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The phenology of vegetation, particularly the length of the growing season ( LOS; i.e., the period from greenup to senescence), is highly sensitive to climate change, which could imply potent feedbacks to the climate system, for example, by altering the ecosystem carbon (C) balance. In recent decades, the largest extensions of LOS have been reported at high northern latitudes, but further warming-induced LOS extensions may be constrained by too short photoperiod or unfulfilled chilling requirements. Here, we studied subarctic grasslands, which cover a vast area and contain large C stocks, but for which LOS changes under further warming are highly uncertain. We measured LOS extensions of Icelandic subarctic grasslands along natural geothermal soil warming gradients of different age (short term, where the measurements started after 5 years of warming and long term, i.e., warmed since ≥50 years) using ground-level measurements of normalized difference vegetation index. We found that LOS linearly extended with on average 2.1 days per °C soil warming up to the highest soil warming levels (ca. +10°C) and that LOS had the potential to extend at least 1 month. This indicates that the warming impact on LOS in these subarctic grasslands will likely not saturate in the near future. A similar response to short- and long-term warming indicated a strong physiological control of the phenological response of the subarctic grasslands to warming and suggested that genetic adaptations and community changes were likely of minor importance. We conclude that the warming-driven extension of the LOSs of these subarctic grasslands did not saturate up to +10°C warming, and hence that growing seasons of high-latitude grasslands are likely to continue lengthening with future warming (unless genetic adaptations or species shifts do occur). This persistence of the warming-induced extension of LOS has important implications for the C-sink potential of subarctic grasslands under climate change. [ABSTRACT FROM AUTHOR]

About 10% of the anthropogenic CO emissions have been absorbed by northern terrestrial ecosystems during the past decades. It has been hypothesized that part of this increasing carbon (C) sink is caused by the alleviation of nitrogen (N) limitation by increasing anthropogenic N inputs. However, little is known about this N-dependent C sink. Here, we studied the effect of chronic seabird-derived N inputs (47-67 kg N ha year) on the net soil organic C (SOC) storage rate of unmanaged Icelandic grasslands on the volcanic Vestmannaeyjar archipelago by using a stock change approach in combination with soil dating. We studied both early developmental (young) soils that had been receiving increased N inputs over a decadal timescale since an eruption in 1963, and well-developed soils, that had been receiving N inputs over a millennial timescale. For the latter, however, the effects on both decadal (topsoil; 40 years) and millennial (total soil profile; 1600 years) SOC storage could be studied, as the age of topsoil and the total soil profile could be determined from volcanic ash layers deposited in 1973 and 395 AD. We found that enhanced N availability-either from accumulation over time, or seabird derived-increased the net SOC storage rate. Under low N inputs, early developmental soils were weak decadal C sinks (0.018 ton SOC ha year), but this increased quickly under ca. 30 years of elevated N inputs to 0.29 ton SOC ha year, thereby equalling the decadal SOC storage rate of the unfertilized well-developed soils. Furthermore, for the well-developed soils, chronically elevated N inputs not only stimulated the decadal SOC storage rate in the topsoil, but also the total millennial SOC storage was consistently higher. Hence, our study suggests that Icelandic grasslands, if not disturbed, can remain C sinks for many centuries under current climatic conditions and that chronically elevated N inputs can induce a permanent strengthening of this sink. [ABSTRACT FROM AUTHOR]

