Carboxylation capacity is the main limitation of carbon assimilation in High Arctic shrubs.

Increases in shrub height, biomass and canopy cover are key whole-plant features of warming-induced vegetation change in tundra. We investigated leaf functional traits underlying photosynthetic capacity of Arctic shrub species, particularly its main limiting processes such as mesophyll conductance. In this nutrient-limited ecosystem, we expect leaf nitrogen concentration to be the main limiting factor for photosynthesis. We measured the net photosynthetic rate at saturated light (Asat) in three Salix species throughout a glacial valley in High-Arctic tundra and used a causal approach to test relationships between leaf stomatal and mesophyll conductances (gsc, gm), carboxylation capacity (Vcmax), nitrogen and phosphorus concentration (Narea, Parea) and leaf mass ratio (LMA). Arctic Salix species showed no difference in Asat compared to a global data set, while being characterized by higher Narea, Parea and LMA. Vcmax, gsc and gm independently increased Asat, with Vcmax as its main limitation. We highlighted a nitrogen-influenced pathway for increasing photosynthesis in the two prostrate mesic habitat species. In contrast, the erect wetland habitat Salix richardsonii mainly increased Asat with increasing gsc. Overall, our study revealed high photosynthetic capacities of Arctic Salix species but contrasting regulatory pathways that may influence shrub ability to respond to environmental changes in High Arctic tundra.

Wildfire effects on mercury fate in soils of North-Western Siberia.

Arctic soils store 49 Gg mercury (Hg) - an extremely toxic heavy metal, whereas soil Hg can be released to the atmosphere by wildfires. For the first time we investigated the effects of wildfires on the fate of soil Hg in North-Western (NW) Siberia based on GIS maps of areas burned during the last 38 years and a field paired comparison of unburned and burned areas in tundra (mosses, lichens, some grasses, and shrubs) and forest-tundra (multi-layered canopy of larch trees, shrubs, mosses, and lichens). These field surveys were deepened by soil controlled burning to assess the Hg losses from organic horizon and mineral soil. The soil Hg stocks in the organic horizon and in the top 10 cm of the mineral soil were 3.3 ± 0.6 and 16 ± 3 mg Hg m-2 for unburned tundra and forest-tundra, respectively. After the burning by wildfires, the soil Hg stocks decreased to 2.4 ± 0.1 and 6.6 ± 0.2 mg Hg m-2 for tundra and forest-tundra, respectively. By the averages annual burned areas in NW Siberia 527 km2, wildfires in tundra and forest-tundra released 0.19 and 2.9 Mg soil Hg per year, respectively, corresponding to 28 % and 59 % of the initial soil Hg stocks. These direct effects of wildfires on Hg volatilization are raised by indirect post-pyrogenic consequences on Hg fate triggered by the vegetation succession and adsorption of atmospheric Hg on the surface of charred biomass. Charred lichens and trees accumulated 4-16 times more Hg compared to the living biomass. Blackened burned vegetation and soil reduced surface albedo and slowly increased soil temperatures in Arctic after wildfires. This created favorable conditions for seeding grasses and shrubs after wildfire and transformed burned high-latitude ecosystems into greener areas, increasing their capacity to trap atmospheric Hg by vegetation, which partly compensate the burning losses of soil Hg.

Diversity of lichen mycobionts and photobionts and their relationships in the Ny-Ålesund region (Svalbard, High Arctic).

