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The initiation of tree growth in spring is typically linked to leaf-out. Bud dormancy drives budburst timing while dormancy of tree roots has largely remained unexplored, although the latter can shape below-ground growth and carbon dynamics. As roots experience different temperatures from buds, their dormancy dynamics and growth timing can differ and need to be studied, in order to better understand above- and below-ground growth responses to climate warming.We evaluated differences in dormancy dynamics between roots and buds in Fagus sylvatica and Populus nigra by quantifying the amount of warmth required to initiate above and below-ground growth from October to February. We furthermore carried out seven experiments, manipulating only the soil temperature prior to or during tree leaf-out to evaluate the potential of warmer roots to advance budburst timing. Soil temperature was manipulated via snow removal, heating buried wires, insulated pots, soil as well as chamber soil warming, using tree seedlings and adult trees of Fagus sylvatica as well as tree seedlings of Betula pendula.Root dormancy was virtually absent in comparison to the much deeper and variable bud dormancy, with roots being able to start growing immediately after being exposed to warm temperatures during the winter. Furthermore, warmer soil temperature advanced budburst in the meta-analysis of all soil temperature manipulation experiments. Therefore, differences in root and bud dormancy dynamics and their interaction likely explain the non-synchronized below and above-ground growth periods, both processes requiring separate predictions under climate warming.

Arctic soils store nearly half of the global soil organic carbon, but what will happen with this large carbon pool if soil macrofauna, able to ingest organic matter accumulated at depths in the soil, establish in the Arctic? The question is justified as emerging evidence suggests that low soil organic carbon (SOC) turnover rates in high latitude ecosystems could, in addition to abiotic factors, partly be due to the current lack of larger detritivores that can stimulate the breakdown of organic matter. Earthworms are large detritivores increasing their distribution into arctic ecosystems. With potential to both increase stabilisation and decomposition of SOC, but also enhance plant productivity, it has been difficult to determine what net effect earthworms have on ecosystem C storage. The scientific debate around this ‘earthworm dilemma’ has however primarily focused on the fate of SOC in response to earthworms, leaving cascading effects on plant productivity largely undiscussed.Here, we use a four-year outdoor experiment to study the effects of introducing earthworms to tundra vegetation types on both plant biomass (above and belowground) and SOC. We found that earthworms invasive to the Arctic: i) reduced the SOC pool beneath herb dominated tundra while they increased the SOC pool under dwarf-shrub dominated tundra; ii) increased the below ground biomass in both vegetation types; and iii) increased the total plant biomass C to the degree that it offset the SOC losses from the herb dominated soil. In the dwarf-shrub vegetation, earthworms increased both the plant C pool and the SOC pool resulting in a net increase of the ecosystem C stock. We highlight that the effect on root growth seems of great importance when predicting how ecosystem C sequestering responds to invasive earthworms. Both through increased plant biomass C but also through increased deposition of persistent root-derived organic matter.

The rewetting of formerly drained peatlands is a strategy to fight against global warming through the reduction of CO2 emissions, although this can lead to elevated CH4 emissions. The interplay between plants, hydrology and microbiomes as ultimate determinants of CH4 dynamics is still poorly understood, despite recent progress in field studies. Using a mesocosm approach, we simulated the re-cultivation of a degraded temperate fen with three different water levels and two different plant over the course of a growing season. Peat samples for microbiome analysis, above- and below-ground plant biomass and gas fluxes were measured in April, June, August and October. Microbiome composition in top and subsoils was determined using 16S rRNA gene amplicon sequencing. We found that peat depth and sampling time were the major factors shaping the microbiome composition dynamics. While plant species had a less strong impact, the difference to bare ground microbiomes was significant, especially in the lower layer. The water status also affected the microbiome, albeit to a much lesser extent. Methanogens were most abundant in the deeper peat and also more abundant in bare ground and Carex rostrata pots, as compared to Juncus inflexus or mixed pots. This was inversely linked to the larger root network size of J. inflexus. The methane emissions correlated positively with the abundance of methanogens and correlated negatively with the root network size. In conclusion, this interdisciplinary study sheds light on how the complex interplay between plants, hydrology and the fen microbiome affect CH4 emissions. It showed that the presence of plants as well as the plant functional type determine the abundance of methanogens and microbiome composition and thereby the resulting CH4 fluxes accordingly.