Most manipulation experiments simulating global change in tundra were short-term or did not measure plant growth directly. Here, we assessed the growth of three shrubs (Cassiope tetragona, Empetrum hermaphroditum and Betula nana) at a subarctic heath in Abisko (Northern Sweden) after 22 years of warming (passive greenhouses), fertilisation (nutrients addition) and shading (hessian fabric), and compare this to observations from the first decade of treatment. We assessed the growth rate of current-year leaves and apical stem (primary growth) and cambial growth (secondary growth), and integrated growth rates with morphological measurements and species coverage. Primary- and total growth of Cassiope and Empetrum were unaffected by manipulations, whereas growth was substantially reduced under fertilisation and shading (but not warming) for Betula. Overall, shrub height and length tended to increase under fertilisation and warming, whereas branching increased mostly in shaded Cassiope. Morphological changes were coupled to increased secondary growth under fertilisation. The species coverage showed a remarkable increase in graminoids in fertilised plots. Shrub response to fertilisation was positive in the short-term but changed over time, likely because of an increased competition with graminoids. More erected postures and large, canopies (requiring enhanced secondary growth for stem reinforcement) likely compensated for the increased light competition in Empetrum and Cassiope but did not avoid growth reduction in the shade intolerant Betula. The impact of warming and shading on shrub growth was more conservative. The lack of growth enhancement under warming suggests the absence of long-term acclimation for processes limiting biomass production. The lack of negative effects of shading on Cassiope was linked to morphological changes increasing the photosynthetic surface. Overall, tundra shrubs showed developmental plasticity over the longer term. However, such plasticity was associated clearly with growth rate trends only in fertilised plots. [ABSTRACT FROM AUTHOR]

Summary: Below and aboveground vegetation dynamics are crucial in understanding how climate warming may affect terrestrial ecosystem carbon cycling. In contrast to aboveground biomass, the response of belowground biomass to long‐term warming has been poorly studied.Here, we characterized the impacts of decadal geothermal warming at two levels (on average +3.3°C and +7.9°C) on below and aboveground plant biomass stocks and production in a subarctic grassland.Soil warming did not change standing root biomass and even decreased fine root production and reduced aboveground biomass and production. Decadal soil warming also did not significantly alter the root–shoot ratio. The linear stepwise regression model suggested that following 10 yr of soil warming, temperature was no longer the direct driver of these responses, but losses of soil N were. Soil N losses, due to warming‐induced decreases in organic matter and water retention capacity, were identified as key driver of the decreased above and belowground production. The reduction in fine root production was accompanied by thinner roots with increased specific root area.These results indicate that after a decade of soil warming, plant productivity in the studied subarctic grassland was affected by soil warming mainly by the reduction in soil N. [ABSTRACT FROM AUTHOR]

Background and Aims The response of subarctic grassland's below-ground to soil warming is key to understanding this ecosystem's adaptation to future climate. Functionally different below-ground plant organs can respond differently to changes in soil temperature (T s). We aimed to understand the below-ground adaptation mechanisms by analysing the dynamics and chemistry of fine roots and rhizomes in relation to plant community composition and soil chemistry, along with the duration and magnitude of soil warming. Methods We investigated the effects of the duration [medium-term warming (MTW; 11 years) and long-term warming (LTW; > 60 years)] and magnitude (0–8.4 °C) of soil warming on below-ground plant biomass (BPB), fine root biomass (FRB) and rhizome biomass (RHB) in geothermally warmed subarctic grasslands. We evaluated the changes in BPB, FRB and RHB and the corresponding carbon (C) and nitrogen (N) pools in the context of ambient, T s < +2 °C and T s > +2 °C scenarios. Key Results BPB decreased exponentially in response to an increase in T s under MTW, whereas FRB declined under both MTW and LTW. The proportion of rhizomes increased and the C–N ratio in rhizomes decreased under LTW. The C and N pools in BPB in highly warmed plots under MTW were 50 % less than in the ambient plots, whereas under LTW, C and N pools in warmed plots were similar to those in non-warmed plots. Approximately 78 % of the variation in FRB, RHB, and C and N concentration and pools in fine roots and rhizomes was explained by the duration and magnitude of soil warming, soil chemistry, plant community functional composition, and above-ground biomass. Plant's below-ground biomass, chemistry and pools were related to a shift in the grassland's plant community composition – the abundance of ferns increased and BPB decreased towards higher T s under MTW, while the recovery of below-ground C and N pools under LTW was related to a higher plant diversity. Conclusion Our results indicate that plant community-level adaptation of below ground to soil warming occurs over long periods. We provide insight into the potential adaptation phases of subarctic grasslands. [ABSTRACT FROM AUTHOR]