Lichens are dual organisms, with one major mycobiont and one major photobiont in each lichen symbiosis, which can survive extreme environmental conditions in the Arctic. However, the diversity and distribution of lichen photobionts in the Arctic remain poorly understood compared to their mycobiont partners. This study explored the diversity of lichen mycobionts and photobionts in 197 lichen samples collected from the Ny-Ålesund region (Svalbard, High Arctic). The nuclear ribosomal internal transcribed spacer (ITS) regions were sequenced and phylogenetic analyses were performed. The relationships between mycobionts and photobionts, as well as the association patterns, were also investigated. A total of 48 species of lichen mycobionts (16 families, nine orders) and 31 species/lineages of photobionts were identified. These 31 photobiont species belonged to one class (Trebouxiophyceae) and five genera, including 22 species of Trebouxia, five species of Asterochloris, two species of Chloroidium, one species of Symbiochloris, and one species of Coccomyxa. The results indicated that most analyzed lichen mycobionts could associate with multiple photobiont species, and the photobionts also exhibited a similar pattern. The results provided an important reference dataset for characterizing the diversity of lichen mycobionts and photobionts in the High Arctic region.

Changes in soil and plant carbon pools after 9 years of experimental summer warming and increased snow depth.

Climate change can have positive and negative effects on the carbon pools and budgets in soil and plant fractions, but net effects are unclear and expected to vary widely within the arctic. We report responses after nine years (2012-2021) of increased snow depth (snow fences) and summer warming (open top chambers) and the combination on soil and plant carbon pools within a tundra ecosystem in West Greenland. Data included characteristics of depth-specific soil samples, including the rhizosphere soil, as well as vegetation responses of NDVI-derived traits, plant species cover and aboveground biomass, litter and roots. Furthermore, natural vegetation growth through the study period was quantified based on time-integrated NDVI Landsat 8 satellite imagery. Our results showed that summer warming resulted in a significant and positive vegetation response driven by the deciduous low shrub Betula nana (no other vascular plant species), while snow addition alone resulted in a significant negative response for Betula. A significant positive effect of summer warming was also observed for moss biomass, possibly driven increasing shade by Betula. The aboveground effects cascaded to belowground traits. The rhizosphere soil characteristics differed from those of the bulk soil regardless of treatment. Only the rhizosphere fraction showed responses to treatment, as soil organic C stock increased in near-surface and top 20 cm with summer warming. We observed no belowground effects from snow addition. The study highlights the plant species response to treatment followed by impacts on belowground C pools, mainly driven by dead fine roots via Betula nana. We conclude that the summer warming treatment and snow addition treatment separately showed opposing effects on ecosystem C pools, with lack of interactive effects between main factors in the combination treatment. Furthermore, changes in soil C are more clearly observed in the rhizosphere soil fraction, which should receive more attention in the future.

Impact of three decades of warming, increased nutrient availability, and increased cloudiness on the fluxes of greenhouse gases and biogenic volatile organic compounds in a subarctic tundra heath.

Climate change is exposing subarctic ecosystems to higher temperatures, increased nutrient availability, and increasing cloud cover. In this study, we assessed how these factors affect the fluxes of greenhouse gases (GHGs) (i.e., methane (CH4), nitrous oxide (N2O), and carbon dioxide (CO2)), and biogenic volatile organic compounds (BVOCs) in a subarctic mesic heath subjected to 34 years of climate change related manipulations of temperature, nutrient availability, and light. GHGs were sampled from static chambers and gases analyzed with gas chromatograph. BVOCs were measured using the push-pull method and gases analyzed with chromatography-mass spectrometry. The soil temperature and moisture content in the warmed and shaded plots did not differ significantly from that in the controls during GHG and BVOC measurements. Also, the enclosure temperatures during BVOC measurements in the warmed and shaded plots did not differ significantly from temperatures in the controls. Hence, this allowed for assessment of long-term effects of the climate treatment manipulations without interference of temperature and moisture differences at the time of measurements. Warming enhanced CH4 uptake and the emissions of CO2, N2O, and isoprene. Increased nutrient availability increased the emissions of CO2 and N2O but caused no significant changes in the fluxes of CH4 and BVOCs. Shading (simulating increased cloudiness) enhanced CH4 uptake but caused no significant changes in the fluxes of other gases compared to the controls. The results show that climate warming and increased cloudiness will enhance CH4 sink strength of subarctic mesic heath ecosystems, providing negative climate feedback, while climate warming and enhanced nutrient availability will provide positive climate feedback through increased emissions of CO2 and N2O. Climate warming will also indirectly, through vegetation changes, increase the amount of carbon lost as isoprene from subarctic ecosystems.