openaire: EC/H2020/819202/EU//SOS.aquaterra Funding Information: We thank the PrimeSCale team for laying the foundation of this research. We further thank Colleen Iversen for being up to date with root representation in models. BW acknowledges ERC StG PRIMETIME (grant no. 101039588). MK acknowledges European Research Council (ERC) under the European Union's Horizon 2020 research and innovation program (grant agreement no. 819202). Publisher Copyright: © 2023 The Authors. New Phytologist © 2023 New Phytologist Foundation. The distribution of roots throughout the soil drives depth-dependent plant–soil interactions and ecosystem processes, particularly in arctic tundra where plant biomass, is predominantly belowground. Vegetation is usually classified from aboveground, but it is unclear whether such classifications are suitable to estimate belowground attributes and their consequences, such as rooting depth distribution and its influence on carbon cycling. We performed a meta-analysis of 55 published arctic rooting depth profiles, testing for differences both between distributions based on aboveground vegetation types (Graminoid, Wetland, Erect-shrub, and Prostrate-shrub tundra) and between ‘Root Profile Types’ for which we defined three representative and contrasting clusters. We further analyzed potential impacts of these different rooting depth distributions on rhizosphere priming-induced carbon losses from tundra soils. Rooting depth distribution hardly differed between aboveground vegetation types but varied between Root Profile Types. Accordingly, modelled priming-induced carbon emissions were similar between aboveground vegetation types when they were applied to the entire tundra, but ranged from 7.2 to 17.6 Pg C cumulative emissions until 2100 between individual Root Profile Types. Variations in rooting depth distribution are important for the circumpolar tundra carbon-climate feedback but can currently not be inferred adequately from aboveground vegetation type classifications.

During litter decomposition, part of the water-soluble components of the material dissolve (leach) rapidly into available water in the environment. Studies on litter decomposition that quantify mass-loss from litterbags integrate leaching and mineralization. In contrast to Lind et al. (2022), we believe that correcting for leaching in (terrestrial) litterbags studies such as the Tea Bag Index will result in more uncertainties than it resolves. This is mainly because leaching is a continuous process and because leached material can still be mineralized after leaching. Further, amount of material that potentially leaches from tea is comparable to other litter types. When correcting for leaching, it is key to be specific about the employed method, just like being specific about the study specific definition of decomposition.

Northern soils play an important role in Earth’s climate system as they store large amounts of carbon that, if released, could strongly increase greenhouse gas levels in our atmosphere. Most research to date has focused on how the turnover of organic matter in these soils is regulated by abiotic factors and few studies have considered the potential role of biotic regulation. Here, we claim that soil organisms’ presence or absence is key to understanding and predicting future climate feedbacks from northern soils. We propose that the arrival of soil organisms with currently ‘missing traits’, i.e., properties that the present community does not have, can alleviate functional limitation and result in greatly enhanced decomposition rates, in parity with effects predicted due to increasing temperatures. We base this argument on a series of emerging evidence suggesting that the dispersal of until-then absent micro-, meso- and macro-organisms (i.e., microbes and invertebrate soil fauna) into new regions and newly-thawed soil layers can drastically affect soil functioning. These new observations make us question the current view that neglects organism driven ‘alleviation effects’ when predicting the future feedbacks between northern ecosystems and our planets’ climate. We therefore advocate for an updated framework in which soil biota and their traits become essential when predicting the fate of soil functions in warming northern ecosystems.

PurposeThe standardized ‘Tea Bag Index’ enables comparisons of litter decomposition rates, a key component of carbon cycling, across ecosystems. However, tea ‘litter’ may leach more than other plant litter, skewing comparisons of decomposition rates between sites with differing moisture conditions. Therefore, some researchers leach tea bags before field incubation. This decreases comparability between studies, and it is unclear if this modification is necessary.MethodsWe submerged green and rooibos tea bags in water, and measured their leaching losses over time (2 min – 72 h). We also compared leaching of tea to leaf and root litter from other plant species, and finally, compared mass loss of pre-leached and standard tea bags in a fully factorial incubation experiment differing in soil moisture (wet and dry) and soil types (sand and peat).ResultsBoth green and rooibos tea leached strongly, levelling-off at about 40% and 20% mass loss, respectively. Mass loss from leaching was highest in green tea followed by leaves of other plants, then rooibos tea, and finally roots of other plants. When incubated for 4 weeks, both teas showed lower mass loss when they had been pre-leached compared to standard tea bags. However, these differences between standard and pre-leached tea bags were similar in moist vs. dry soils, both in peat and in sand.ConclusionsThus, despite large leaching losses, we conclude that leaching tea bags before field or lab incubation is not necessary to compare decomposition rates between systems, ranging from as much as 5% to 25% soil moisture.