Terrestrial carbon cycle feedbacks to global warming are major uncertainties in climate models. For in-depth understanding of changes in soil organic carbon ( SOC) after soil warming, long-term responses of SOC stabilization mechanisms such as aggregation, organo-mineral interactions and chemical recalcitrance need to be addressed. This study investigated the effect of 6 years of geothermal soil warming on different SOC fractions in an unmanaged grassland in Iceland. Along an extreme warming gradient of +0 to ~+40 °C, we isolated five fractions of SOC that varied conceptually in turnover rate from active to passive in the following order: particulate organic matter ( POM), dissolved organic carbon ( DOC), SOC in sand and stable aggregates ( SA), SOC in silt and clay ( SC- rSOC) and resistant SOC ( rSOC). Soil warming of 0.6 °C increased bulk SOC by 22 ± 43% (0-10 cm soil layer) and 27 ± 54% (20-30 cm), while further warming led to exponential SOC depletion of up to 79 ± 14% (0-10 cm) and 74 ± 8% (20-30) in the most warmed plots (~+40 °C). Only the SA fraction was more sensitive than the bulk soil, with 93 ± 6% (0-10 cm) and 86 ± 13% (20-30 cm) SOC losses and the highest relative enrichment in 13C as an indicator for the degree of decomposition (+1.6 ± 1.5‰ in 0-10 cm and +1.3 ± 0.8‰ in 20-30 cm). The SA fraction mass also declined along the warming gradient, while the SC fraction mass increased. This was explained by deactivation of aggregate-binding mechanisms. There was no difference between the responses of SC- rSOC (slow-cycling) and rSOC (passive) to warming, and 13C enrichment in rSOC was equal to that in bulk soil. We concluded that the sensitivity of SOC to warming was not a function of age or chemical recalcitrance, but triggered by changes in biophysical stabilization mechanisms, such as aggregation. [ABSTRACT FROM AUTHOR]

Global warming is predicted to impact high-latitude and high-altitude forests severely, jeopardizing their overall functioning and carbon storage, both of which depend on the warming response of tree fine root systems. This paper investigates the effect of soil warming on the biomass, morphology and colonizing ectomycorrhizal community of spruce fine and absorptive fine roots. We compare the responses of spruce roots growing at a man-made long-term soil warming (+ 4°C) experiment to results obtained from a geothermal soil temperature gradient (+ 1 to + 14°C) extending to the forest die-off edge, to shed light on the generalizability of the warming response and reveal any thresholds in acclimation ability. Trees in warmer soils formed longer and less-branched absorptive roots with higher specific root length and area, and lower root tissue density in both spruce stands, irrespective of warming method and location. Soil warming at the experimental warming site also supported the occurrence of a more varied EcM community and an increase in the abundance of Tomentella spp., indicating a shift in nutrient foraging. Fine and absorptive fine root biomass decreased toward warmer soil, with a sharp reduction occurring between + 4 and + 6°C from the ambient and leading to the collapse of the fine root system at the geothermal gradient. At the experimental warming site, the applied + 4°C warming had no effect on fine and absorptive fine root biomass. The similar fine root responses at the two warming sites suggest that the observations possibly reflect general acclimation patterns in spruce forests to global warming. [ABSTRACT FROM AUTHOR]

Abstract: Human activities have caused global changes of atmospheric chemistry resulting in increased temperature especially in the colder regions of the northern hemisphere. Since warming of the environment can have drastic effects on terrestrial ecosystems it is important to experimentally evaluate the extent of such effects in long‐term field‐based experiments. In this study we make use of both recent (short‐term) and long‐term geothermal warming of Icelandic soils to examine the responses of Collembola, an ecologically important group of soil invertebrates, to warming. On the basis of metabolic scaling theory, we hypothesized that species of small size would be more successful in warmed soils than species of larger size. Further we expected that top‐soil‐dwelling species would benefit more from warming than deep‐soil‐dwelling species. In order to test these hypotheses we sampled Collembola along replicated gradients of increasing temperature in areas that had been heated for about 6 years and more than 50 years respectively. Collembola were identified to species level, counted and the community‐weighted mean trait scores for six functional and ecological traits were calculated. Results show that both short‐term and long‐term soil warming caused a shift towards a higher relative abundance of species with small body size. Furthermore, abundance of top‐soil‐dwelling Collembola tended to increase after short‐term warming, but the opposite was observed after long‐term warming. Using trait‐based diversity indices (FRic and RaoQ), we show that functional richness and diversity of Collembola communities was significantly reduced (almost halved) as a result of short‐term soil warming to about 10°C above normal, but this effect was not detected in plots equally warmed for more than 50 years. This indicates that the functional diversity of Collembola communities have high resilience towards soil warming in a long‐term perspective. A plain language summary is available for this article. [ABSTRACT FROM AUTHOR]

