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Spatial variation in plant chemical defence towards herbivores can help us understand variation in herbivore top–down control of shrubs in the Arctic and possibly also shrub responses to global warming. Less defended, non-resinous shrubs could be more influenced by herbivores than more defended, resinous shrubs. However, sparse field measurements limit our current understanding of how much of the circum-Arctic variation in defence compounds is explained by taxa or defence functional groups (resinous/non-resinous). We measured circum-Arctic chemical defence and leaf digestibility in resinous (Betula glandulosa, B. nana ssp. exilis) and non-resinous (B. nana ssp. nana, B. pumila) shrub birches to see how they vary among and within taxa and functional groups. Using liquid chromatography–mass spectrometry (LC–MS) metabolomic analyses and in vitro leaf digestibility via incubation in cattle rumen fluid, we analysed defence composition and leaf digestibility in 128 samples from 44 tundra locations.

We found biogeographical patterns in anti-herbivore defence where mean leaf triterpene concentrations and twig resin gland density were greater in resinous taxa and mean concentrations of condensing tannins were greater in non-resinous taxa. This indicates a biome-wide trade-off between triterpene- or tannin-dominated defences. However, we also found variations in chemical defence composition and resin gland density both within and among functional groups (resinous/non-resinous) and taxa, suggesting these categorisations only partly predict chemical herbivore defence. Complex tannins were the only defence compounds negatively related to in vitro digestibility, identifying this previously neglected tannin group as having a potential key role in birch anti-herbivore defence.

We conclude that circum-Arctic variation in birch anti-herbivore defence can be partly derived from biogeographical distributions of birch taxa, although our detailed mapping of plant defence provides more information on this variation and can be used for better predictions of herbivore effects on Arctic vegetation.

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.

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.

Large herbivores influence plant community structure and ecosystem processes in many ecosystems. In large parts of the Arctic, reindeer (or caribou) are the only large herbivores present. Recent studies show that reindeer have the potential to mitigate recent warming‐induced shrub encroachment in the Arctic and the associated greening of high‐latitude ecosystems. This will potentially have large scale consequences for ecosystem productivity and carbon cycling.

To date, information on variation in the interactions between reindeer and plants across Arctic landscapes has been scarce. We utilized a network of experimental sites across a latitudinal gradient in the Scandinavian mountains where reindeer have been excluded from 59 study plots for at least 15 years. We used this study system to test the effect of long‐term exclusion of reindeer on the abundance of major plant functional groups, the greenness indexes Leaf Area Index (LAI) and Normalized Difference Vegetation Index (NDVI), soil mineral nitrogen (N) and phosphorous (P), and species richness, and to determine whether the effect of reindeer exclusion is dependent on reindeer density, productivity, soil fertility or climate.

We found that NDVI and LAI, lichen and deciduous shrub abundances were largely reduced while soil mineral N was enhanced by reindeer. The direction and amplitude of other plant functional group responses to reindeer exclusion differed between forest and tundra as well as shrub‐ and herbaceous‐dominated vegetation. Higher reindeer densities were related to decreased plant species richness in low‐productive sites and to increased species richness in productive sites.

The relative reduction in LAI and associated absolute reductions of deciduous shrubs in response to reindeer were positively related to reindeer density, while the relative reduction in NDVI was not. Further, relative reductions in LAI and NDVI in response to reindeer were unrelated to climate and soil fertility.

Synthesis. Our results provide long‐term experimental evidence highlighting the role of reindeer density in regulating plant species richness, global climate change induced greenness patterns and shrub encroachment at regional scales in the Arctic. These findings emphasize the need to consider reindeer in models predicting vegetation patterns and changes in high‐latitude ecosystems.

Trophic interactions within food webs affect species distributions, coexistence, and provision of ecosystem services but can be strongly impacted by climatic changes. Understanding these impacts is therefore essential for managing ecosystems and sustaining human well-being. Here, we conducted a global synthesis of terrestrial, marine, and freshwater studies to identify key gaps in our knowledge of climate change impacts on food webs and determine whether the areas currently studied are those most likely to be impacted by climate change. We found research suffers from a strong geographic bias, with only 3.5% of studies occurring in the tropics. Importantly, the distribution of sites sampled under projected climate changes was biased-areas with decreases or large increases in precipitation and areas with low magnitudes of temperature change were under-represented. Our results suggest that understanding of climate change impacts on food webs could be broadened by considering more than two trophic levels, responses in addition to species abundance and biomass, impacts of a wider suite of climatic variables, and tropical ecosystems. Most importantly, to enable better forecasts of biodiversity responses to dimate change, we identify critically under-represented geographic regions and climatic conditions which should be prioritized in future research.