Understanding variation in root and shoot growth phenology among species is crucial to understanding underlying mechanisms of temporal niche differentiation. However, little is known about the relationship between root and shoot phenology, or how this relationship varies among functional traits. We examined fine root and shoot phenology of 42 seedlings representing a variety of woody species that inhabit the cool temperate forests of northern Japan. Some aspects of phenology were common to the pool of species examined: we found positive relationships between root and shoot phenology for the end of growth, and for the duration of growth, but not for the start of growth. Further, seedlings that started root growth relatively early also ended root growth relatively late. Other aspects of phenology varied predictably with functional traits, i.e. leaf habit and successional status: first, root growth in evergreen species started significantly earlier and ended later than in deciduous species; second, early successional species had the longest duration of shoot growth among all successional types. Our results suggest that niche differentiation may be promoted by differences in phenology between root and shoots, likely contributing to the co‐existence of woody seedlings in temperate forests.

1. Root phenology influences the timing of plant resource acquisition and carbon fluxes into the soil. This is particularly important in fen peatlands, in which peat is primarily formed by roots and rhizomes of vascular plants. However, most fens in Central Europe are drained for agriculture, leading to large carbon losses, and further threatened by increasing frequency and intensity of droughts. Rewetting fens aims to restore the original carbon sink, but how root phenology is affected by drainage and rewetting is largely unknown. 2. We monitored root phenology with minirhizotrons in drained and rewetted fens (alder forest, percolation fen and coastal fen) as well as its soil temperature and water table depth during the 2018 drought. For each fen type, we studied a drained site and a site that was rewetted ~25 years ago, while all the sites studied had been drained for almost a century. 3. Overall, the growing season was longer with rewetting, allowing roots to grow over a longer period in the year and have a higher root production than under drainage. With increasing depth, the growing season shifted to later in time but remained a similar length, and the relative importance of soil temperature for root length changes increased with soil depth. 4. Synthesis and applications. Rewetting extended the growing season of roots, highlighting the importance of phenology in explaining root productivity in peatlands. A longer growing season allows a longer period of carbon sequestration in form of root biomass and promotes the peatlands' carbon sink function, especially through longer growth in deep soil layers. Thus, management practices that focus on rewetting peatland ecosystems are necessary to maintain their function as carbon sinks, particularly under drought conditions, and are a top priority to reduce carbon emissions and address climate change.

Start and end of the growing season determine important ecosystem processes, but their drivers may differ above-and belowground, between autumn and spring, and between ecosystems. Here, we compare above-and belowground spring and autumn phenology, and their abiotic drivers (temperature, water level, and soil moisture) in four temperate ecosystems (beech forest, alder carr, phragmites reed, and sedge reed). Root growth was measured in-situ with minirhizotrons and compared with aboveground phenology assessed with dendrometer data and NDVI. Synchrony of above- and belowground phenology depended on ecosystem. Onset of root growth was later than shoot growth in all three peatlands (12–33 days), but similar in the beech forest. The growing season ended earlier belowground in the two forested ecosystems (beech forest: 27 days, understory of the alder carr: 55 days), but did not differ in the phragmites reed. Generally, root production was correlated with soil temperature (especially in spring) and water level in the peatlands, while abiotic factors were less correlated with leaf activity or root production in either spring or autumn in the beech forest. Root production on organic soils was ten times higher compared to the zonal broadleaf deciduous forest on mineral soils, highlighting the importance of peatlands. Belowground phenology cannot be projected from aboveground phenology and measuring root phenology is crucial to understand temporal dynamics of production and carbon fluxes.