Global change is expected to affect soil microbial communities through their responsiveness to temperature. It has been proposed that prolonged exposure to elevated temperatures may lead to progressively larger effects on soil microbial community composition. However, due to the relatively short-term nature of most warming experiments, this idea has been challenging to evaluate. The present study took the advantage of natural geothermal gradients (from +1°C to +19°C above ambient) in two subarctic grasslands to test the hypothesis that long-term exposure (>50 years) intensifies the effect of warming on microbial community composition compared to short-term exposure (5-7 years). Community profiles from amplicon sequencing of bacterial and fungal rRNA genes did not support this hypothesis: significant changes relative to ambient were observed only starting from the warming intensity of +9°C in the long termand +7°C/+3°C in the short term, for bacteria and fungi, respectively. Our results suggest that microbial communities in high-latitude grasslands will not undergo lasting shifts in community composition under the warming predicted for the coming 100 years (+2.2°C to +8.3°C). [ABSTRACT FROM AUTHOR]

Nitrous oxide (N 2 O) and methane (CH 4 ) emissions were measured along three natural geothermal soil temperature (T s ) gradients in freely drained upland soils in a volcanic area in Iceland. Two of the T s gradients (underneath a grassland (GN) and a forest site (FN), respectively) were recently formed (in May 2008) and thus subjected to relatively short-term warming. The third T s gradient, underneath another grassland site (GO), had been subjected to long-term soil warming (over at least 45 years). The N 2 O and CH 4 emissions were measured using the static chamber method. In addition, subsurface soil gas concentrations (5–20 cm) were studied. N 2 O emissions from GN were slightly higher than those from GO in the temperature elevation range up to +5 ° C, while CH 4 uptake rates were similar. Under moderate soil warming (<+5 ° C) there were no significant increases in gas flux rates within any of the sites, but when soil warming exceeded +20 ° C, both N 2 O and CH 4 emissions increased significantly at all sites. While net uptake of CH 4 (up to −0.15 mg CH 4 m −2 h −1 ) and occasional N 2 O uptake (up to −12 μg N 2 O m −2 h −1 ) were measured in the unwarmed plots at all sites, net emissions were only measured from the warmest plots (up to 2600 μg N 2 O m −2 h −1 and up to 1.3 mg CH 4 m −2 h −1 ). The subsurface soil N 2 O concentrations increased with soil warming, indicating enhanced N-turnover. Subsurface soil CH 4 concentrations initially decreased under moderate soil warming (up to +5 ° C), but above that threshold they also increased significantly. A portion of the N 2 O and CH 4 emitted from the warmest plots may, however, be geothermally derived, this should be further confirmed with isotope studies. In conclusion, our research suggests that moderate increases in soil temperature (up to +5 ° C) may not significantly increase N 2 O and CH 4 emissions at these upland soils, both in the short and longer term. However, warming trends exceeding +5 ° C as predicted for 2100 in pessimistic scenarios may cause increased trace gas emissions and thus significant positive feedbacks to climate change. [ABSTRACT FROM AUTHOR]