Metabarcoding using nanopore sequencing enables identification of diverse and zoonotic vector-borne pathogens from neglected regions: A case study investigating dogs from Bhutan.

The diversity and prevalence of canine vector-borne pathogens (VBPs) in Bhutan have to date remained unexplored, whilst recent epidemiological surveys in other South Asian nations have found diseases caused by VBPs to be rife in local dog populations. Importantly, many of such VBPs can infect people as well, with a building body of evidence identifying potentially zoonotic rickettsial organisms infecting humans in Bhutan. Given the lack of data on canine pathogens in Bhutan we employed a suite of deep-sequencing metabarcoding methods using Oxford Nanopore Technologies' MinION™ device to holistically characterise the bacterial, apicomplexan and filarial worm blood-borne pathogens of dogs in the country's south. Of the 95 stray, owned and community dogs sampled 78% (95% CI = 69%-85%) were infected with at least one VBP. Pathogen species detected were highly diverse including the bacteria Mycoplasma haemocanis in 16% (95% CI: 10-24%), Ehrlichia canis in 4% (95% CI: 2-10%), Anaplasma platys in 2% (95% CI: 0.5-7%) of dogs as well as the zoonotic species Bartonella clarridgeiae in 1% (95% CI: 0.1-6%), a potentially novel Bartonella spp. and an Ehrlichia chaffeensis-like bacterium, both in 1% (95% CI: 0.1-6%) of dogs. The apicomplexan haemoparasites Hepatozoon canis in 62% (95% CI: 52-71%), Babesia gibsoni in 45% (95% CI: 36-55%) and Babesia vogeli in 3% (95% CI: 1-9%) of dogs were also detected. Finally, 5% (95% CI: 2-12%) of dogs were found to be infected with the filarioid Acanthocheilonema reconditum and 1% (95% CI: 0.1-6%) with zoonotic Dirofilaria sp. hongkongensis. One canine was found positive to the filarioid Setaria tundra, a species normally found infecting cervids. The elucidated diversity of VBP communities highlights the strength of assumption-free diagnostics, such as metabarcoding, in detecting rare, novel, and unexpected pathogens. This approach to identifying pathogen diversity is of critical importance when investigating regions and populations that have thus far been neglected, with the findings aiding the development of future One Health informed strategies for disease control.

Warming alters cascading effects of a dominant arthropod predator on fungal community composition in the Arctic.

Rapid climate change in the Arctic is altering microbial structure and function, with important consequences for the global ecosystem. Emerging evidence suggests organisms in higher trophic levels may also influence microbial communities, but whether warming alters these effects is unclear. Wolf spiders are dominant Arctic predators whose densities are expected to increase with warming. These predators have temperature-dependent effects on decomposition via their consumption of fungal-feeding detritivores, suggesting they may indirectly affect the microbial structure as well. To address this, we used a fully factorial mesocosm experiment to test the effects of wolf spider density and warming on litter microbial structure in Arctic tundra. We deployed replicate litter bags at the surface and belowground in the organic soil profile and analyzed the litter for bacterial and fungal community structure, mass loss, and nutrient characteristics after 2 and 14 months. We found there were significant interactive effects of wolf spider density and warming on fungal but not bacterial communities. Specifically, higher wolf spider densities caused greater fungal diversity under ambient temperature but lower fungal diversity under warming at the soil surface. We also observed interactive treatment effects on fungal composition belowground. Wolf spider density influenced surface bacterial composition, but the effects did not change with warming. These findings suggest a widespread predator can have indirect, cascading effects on litter microbes and that effects on fungi specifically shift under future expected levels of warming. Overall, our study highlights that trophic interactions may play important, albeit overlooked, roles in driving microbial responses to warming in Arctic terrestrial ecosystems. IMPORTANCE: The Arctic contains nearly half of the global pool of soil organic carbon and is one of the fastest warming regions on the planet. Accelerated decomposition of soil organic carbon due to warming could cause positive feedbacks to climate change through increased greenhouse gas emissions; thus, changes in ecological dynamics in this region are of global relevance. Microbial structure is an important driver of decomposition and is affected by both abiotic and biotic conditions. Yet how activities of soil-dwelling organisms in higher trophic levels influence microbial structure and function is unclear. In this study, we demonstrate that predicted changes in abundances of a dominant predator and warming interactively affect the structure of litter-dwelling fungal communities in the Arctic. These findings suggest predators may have widespread, indirect cascading effects on microbial communities, which could influence ecosystem responses to future climate change.