Peatlands cover only 3% of the lands surface, but store roughly a third of the global soil carbon due to inhibited decomposition rates. Over a third of the peatland area in Europe are fens, in which the peat is primarily formed by roots and rhizomes of vascular plants. These fens have been subjected to widespread drainage and conversion into agricultural areas. As a result, they continuously emit large amount of greenhouse gases. One strategy of mitigating the emissions, and ideally restoring the original sink function, is to rewet fen peatlands. However, it remains uncertain how rewetting changes decomposition rates compared to the drained state, and what the underlying biogeochemical processes and organic matter transformations during litter decomposition and peat formation are. We here present decomposition rates of root material in different depth, over 6 months, a year, and two years in different drained and rewetted fen ecosystems (percolation fen, coastal fen, alder forest). In addition to mass loss, we also assessed the composition of carbon compounds over time.

Soils are warming as air temperatures rise across the Arctic and Boreal region concurrent with the expansion of tall-statured shrubs and trees in the tundra. Changes in vegetation structure and function are expected to alter soil thermal regimes, thereby modifying climate feedbacks related to permafrost thaw and carbon cycling. However, current understanding of vegetation impacts on soil temperature is limited to local or regional scales and lacks the generality necessary to predict soil warming and permafrost stability on a pan-Arctic scale. Here we synthesize shallow soil and air temperature observations with broad spatial and temporal coverage collected across 106 sites representing nine different vegetation types in the permafrost region. We showed ecosystems with tall-statured shrubs and trees (>40 cm) have warmer shallow soils than those with short-statured tundra vegetation when normalized to a constant air temperature. In tree and tall shrub vegetation types, cooler temperatures in the warm season do not lead to cooler mean annual soil temperature indicating that ground thermal regimes in the cold-season rather than the warm-season are most critical for predicting soil warming in ecosystems underlain by permafrost. Our results suggest that the expansion of tall shrubs and trees into tundra regions can amplify shallow soil warming, and could increase the potential for increased seasonal thaw depth and increase soil carbon cycling rates and lead to increased carbon dioxide loss and further permafrost thaw.

Background and AimsForest peatlands represent 25 % of global peatlands and store large amounts of carbon (C) as peat. Traditionally they have been drained in order to increase forestry yield, which may cause large losses of C from the peat. Rewetting aims to stop these losses and to restore the initial storage function of the peatlands. As roots represent major peat-forming elements in these systems, we sampled roots with diameter MethodsWe cored soil next to Alnus glutinosa stems and sorted root biomass into Key ResultsRoot biomass in the rewetted site was more than double that in the drained site. This difference was mostly driven by very fine roots ConclusionsThe size-dependent opposite patterns between root biomass and their functional characteristics under contrasting water regimes indicate differences between fine and coarse roots in their response to environmental changes. Root age distribution points to similar root turnover rates between the sites, while higher root biomass in the rewetted site clearly indicates larger tree C stocks below ground under rewetting, supporting the C sink function of the ecosystem.

Peatlands are effective carbon sinks as more biomass is produced than decomposed under the prevalent anoxic conditions. Draining peatlands coupled with warming releases stored carbon, and subsequent rewetting may or may not restore the original carbon sink. Yet, patterns of plant production and decomposition in rewetted peatlands and how they compare to drained conditions remain largely unexplored. Here, we measured annual above- and belowground biomass production and decomposition in three different drained and rewetted peatland types: alder forest, percolation fen and coastal fen during an exceptionally dry year. We also used standard plant material to compare decomposition between the sites, regardless of the decomposability of the local plant material. Rewetted sites showed higher root and shoot production in the percolation fen and higher root production in the coastal fen, but similar root and leaf production in the alder forest. Decomposition rates were generally similar in drained and rewetted sites, only in the percolation fen and alder forest did aboveground litter decompose faster in the drained sites. The rewetted percolation fen and the two coastal sites had the highest projected potential for organic matter accumulation. Roots accounted for 23–66% of total biomass production, and belowground biomass, rather than aboveground biomass, was particularly important for organic matter accumulation in the coastal fens. This highlights the significance of roots as main peat-forming element in these graminoid-dominated fen peatlands and their crucial role in carbon cycling, and shows that high biomass production supported the peatlands’ function as carbon sink even during a dry year.

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.