The effect of warming is stronger in arctic and sub-arctic latitudes than in temperate and tropical zones. We studied soil metabolomes along two soil-warming gradients (0 to +15 °C). One temperature gradient has been present for at least 50 years and possibly even centuries (long-term treatment), while the second gradient was created after a shallow crustal earthquake in 2008 (short-term treatment). Soil metabolomes at the two sites responded differently to warming. At the short-term warmed site, warming of ≤+3 °C already shifted soil metabolomic profiles relative to the controls, whereas at the long-term warmed site the soil metabolome only shifted at temperatures ≥+5 °C. Saccharides and amino acids, primary metabolites involved in protective mechanisms against heat, were the main compounds accumulated at the highest soil warming levels. Some secondary metabolites associated with a broad spectrum of stressors, like phenolic acids and terpenes, were also up-regulated. Across the IPCC scenario's, most climate models predict a substantial rise in mean annual temperature of up to 8 °C in the Arctic region by the end of the 21st century. Our results suggest that temperature increases of ≥+5 °C would permanently alter soil metabolomic profile, whereas smaller temperature increases of (≤+3 °C) would affect soil metabolome profile transiently, not permanently. • The time and intensity of warming exposure affect differently soil function. • Soil function could acclimatize to smaller temperature increases of ≤+3 °C. • Temperature increases of ≥+5 °C could permanently alter soil function. • Under long-term warming: ≥+5 °C, soil contained more metabolites against heat stress. [ABSTRACT FROM AUTHOR]

Although ongoing research has revealed some of the main drivers behind global spatial patterns of microbial communities, spatio-temporal dynamics of these communities still remain largely unexplored. Here, we investigate spatio-temporal variability of both bacterial and eukaryotic soil microbial communities at local and intercontinental scales. We compare how temporal variation in community composition scales with spatial variation in community composition, and explore the extent to which bacteria, protists, fungi and metazoa have similar patterns of temporal community dynamics. All soil microbial groups displayed a strong correlation between spatial distance and community dissimilarity, which was related to the ratio of organism to sample size. Temporal changes were variable, ranging from equal to local between-sample variation, to as large as that between communities several thousand kilometers apart. Moreover, significant correlations were found between bacterial and protist communities, as well as between protist and fungal communities, indicating that these microbial groups change in tandem, potentially driven by interactions between them. We conclude that temporal variation can be considerable in soil microbial communities, and that future studies need to consider temporal variation in order to reliably capture all drivers of soil microbiome changes. [ABSTRACT FROM AUTHOR]

• Temperature gradients in geothermal habitats serve as local models of global warming • We studied temperature-dependent traits in springtails from two geothermal habitats • Springtail traits and their plasticity varied within and between forest and grassland • Habitat properties may shape thermal phenotypes to an extent comparable with warming Common-garden experiments with populations sampled along natural thermal gradients help to reveal local adaptation, disentangle environmental and genetic effects, and ultimately predict, by analogy, future biotic responses to climate change. In this regard, geothermal habitats are useful model systems as they exhibit dramatic changes in soil temperature. The springtail Protaphorura pseudovanderdrifti has apparently coped with such local geothermal warming in Iceland, as this species occurs along a more than half-century-old geothermal gradient in a grassland and persists along a newly emerged temperature gradient in a previously non-geothermal planted spruce forest. We measured thermal reaction norms for development and walking speed and acute cold shock tolerance of P. pseudovanderdrifti originating from the grassland and forest geothermal gradients. Temperature-dependent juvenile development showed little variation among subpopulations from the recently warmed forest, probably due to insufficient evolutionary time, but springtails from the warmed grassland plots had significantly steeper reaction norms than their counterparts from the corresponding unwarmed plot. In contrast, cold tolerance and locomotory activity showed no conclusive clinal pattern despite significant within-habitat variation. There appeared to be significant differences between habitats, as springtails from the forest had more temperature-sensitive developmental rate and locomotory activity, walked faster, and exhibited more variable cold tolerance than grassland springtails did. The planting of a forest, therefore, seems to have exerted a stronger effect on the thermal phenotype of P. pseudovanderdrifti than the emergence of a geothermal gradient. Thus, habitat properties may be no less important in shaping thermal reaction norms than the mean temperature. These local-scale findings suggest that, in addition to warming per se, global transformation of communities may drive the evolution of thermal phenotypes to an extent comparable with the effect of rising environmental temperature. [ABSTRACT FROM AUTHOR]