Phosphorus Interactions with Iron in Undisturbed and Disturbed Arctic Tundra Ecosystems.

Phosphorus (P) limitation often constrains biological processes in Arctic tundra ecosystems. Although adsorption to soil minerals may limit P bioavailability and export from soils into aquatic systems, the contribution of mineral phases to P retention in Arctic tundra is poorly understood. Our objective was to use X-ray absorption spectroscopy to characterize P speciation and associations with soil minerals along hillslope toposequences and in undisturbed and disturbed low-lying wet sedge tundra on the North Slope, AK. Biogenic mats comprised of short-range ordered iron (Fe) oxyhydroxides were prevalent in undisturbed wet sedge meadows. Upland soils and pond sediments impacted by gravel mining or thermokarst lacked biogenic Fe mats and were comparatively iron poor. Phosphorus was primarily contained in organic compounds in hillslope soils but associated with Fe(III) oxyhydroxides in undisturbed wet sedge meadows and calcium (Ca) in disturbed pond sediments. We infer that phosphate mobilized through organic decomposition binds to Fe(III) oxyhydroxides in wet sedge, but these associations are disrupted by physical disturbance that removes Fe mats. Increasing disturbances of the Arctic tundra may continue to alter the mineralogical composition of soils at terrestrial-aquatic interfaces and binding mechanisms that could inhibit or promote transport of bioavailable P from soils to aquatic ecosystems.

Greening of Svalbard.

Svalbard, located between 76°30'N and 80°50'N, is among the regions in the world with the most rapid temperature increase. We processed a cloud-free time-series of MODIS-NDVI for Svalbard. The dataset is interpolated to daily data during the 2000-2022 period with 232 m pixel resolution. The onset of growth, with a clear phenological definition, has been mapped each year. Then the integrated NDVI from the onset (O) of growth each year to the time of average (2000-2022) peak (P) of growth (OP NDVI) have been calculated. OP NDVI has previously shown high correlation with field-based tundra productivity. Daily mean temperature data from 11 meteorological stations are compared with the NDVI data. The OP NDVI values show very high and significant correlation with growing degree days computed from onset to time of peak of growth for all the meteorological stations used. On average for the entire Svalbard, the year 2016 first had the highest greening (OP NDVI values) recorded since the year 2000, then the greening in 2018 surpassed 2016, then 2020 surpassed 2018, and finally 2022 was the year with the overall highest greening by far for the whole 2000-2022 period. This shows a rapid recent greening of Svalbard very strongly linked to temperature increase, although there are regional differences: the eastern parts of Svalbard show the largest variability between years, most likely due to variability in the timing of sea-ice break-up in adjacent areas. Finally, we find that areas dominated by manured moss-tundra in the polar desert zone require new methodologies, as moss does not share the seasonal NDVI dynamics of tundra communities.

Pixel walking along the boreal forest-Arctic tundra ecotone: Large scale ground-truthing of satellite-derived greenness (NDVI).