In their natural state peatlands are effective carbon sinks as more biomass is produced than decomposed under the prevalent anoxic conditions. Draining peatlands results in release of the stored carbon. Rewetting may or may not restore the original carbon sink. Patterns of plant production and decomposition in rewetted peatlands and how they compare to the drained state remain largely unexplored.We measured annual above- and belowground biomass production and decomposition in three different drained and rewetted peatland types: alder forest, percolation fen and coastal fen. We also used standard material (green and rooibos tea) to compare decomposition between the sites, regardless of the decomposability of the local plant material.Rewetted sites had higher root and shoot production in the percolation fen, and higher root production in the coastal fen but similar root and leaf production in the alder forest (excluding woody biomass). Decomposition rates were similar in drained and rewetted sites, only in the percolation fen and alder forest aboveground litter decomposed faster in the drained sites. The rewetted percolation fen and the two coastal sites have the highest projected potential for organic matter accumulation due to high production and low decomposition rates. Roots accounted for 23–66% of total biomass production, and the importance of belowground biomass, rather than aboveground biomass, for organic matter accumulation increased with time. This highlights the significance of roots as main peat forming element in these graminoid-dominated fen peatlands and their crucial role in carbon cycling. Notably, increased production compensated for loss by decomposition even during the exceptionally dry year 2018.Rewetted sites generally had a more productive plant community compared to drained sites, only tree stem biomass increment was higher in the drained alder forest site. High biomass production supported the peatlands’ function as carbon sink even during a dry year and roots were more important than shoots in establishing this sink, especially in the graminoid dominated sites. Rewetted peatlands may cope better with the extreme weather conditions that will occur more frequently in the future, emphasizing the case for rewetting those systems

Snow depth increases observed and predicted in the sub-arctic are of critical importance for the dynamics of lowland permafrost and vegetation. Snow acts as an insulator that protects vegetation but may lead to permafrost degradation. In the Abisko area, in northernmost Sweden, there has been an increasing trend in snow depth during the last Century. Downscaled climate scenarios predict an increase in precipitation by 1.5 - 2% per decade for the coming 60 years. The observed changes in snow cover have affected peat mires in this area as thawing of permafrost, increases in active layer thickness and associated vegetation changes have been reported during the last decades. An experimental manipulation was set up at one of these lowland permafrost sites in the Abisko area (68°20’48’’N, 18°58’16’’E) 15 years ago, to simulate projected future increases in winter precipitation and to study their effect on permafrost and vegetation. The snow cover has been more than twice as thick in manipulated plots compared to control plots and it has had a large impact on permafrost and vegetation. It resulted in statistically significant differences in mean winter and minimum ground temperatures between the control and the manipulated plots. Already after three years there was a statistically significant difference between active layer thickness in the manipulated plots compared to the control plots. In 2019, the active layer thickness in the control plots were around 70 cm whereas in the manipulated plots it was 110 cm. The increased active layer thickness has led to surface subsidence due to melting of ground ice in all the manipulated plots. The increased snow thickness has prolonged the duration of the snow cover in spring with up to 22 days. However, this loss in early season photosynthesis was well compensated for by the increased absorption of PAR and higher light use efficiency throughout the whole growing seasons in the manipulated plots. Eriophorum vaginatum is a species that has been especially favored in the manipulated plots. It has increased both in number and in size. Underneath the soil surface, the roots have also been affected. There has been a strong increase in total root length and growth in the active layer, and deep roots has invaded the newly thawed permafrost in the manipulated plots. The increased active layer thickness has also had an effect on the bacterial community composition in the newly thawed areas. According to past, century-long patterns of increasing snow depth and projections of continuing increases, it is very likely that the changes in permafrost and vegetation that have been demonstrated by this experimental treatment will occur in the future under natural conditions.

Of all terrestrial ecosystems, peatlands store carbon most effectively in long-term scales of millennia. However, many peatlands have been drained for peat extraction or agricultural use. This converts peatlands from sinks to sources of carbon, causing approx. 5% of the anthropogenic greenhouse effect and additional negative effects on other ecosystem services. Rewetting peatlands can mitigate climate change and may be combined with management in the form of paludiculture. Rewetted peatlands, however, do not equal their pristine ancestors and their ecological functioning is not understood. This holds true especially for groundwater-fed fens. Their functioning results from manifold interactions and can only be understood following an integrative approach of many relevant fields of science, which we merge in the interdisciplinary project WETSCAPES. Here, we address interactions among water transport and chemistry, primary production, peat formation, matter transformation and transport, microbial community, and greenhouse gas exchange using state of the art methods. We record data on six study sites spread across three common fen types (Alder forest, percolation fen, and coastal fen), each in drained and rewetted states. First results revealed that indicators reflecting more long-term effects like vegetation and soil chemistry showed a stronger differentiation between drained and rewetted states than variables with a more immediate reaction to environmental change, like greenhouse gas (GHG) emissions. Variations in microbial community composition explained differences in soil chemical data as well as vegetation composition and GHG exchange. We show the importance of developing an integrative understanding of managed fen peatlands and their ecosystem functioning.