Climate change is affecting the species composition and functioning of Arctic and sub-Arctic plant and soil communities. Here we studied patterns in soil microarthropod (springtails and mites) communities across a gradient of increasing elevation that spanned 450 m, across which mean temperature declined by approximately 2.5 degrees C, in sub-Arctic Sweden. Across this gradient we characterized microarthropod communities in each of two types of vegetation, i.e., heath and meadow, to determine whether their responses to declining temperature differed with vegetation type. Mite abundance declined with increasing elevation, while springtail abundance showed the opposite response. Springtail communities were dominated by larger species at higher elevation. Mite abundance was unaffected by vegetation type, while springtail abundance was 53% higher in the heath than meadow vegetation across the gradient. Springtails but not mites responded differently to elevation in heath and meadow vegetation; hemi-edaphic species dominated in the heath at higher elevation while epiedaphic species dominated in the meadow. Our results suggest that sub-Arctic mite and springtail communities will likely respond in contrasting ways to changes in vegetation and soil properties resulting from climate warming.

Effective societal responses to rapid climate change in the Arctic rely on an accurate representation of region-specific ecosystem properties and processes. However, this is limited by the scarcity and patchy distribution of field measurements. Here, we use a comprehensive, geo-referenced database of primary field measurements in 1,840 published studies across the Arctic to identify statistically significant spatial biases in field sampling and study citation across this globally important region. We find that 31% of all study citations are derived from sites located within 50 km of just two research sites: Toolik Lake in the USA and Abisko in Sweden. Furthermore, relatively colder, more rapidly warming and sparsely vegetated sites are under-sampled and under-recognized in terms of citations, particularly among microbiology-related studies. The poorly sampled and cited areas, mainly in the Canadian high-Arctic archipelago and the Arctic coastline of Russia, constitute a large fraction of the Arctic ice-free land area. Our results suggest that the current pattern of sampling and citation may bias the scientific consensuses that underpin attempts to accurately predict and effectively mitigate climate change in the region. Further work is required to increase both the quality and quantity of sampling, and incorporate existing literature from poorly cited areas to generate a more representative picture of Arctic climate change and its environmental impacts.

Questions: Roots represent a considerable proportion of biomass, primary production and litter input in arctic tundra, and plant allocation of biomass to above- or below-ground tissue in response to climate change is a key factor in the future C balance of these ecosystems. According to optimality theory plants allocate C to the above- or below-ground structure that captures the most limiting resource. We used an elevational gradient to test this theory and as a space-for-time substitution to inform on tundra carbon allocation patterns under a shifting climate, by exploring if increasing elevation was positively related to the root:shoot ratio, as well as a larger plant allocation to adsorptive over storage roots.

Location: Arctic tundra heath dominated by Empetrum hermaphroditum close to Abisko, Sweden.

Methods: We measured root:shoot and fine:coarse root ratios of the plant communities along an elevational gradient by sampling above- and below-ground biomass, further separating root biomass into fine (<1 mm) and coarse roots.

Results: Plant biomass was higher at the lower elevations, but the root:shoot ratio did not vary with elevation. Resource allocation to fine relative to coarse roots increased with elevation, resulting in a fine:coarse root ratio that more than doubled with increasing elevation.

Conclusions: Contrary to previous works, the root:shoot ratio along this elevational gradient remained stable. However, communities along our study system were dominated by the same species at each elevation, which suggests that when changes in the root:shoot ratio occur with elevation these changes may be driven by differences in allocation patterns among species and thus turnover in plant community structure. Our results further reveal that the allocation of biomass to fine relative to coarse roots can differ between locations along an elevational gradient, even when overall above- vs below-ground biomass allocation does not. Given the functionally different roles of fine vs coarse roots this could have large implications for below-ground C cycling. Our results highlight the importance of direct effects vs indirect effects (such as changes in plant community composition and nutrient availability) of climate change for future C allocation above and below ground.

The above mentioned article was originally scheduled for publication in the special issue on Ecology of Tundra Arthropods with guest editors Toke T. Hoye . Lauren E. Culler. Erroneously, the article was published in Polar Biology, Volume 40, Issue 11, November, 2017. The publisher sincerely apologizes to the guest editors and the authors for the inconvenience caused.

Shrubs commonly form islands of fertility and are expanding their distribution and dominance in the arctic due to climate change, yet how soil properties may be influenced when different species of shrubs expand under warmer climates remains less explored. Important plant traits, such as their associated root community, are linked to functionally different and dominant shrub species in the arctic and these traits likely shape biogeochemical cycling in areas of shrub expansion. Using an elevational gradient as a proxy for warming, we explored how biochemical processes beneath two important arctic shrubs varied under warmer (low elevation) and cooler (high elevation) climates. Interestingly, the influence of elevation on biogeochemistry varied between the two shrubs. At the low elevation, Betula nana L., an ectomycorrhizal shrub, had high carbon (C) degrading enzyme activities, and relatively low potential net nitrogen (N) mineralization rates. Conversely, Empetrum nigrum ssp. hermaphroditum Hagerup, an cricoid mycorrhizal dwarf-shrub, had higher enzyme activities and net N immobilization rates at the higher elevation. Further, E. nigrum ssp. hermpahroditum appeared to have a more closed C and nutrient cycle than B. nana-enzymes degrading C, N, and phosphorus were tightly correlated with each other and with total C and ammonium concentrations in the humus beneath E. nigrum ssp. hermaphroditum, but not beneath B. nana. Our results suggest differences in the warming responses of C and N cycling beneath shrub species across an arctic tundra landscape.