In this Technical Advance, we describe a novel method to improve ecological interpretation of remotely sensed vegetation greenness measurements that involved sampling 24,395 Landsat pixels (30 m) across 639 km of Alaska's central Brooks Range. The method goes well beyond the spatial scale of traditional plot-based sampling and thereby more thoroughly relates ground-based observations to satellite measurements. Our example dataset illustrates that, along the boreal-Arctic boundary, vegetation with the greatest Landsat Normalized Difference Vegetation Index (NDVI) is taller than 1 m, woody, and deciduous; whereas vegetation with lower NDVI tends to be shorter, evergreen, or non-woody. The field methods and associated analyses advance efforts to inform satellite data with ground-based vegetation observations using field samples collected at spatial scales that closely match the resolution of remotely sensed imagery.

Shifts in mycorrhizal types of fungi and plants in response to fertilisation, warming and herbivory in a tundra grassland.

Climate warming is severely affecting high-latitude regions. In the Arctic tundra, it may lead to enhanced soil nutrient availability and interact with simultaneous changes in grazing pressure. It is presently unknown how these concurrently occurring global change drivers affect the root-associated fungal communities, particularly mycorrhizal fungi, and whether changes coincide with shifts in plant mycorrhizal types. We investigated changes in root-associated fungal communities and mycorrhizal types of the plant community in a 10-yr factorial experiment with warming, fertilisation and grazing exclusion in a Finnish tundra grassland. The strongest determinant of the root-associated fungal community was fertilisation, which consistently increased potential plant pathogen abundance and had contrasting effects on the different mycorrhizal fungal types, contingent on other treatments. Plant mycorrhizal types went through pronounced shifts, with warming favouring ecto- and ericoid mycorrhiza but not under fertilisation and grazing exclusion. Combination of all treatments resulted in dominance by arbuscular mycorrhizal plants. However, shifts in plant mycorrhizal types vs fungi were mostly but not always aligned in their magnitude and direction. Our results show that our ability to predict shifts in symbiotic and antagonistic fungal communities depend on simultaneous consideration of multiple global change factors that jointly alter plant and fungal communities.

Upscaling vascular aboveground biomass and topsoil moisture of subarctic fens from Unoccupied Aerial Vehicles (UAVs) to satellite level.

Arctic and subarctic ecosystems are experiencing rapid changes in vegetation composition and productivity due to global warming. Tundra wetlands are especially susceptible to these changes, which may trigger shifts in soil moisture dynamics. It is therefore essential to accurately map plant biomass and topsoil moisture. In this study, we mapped total, wood, and leaf above ground biomass and topsoil moisture in subarctic tundra wetlands located between Norway and Finland by linking models derived from Unoccupied Aerial Vehicles with multiple satellite data sources using the Extreme Gradient Boosting algorithm. The most accurate predictions for topsoil moisture (R2 = 0.73) used a set of red edge-based vegetation indices with a spatial resolution of 20 m per pixel. On the contrary, wood biomass showed the lowest accuracies across all tested models (R2 = 0.38). We found a trade-off between the spatial resolution and the performance of upscaling models, where smaller pixel sizes generally led to lower accuracies. However, we were able to compensate for reduced accuracy at smaller pixel sizes using Copernicus phenology metrics. A modelling uncertainty assessment revealed that the uncertainty of predictions increased with decreasing pixel sizes and increasing number of co-predictors. Our approach could improve efforts to map and monitor changes in vegetation at regional to pan-Arctic scales.

Dirofilaria sp. and Blood Meal Analysis in Mosquitoes Collected in Vojvodina and Macva, and the First Report of Setaria tundra (Issaitshikoff & Rajewskaya, 1928) in Serbia.

