Arctic plant growth is predominantly nitrogen (N) limited. This limitation is generally attributed to slow soil microbial processes due to low temperatures. Here, we show that arctic plant-soil N cycling is also substantially constrained by the lack of larger detritivores (earthworms) able to mineralize and physically translocate litter and soil organic matter. These new functions provided by earthworms increased shrub and grass N concentration in our common garden experiment. Earthworm activity also increased either the height or number of floral shoots, while enhancing fine root production and vegetation greenness in heath and meadow communities to a level that exceeded the inherent differences between these two common arctic plant communities. Moreover, these worming effects on plant N and greening exceeded reported effects of warming, herbivory and nutrient addition, suggesting that human spreading of earthworms may lead to substantial changes in the structure and function of arctic ecosystems. Arctic plant growth is predominantly nitrogen limited, where the slow nitrogen turnover in the soil is commonly attributed to the cold arctic climate. Here the authors show that the arctic plant-soil nitrogen cycling is also constrained by the lack of larger detritivores like earthworms.

Permafrost thaw resulting from climate warming is threatening to release carbon from high latitude peatlands. The aim of this research was to determine subsidence rates linked to permafrost thaw in sub-Arctic peatlands in Sweden using historical orthophotographic (orthophotos), Unoccupied Aerial Vehicle (UAV), and Interferometric Synthetic Aperture Radar (InSAR) data. The orthophotos showed that the permafrost palsa on the study sites have been contracting in their areal extent, with the greatest rates of loss between 2002 and 2008. The surface motion estimated from differential digital elevation models from the UAV data showed high levels of subsidence (maxi-mum of −25 cm between 2017 and 2020) around the edges of the raised palsa plateaus. The InSAR data analysis showed that raised palsa areas had the greatest subsidence rates, with maximum subsidence rates of 1.5 cm between 2017 and 2020; however, all wetland vegetation types showed sub-sidence. We suggest that the difference in spatial units associated with each sensor explains parts of the variation in the subsidence levels recorded. We conclude that InSAR was able to identify the areas most at risk of subsidence and that it can be used to investigate subsidence over large spatial extents, whereas UAV data can be used to better understand the dynamics of permafrost degradation at a local level. These findings underpin a monitoring approach for these peatlands.

Plant-associated fungi have elementary roles in ecosystem productivity. There is little information on the interactions between arbuscular mycorrhizal (AM) fungal symbiosis, fine endophytic (FE) and dark septate endophytic (DSE) fungi, and their host plants in cold climate systems. In particular, the environmental filters potentially driving the relative abundance of these root symbionts remain unknown. We investigated the interlinkage of plant and belowground fungal responses to simulated herbivory (clipping, fertilization, and trampling) in a subarctic meadow system. AM and FE frequency in the two target plant roots, Potentilla crantzii and Saussurea alpina, was unaffected by simulated herbivory, highlighting the importance and resilience of arbuscule forming mycorrhizas in a range of environmental conditions. Fertilization and trampling increased DSE colonization in P. crantzii roots although generally P. crantzii performance was reduced in these plots. The idiosyncratic responses by DSE fungal frequency in the two host plants in our experiment indicate that the host plant identity has a pivotal role in the DSE fungus–plant outcome. DSE fungal frequency did not respond to environmental manipulations in a manner similar to arbuscular mycorrhizas, suggesting that they have a different role in plant ecology.

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.

Large and small mammalian herbivores are present in most vegetated areas in the Arctic and often have large impacts on plant community composition and ecosystem functioning. The relative importance of different herbivores and especially how their specific impact on the vegetation varies across the Arctic is however poorly understood. Here, we investigate how large and small herbivores influence vegetation density and plant community composition in four arctic vegetation types in Scandinavia and Alaska. We used a unique set of exclosures, excluding only large (reindeer and muskoxen) or all mammalian herbivores (also voles and lemmings) for at least 20 years. We found that mammalian herbivores in general decreased leaf area index, NDVI, and abundance of vascular plants in all four locations, even though the strength of the effect and which herbivore type caused these effects differed across locations. In three locations, herbivore presence caused contrasting plant communities, but not in the location with lowest productivity. Large herbivores had a negative effect on plant height, whereas small mammalian herbivores increased species diversity by decreasing dominance of the initially dominating plant species. Above- or belowground disturbances caused by herbivores were found to play an important role in shaping the vegetation in all locations. Synthesis: Based on these results, we conclude that both small and large mammalian herbivores influence vegetation in Scandinavia and Alaska in a similar way, some of which can mitigate effects of climate change. We also see important differences across locations, but these depend rather on local herbivore and plant community composition than large biogeographical differences among continents.

In arctic ecosystems, climate change has increased plant productivity. As arctic carbon (C) stocks predominantly are located belowground, the effects of greater plant productivity on soil C storage will significantly determine the net sink/source potential of these ecosystems, but vegetation controls on soil CO2 efflux remain poorly resolved.

In order to identify the role of canopy‐forming species in belowground C dynamics, we conducted a girdling experiment with plots distributed across 1 km2 of treeline birch (Betula pubescens ) forest and willow (Salix lapponum ) patches in northern Sweden and quantified the contribution of canopy vegetation to soil CO2 fluxes and belowground productivity.

Girdling birches reduced total soil CO2 efflux in the peak growing season by 53%, which is double the expected amount, given that trees contribute only half of the total leaf area in the forest. Root and mycorrhizal mycelial production also decreased substantially. At peak season, willow shrubs contributed 38% to soil CO2 efflux in their patches.

Our findings indicate that C, recently fixed by trees and tall shrubs, makes a substantial contribution to soil respiration. It is critically important that these processes are taken into consideration in the context of a greening arctic because productivity and ecosystem C sequestration are not synonymous.

