathogens (Witzell & Martín, 2008), oxidative stress and light damage (Close & McArthur, 2002). Plants produce an array of different chemical groups of compounds, but in infertile lowproductive systems carbon-based secondary compounds (CBSCs) are Received: 19 November 2019 | Accepted: 18 October 2020 DOI: 10.1111/1365-2435.13712

Lichens and bryophytes are abundant primary producers in high latitude and high elevation ecosystems, and they play an important role in ecosystem processes such as decomposition and nutrient cycling. Despite their importance, little is known about the decomposability of lichens and bryophytes either among or within species, at the whole community level, or how this decomposability is affected by their functional traits. Here, we studied decomposability of lichens and bryophytes at the community‐level and individual species‐level (using 21 species and genera) collected from an elevational gradient in alpine Norway. In order to isolate the elevation effect on litter quality, we used a standardized laboratory bioassay to measure decomposability. In contrast to our expectations, we found that community‐level decomposability of lichens and bryophytes increased with elevation and thus decreasing temperature. In contrast, phosphorus release from the litter decreased with elevation while nitrogen release was unresponsive. Decomposability was explained by nutrient concentrations, litter pH and primary producer group identity (lichens versus bryophytes) at both the individual species and community levels. Species turnover (changes in species composition and abundance) was the main driver of decomposability across elevation at the community level, despite some of the traits explaining decomposability showing high intraspecific variability. Our study highlights the importance of among‐species variation in determining lichen and bryophyte decomposability. Further, the higher decomposability that we found for higher elevations suggests that global warming might result in a shift towards slower decomposable lichen and bryophyte species.

Surface sediments from 18 boxcores across the Canadian Arctic provide the first assessment of modern non-pollen, non-dinocyst palynomorph regional distribution patterns. Palynomorphs are abundant and well-preserved (~400–21,900 ind./g; average 5730 ind./g), comprising acritarchs, zoomorphs (invertebrate eggs, egg capsules, mouthparts), foraminiferal linings, tintinnid loricae, ciliate cysts, and freshwater algae. Acritarch P and Halodinium minor were the most abundant and widely distributed acritarchs (avg. ~410 and ~340 ind./g, respectively). Invertebrate eggs (avg. ~570 ind./g) and foraminiferal linings (avg. ~1600 ind./g) were prominent. Six tintinnid forms were identified, cf. Parafavella parumdentata (avg. ~40 ind./g; 10 stations) showing the widest distribution. Ciliate cysts included Strombidium? cyst A, a spherical, flask-shaped cell with short spines and a rounded base (avg. ~120 ind./g; 16 stations). Pediastrum sp. (avg. ~140 ind./g) and Botryococcus sp. (avg. ~1040 ind./g) were especially abundant in the west, likely delivered via rivers. Out of 14 environmental parameters only sedimentary biogenic silica (%Si(opal)) was significant in explaining palynomorph variation. At group level, tinitinnids showed the strongest positive correlation to %Si(opal). Individually, the tintinnid Acanthostomella? sp. 1, ciliate cyst Fusopsis? cyst A, and the acritarch Palaeostomocystis fritilla were almost solely and positively explained by %Si(opal). This suggests that %Si(opal) strongly determines palynomorph variability, particularly for microzooplankton, implying that siliceous organisms (diatoms) play an important role in shaping palynomorph content in the sedimentary record. Our data suggest that palynomorphs represent a valuable addition to organic-walled microfossils (dinocysts) routinely used in this region, offering a unique multiproxy glimpse into the ecosystem.

We studied the effects of different mat-forming lichens on microclimate and litter decomposition in an alpine ecosystem where lichens constitute a major part of the vegetation. We used four lichens with contrasting colour and water-holding capacity. We recorded soil temperature and moisture, and decomposition rate of plant litter under the different lichen mats and in bare soil. While soil temperature and freeze-thaw cycles were reduced under all lichen mats compared to bare soil, Cladonia rangiferina/Cladonia stygia insulated stronger than other lichens. Litter decomposition was faster under Flavocetraria nivalis than under Alectoria ochroleuca, but this was not related to lichen traits or microclimate. We conclude that insulation by lichen mats is stronger with higher water-holding capacity, and we suggest that microbial communities associated with different lichens have a larger effect on litter decomposition than soil conditions. Our findings highlight the importance of mat-forming lichens on microclimate and ecosystem processes in alpine environments.