Dirofilaria immitis and D. repens are the two most widespread and important species of mosquito-borne nematodes, posing a significant threat to veterinary health and particularly affecting canines and felines. While D. immitis causes cardiopulmonary dirofilariasis, D. repens causes subcutaneous infections in dogs and other carnivores. Despite the extensive knowledge on these parasites, little is known about their natural vectors in Serbia. The parasite Setaria tundra, known to infect deer, has not yet been detected in Serbia but has been documented in neighboring countries. Thus, the aim of this study was to (i) further map out Dirofilaria sp. hotspots in the Vojvodina Province and detect S. tundra for the first time, (ii) detect positive mosquito species that can provide insights into how the nematodes spread in Serbia, and (iii) analyze the blood-fed female mosquitoes of species found to be infected, in order to identify the potential source of parasite infection. A total of 2902 female mosquitoes were collected across 73 locations during 2021 and 2022. Molecular biology methods, based on conventional PCR, were used to analyze non-blood-fed (2521 specimens) and blood-fed (381 specimens) female mosquitos, in order to detect filarial nematode presence and identify blood-meal sources, respectively. When the parasite genome was detected, the amplicon (cox1 gene, 650 bp fragment) was sent for Sanger sequencing, further confirming the presence of nematodes and species assignation. D. immitis was detected in three Culex pipiens mosquitoes collected in Zrenjanin (August 2021) and Glogonj and Svetozar Miletic (both in July 2021). Additionally, Setaria tundra was detected in Aedes vexans collected in Idos (mid-August 2021) and Aedes caspius, which was collected in Mali Idos (end of July 2021). This work identifies two new locations where D. immitis occurs in Vojvodina, and is the first report of S. tundra in Serbian territory. Blood-meal analysis provided insights into the preferences of mosquitoes that were positive for Dirofilaria sp. and S. tundra.

Effects of resource availability and interspecific interactions on Arctic and red foxes' winter use of ungulate carrion in the Fennoscandian low-Arctic tundra.

In the Arctic tundra, predators face recurrent periods of food scarcity and often turn to ungulate carcasses as an alternative food source. As important and localized resource patches, carrion promotes co-occurrence of different individuals, and its use by predators is likely to be affected by interspecific competition. We studied how interspecific competition and resource availability impact winter use of carrion by Arctic and red foxes in low Arctic Fennoscandia. We predicted that the presence of red foxes limits Arctic foxes' use of carrion, and that competition depends on the availability of other resources. We monitored Arctic and red fox presence at supp lied carrion using camera traps. From 2006 to 2021, between 16 and 20 cameras were active for 2 months in late winter (288 camera-winters). Using a multi-species dynamic occupancy model at a week-to-week scale, we evaluated the use of carrion by foxes while accounting for the presence of competitors, rodent availability, and supplemental feeding provided to Arctic foxes. Competition affected carrion use by increasing both species' probability to leave occupied carcasses between consecutive weeks. This increase was similar for the two species, suggesting symmetrical avoidance. Increased rodent abundance was associated with a higher probability of colonizing carrion sites for both species. For Arctic foxes, however, this increase was only observed at carcasses unoccupied by red foxes, showing greater avoidance when alternative preys are available. Supplementary feeding increased Arctic foxes' carrion use, regardless of red fox presence. Contrary to expectations, we did not find strong signs of asymmetric competition for carrion in winter, which suggests that interactions for resources at a short time scale are not necessarily aligned with interactions at the scale of the population. In addition, we found that competition for carcasses depends on the availability of other resources, suggesting that interactions between predators depend on the ecological context.

Bacterial and fungal communities in sub-Arctic tundra heaths are shaped by contrasting snow accumulation and nutrient availability.