In the face of climate change, it is important to estimate changes in key ecosystem properties such as plant biomass and gross primary productivity (GPP). Ground truth estimates and especially experiments are performed at small spatial scales (0.01-1 m(2)) and scaled up using coarse scale satellite remote sensing products. This will lead to a scaling bias for non-linearly related properties in heterogeneous environments when the relationships are not developed at the same spatial scale as the remote sensing products. We show that unmanned aerial vehicles (UAVs) can reliably measure normalized difference vegetation index (NDVI) at centimeter resolution even in highly heterogeneous Arctic tundra terrain. This reveals that this scaling bias increases most at very fine resolution, but UAVs can overcome this by generating remote sensing products at the same scales as ecological changes occur. Using ground truth data generated at 0.0625 m(2)and 1 m(2)with Landsat 30 m scale satellite imagery the resulting underestimation is large (8.9%-17.0% for biomass and 5.0%-9.7% for GPP(600)) and of a magnitude comparable to the expected effects of decades of climate change. Methods to correct this upscaling bias exist but rely on sub-pixel information. Our data shows that this scale-dependency will vary strongly between areas and across seasons, making it hard to derive generalized functions compensating for it. This is particularly relevant to Arctic greening with a predominantly heterogeneous land cover, strong seasonality and much experimental research at sub-meter scale, but also applies to other heterogeneous landscapes. These results demonstrate the value of UAVs for satellite validation. UAVs can bridge between plot scale used in ecological field investigations and coarse scale in satellite monitoring relevant for Earth System Models. Since future climate changes are expected to alter landscape heterogeneity, seasonally updated UAV imagery will be an essential tool to correctly predict landscape-scale changes in ecosystem properties.

Understanding how herbivores shape plant biomass and distribution is a core challenge in ecology. Yet, the lack of suitable remote sensing technology limits our knowledge of temporal and spatial impacts of mammal herbivores in the Earth system. The regular interannual density fluctuations of voles and lemmings are exceptional with their large reduction of plant biomass in Arctic landscapes during peak years (12–24%) as previously shown at large spatial scales using satellites. This provides evidence that herbivores are important drivers of observed global changes in vegetation productivity. Here, we use a novel approach with repeated unmanned aerial vehicle (UAV) flights, to map vegetation impact by rodents, indicating that many important aspects of vegetation dynamics otherwise hidden by the coarse resolution of satellite images, including plant–herbivore interactions, can be revealed using UAVs. We quantify areas impacted by rodents at four complex Arctic landscapes with very high spatial resolution UAV imagery to get a new perspective on how herbivores shape Arctic ecosystems. The area impacted by voles and lemmings is indeed substantial, larger at higher altitude tundra environments, varies between habitats depending on local snow cover and plant community composition, and is heterogeneous even within habitats at submeter scales. Coupling this with spectral reflectance of vegetation (NDVI), we can show that the impact on central ecosystem properties like GPP and biomass is stronger than currently accounted for in Arctic ecosystems. As an emerging technology, UAVs will allow us to better disentangle important information on how herbivores maintain spatial heterogeneity, function and diversity in natural ecosystems.

Rising temperatures can influence ecosystem processes both directly and indirectly, through effects on plant species and communities. An improved understanding of direct versus indirect effects of warming on ecosystem processes is needed for robust predictions of the impacts of climate change on terrestrial ecosystem carbon (C) dynamics.To explore potential direct and indirect effects of warming on C dynamics in arctic tundra heath, we established a warming (open top chambers) and dominant plant species (Empetrum hermaphroditum Hagerup) removal experiment at a high and low elevation site. We measured the individual and interactive effects of warming, dominant species removal and elevation on plant species cover, the normalized difference vegetation index (NDVI), leaf area index (LAI), temperature, soil moisture and instantaneous net ecosystem CO2 exchange.We hypothesized that ecosystems would be stronger CO2 sinks at the low elevation site, and that warming and species removal would weaken the CO2 sink because warming should increase ecosystem respiration (ER) and species removal should reduce gross primary productivity (GPP). Furthermore, we hypothesized that warming and species removal would have the greatest impact on processes at the high elevation where site temperature should be most limiting and dominant species may buffer the overall community to environmental stress more compared to the low elevation site where plants are more likely to compete with the dominant species.The instantaneous CO2 flux, which reflected a weak CO2 sink, was similar at both elevations. Neither experimental warming nor dominant species removal significantly changed GPP or instantaneous net ecosystem CO2 exchange even though species removal significantly reduced ER, NDVI and LAI.Our results show that even the loss of dominant plant species may not result in significant landscape‐scale responses of net ecosystem CO2 exchange to warming. They also show that NDVI and LAI may be limited in their ability to predict changes in GPP in these tundra heaths systems. Our study highlights the need for more detailed vegetation analyses and ground‐truthed measurements in order to accurately predict direct and indirect impacts of climatic change on ecosystem C dynamics.

The majority of variation in six traits critical to the growth, survival and reproduction of plant species is thought to be organised along just two dimensions, corresponding to strategies of plant size and resource acquisition. However, it is unknown whether global plant trait relationships extend to climatic extremes, and if these interspecific relationships are confounded by trait variation within species. We test whether trait relationships extend to the cold extremes of life on Earth using the largest database of tundra plant traits yet compiled. We show that tundra plants demonstrate remarkably similar resource economic traits, but not size traits, compared to global distributions, and exhibit the same two dimensions of trait variation. Three quarters of trait variation occurs among species, mirroring global estimates of interspecific trait variation. Plant trait relationships are thus generalizable to the edge of global trait-space, informing prediction of plant community change in a warming world.