Climate change is affecting winter snow conditions significantly in northern ecosystems but the effects of the changing conditions for soil microbial communities are not well-understood. We utilized naturally occurring differences in snow accumulation to understand how the wintertime subnivean conditions shape bacterial and fungal communities in dwarf shrub-dominated sub-Arctic Fennoscandian tundra sampled in mid-winter, early, and late growing season. Phospholipid fatty acid (PLFA) and quantitative PCR analyses indicated that fungal abundance was higher in windswept tundra heaths with low snow accumulation and lower nutrient availability. This was associated with clear differences in the microbial community structure throughout the season. Members of Clavaria spp. and Sebacinales were especially dominant in the windswept heaths. Bacterial biomass proxies were higher in the snow-accumulating tundra heaths in the late growing season but there were only minor differences in the biomass or community structure in winter. Bacterial communities were dominated by members of Alphaproteobacteria, Actinomycetota, and Acidobacteriota and were less affected by the snow conditions than the fungal communities. The results suggest that small-scale spatial patterns in snow accumulation leading to a mosaic of differing tundra heath vegetation shapes bacterial and fungal communities as well as soil carbon and nutrient availability.

Analysis of taiga and tundra lake browning trends from 2002 to 2021 using MODIS data.

Lakes in taiga and tundra regions may be silently undergoing changes due to global warming. One of those changes is browning in lake color. The browning interacts with the carbon cycle, ecosystem dynamics, and water quality in freshwater systems. However, spatiotemporal variabilities of browning in these regions have not been well documented. Using MODIS remote sensing reflectance at near ultraviolet wavelengths from 2002 to 2021 on the Google Earth Engine platform, we quantified long-term browning trends across 7616 lakes (larger than 10 km2) in taiga and tundra biomes. These lakes showed an overall decreased trend in browning (Theil-Sen Slope = 0.00015), with ∼36% of these lakes showing browning trends, and ∼1% of these lakes showing statistically significant (p-value <0.05) browning trends. The browning trends more likely occurred in small lakes in high latitude, low ground ice content regions, where air temperature increased and precipitation decreased. While temperature is projected to increase in response to climate change, our results provide one means to understand how biogeochemical cycles and ecological dynamics respond to climate change.

Vegetation biomass and topography are associated with seasonal habitat selection and fall translocation behavior in Arctic hares.

Habitat selection theory suggests that environmental features selected at coarse scales reveal fundamental factors affecting animal fitness. When these factors vary across seasons, they may lead to large-scale movements, including long-distance seasonal migrations. We analyzed the seasonal habitat selection of 25 satellite-tracked Arctic hares from a population on Ellesmere Island (Nunavut, Canada) that relocated over 100 km in the fall. Since no other lagomorph is known to perform such extensive movements, this population offered an ideal setting to test animal movement and habitat selection theory. On summer grounds hares selected low elevation areas, while on winter grounds they selected high vegetation biomass, high elevation, and steep slopes. During fall relocation, they alternated between stopover and traveling behavioral states (ratio 2:1). Stopover locations were characterized by higher vegetation heterogeneity and lower rugosity than traveling locations, while vegetation biomass and elevation interacted to explain stopover locations in a more complex way. The selected combination of environmental features thus varied across seasons and behavioral states, in a way broadly consistent with predictions based on the changing food and safety needs of hares. Although causality was not demonstrated, our results improve our understanding of long-distance movements and habitat selection in Arctic hares, as well as herbivore ecology in the polar desert. Results also provide strong support to animal movement and habitat selection theory, by showing how some important hypotheses hold when tested in a species phylogenetically distinct from most animal models used in this research field.

Plant-soil interactions alter nitrogen and phosphorus dynamics in an advancing subarctic treeline.

Treelines advance due to climate warming. The impacts of this vegetation shift on plant-soil nutrient cycling are still uncertain, yet highly relevant as nutrient availability stimulates tree growth. Here, we investigated nitrogen (N) and phosphorus (P) in plant and soil pools along two tundra-forest transects on Kola Peninsula, Russia, with a documented elevation shift of birch-dominated treeline by 70 m during the last 50 years. Results show that although total N and P stocks in the soil-plant system did not change with elevation, their distribution was significantly altered. With the transition from high-elevation tundra to low-elevation forest, P stocks in stones decreased, possibly reflecting enhanced weathering. In contrast, N and P stocks in plant biomass approximately tripled and available P and N in the soil increased fivefold toward the forest. This was paralleled by decreasing carbon (C)-to-nutrient ratios in foliage and litter, smaller C:N:P ratios in microbial biomass, and lower enzymatic activities related to N and P acquisition in forest soils. An incubation experiment further demonstrated manifold higher N and P net mineralization rates in litter and soil in forest compared to tundra, likely due to smaller C:N:P ratios in decomposing organic matter. Overall, our results show that forest expansion increases the mobilization of available nutrients through enhanced weathering and positive plant-soil feedback, with nutrient-rich forest litter releasing greater amounts of N and P upon decomposition. While the low N and P availability in tundra may retard treeline advances, its improvement toward the forest likely promotes tree growth and forest development.

Acclimation of subarctic vegetation to warming and increased cloudiness.

Subarctic ecosystems are exposed to elevated temperatures and increased cloudiness in a changing climate with potentially important effects on vegetation structure, composition, and ecosystem functioning. We investigated the individual and combined effects of warming and increased cloudiness on vegetation greenness and cover in mesocosms from two tundra and one palsa mire ecosystems kept under strict environmental control in climate chambers. We also investigated leaf anatomical and biochemical traits of four dominant vascular plant species (Empetrum hermaphroditum, Vaccinium myrtillus, Vaccinium vitis-idaea, and Rubus chamaemorus). Vegetation greenness increased in response to warming in all sites and in response to increased cloudiness in the tundra sites but without associated increases in vegetation cover or biomass, except that E. hermaphroditum biomass increased under warming. The combined warming and increased cloudiness treatment had an additive effect on vegetation greenness in all sites. It also increased the cover of graminoids and forbs in one of the tundra sites. Warming increased leaf dry mass per area of V. myrtillus and R. chamaemorus, and glandular trichome density of V. myrtillus and decreased spongy intercellular space of E. hermaphroditum and V. vitis-idaea. Increased cloudiness decreased leaf dry mass per area of V. myrtillus, palisade thickness of E. hermaphroditum, and stomata density of E. hermaphroditum and V. vitis-idaea, and increased leaf area and epidermis thickness of V. myrtillus, leaf shape index and nitrogen of E. hermaphroditum, and palisade intercellular space of V. vitis-idaea. The combined treatment caused thinner leaves and decreased leaf carbon for V. myrtillus, and increased leaf chlorophyll of E. hermaphroditum. We show that under future warmer increased cloudiness conditions in the Subarctic (as simulated in our experiment), vegetation composition and distribution will change, mostly dominated by graminoids and forbs. These changes will depend on the responses of leaf anatomical and biochemical traits and will likely impact carbon gain and primary productivity and abiotic and biotic stress tolerance.

Taking the beat of the Arctic: are lemming population cycles changing due to winter climate?

Reports of fading vole and lemming population cycles and persisting low populations in some parts of the Arctic have raised concerns about the spread of these fundamental changes to tundra food web dynamics. By compiling 24 unique time series of lemming population fluctuations across the circumpolar region, we show that virtually all populations displayed alternating periods of cyclic/non-cyclic fluctuations over the past four decades. Cyclic patterns were detected 55% of the time (n = 649 years pooled across sites) with a median periodicity of 3.7 years, and non-cyclic periods were not more frequent in recent years. Overall, there was an indication for a negative effect of warm spells occurring during the snow onset period of the preceding year on lemming abundance. However, winter duration or early winter climatic conditions did not differ on average between cyclic and non-cyclic periods. Analysis of the time series shows that there is presently no Arctic-wide collapse of lemming cycles, even though cycles have been sporadic at most sites during the last decades. Although non-stationary dynamics appears a common feature of lemming populations also in the past, continued warming in early winter may decrease the frequency of periodic irruptions with negative consequences for tundra ecosystems.