Belowground plant structures are inherently difficult to observe in the field. Sedge peat that mainly consists of partly decayed roots and rhizomes offers a particularly challenging soil matrix to study (live) plant roots. To obtain information on belowground plant morphology, research commonly relies on rhizotrons, excavations, or computerized tomography scans (CT). However, all of these methods have certain limitations. For example, CT scans of peat cores cannot sharply distinguish between plant material and water, and rhizotrons do not provide a 3D structure of the root system. Here, we developed a low-cost approach for 3D visualization of the root system in peat monoliths. Two large diameter (20 cm) peat cores were extracted, frozen and two smaller peat monoliths (47 &times, 6.5 &times, 13 cm) were taken from each core. Slices of 0.5 mm or 1 mm were cut from one of the frozen monoliths, respectively, using a paper block cutter and the freshly cut surface of the monolith was photographed after each cut. A 3D model of the fresh (live) roots and rhizomes was reconstructed from the resulting images of the thinner slices based on computerized image analysis, including preprocessing, filtering, segmentation and 3D visualization using the open-source software Fiji, Drishti, and Ilastik. Digital volume measurements on the models produced similar data as manual washing out of roots from the adjacent peat monoliths. The constructed 3D models provide valuable insight into the three-dimensional structure of the root system in the peat matrix.

In the last decades, rewetting of drained peatlands is on the rise worldwide, to restore the significant carbon sink function. Rewetted peatlands differ substantially from their pristine counterparts and can, thus, be considered as novel ecosystems. Despite the increasing understanding of peat microbiomes, little is known about the seasonal dynamics and network interactions of the microbial communities in these novel ecosystems, especially in rewetted groundwater-fed peatlands, i.e. fens. Here, we investigated the seasonal dynamics in both prokaryotic and eukaryotic microbiomes in three common types of fens in Northern Germany, namely percolation fen, alder forest and coastal fen. The eukaryotic microbiomes, including fungi, protists and metazoa, showed significant changes of their community structures across the seasons in contrast to largely unaffected prokaryotic microbiomes. The co-occurrence network in the summer showed a distinct topology compared to networks in the other seasons, which was driven by the increased connections among protists, as well as between protists and the other microbial groups. Our results also indicated that the dynamics in eukaryotic microbiomes differed between fen types, specifically in terms of saprotrophs, arbuscular mycorrhiza and grazers of bacteria. Our study provides the insight that microbial eukaryotes mainly define the seasonal dynamics of microbiomes in rewetted fen peatlands. Accordingly, future research should unravel the importance of eukaryotes for biogeochemical processes, especially the under-characterized protists and metazoa, in these novel yet poorly understood ecosystems.

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 t ...

International audience; Climate change-driven increases in winter temperatures positively affect conditions for shrub growth in arctic tundra by decreasing plant frost damage and stimulation of nutrient availability. However, the extent to which shrubs may benefit from these conditions may be strongly dependent on the following spring climate. Species-specific differences in phenology and spring frost sensitivity likely affect shrub growth responses to warming. Additionally, effects of changes in winter and spring climate may differ over small spatial scales, as shrub growth may be dependent on natural variation in snow cover, shrub density and cryoturbation. We investigated the effects of winter warming and altered spring climate on growing-season performance of three common and widespread shrub species in cryoturbated non-sorted circle arctic tundra. By insulating sparsely vegetated non-sorted circles and parts of the surrounding heath with additional snow or gardening fleeces, we created two climate change scenarios: snow addition increased soil temperatures in autumn and winter and delayed snowmelt timing without increasing spring temperatures, whereas fleeces increased soil temperature similarly in autumn and winter, but created warmer spring conditions without altering snowmelt timing. Winter warming affected shrub performance, but the direction and magnitude were species-specific and dependent on spring conditions. Spring warming advanced, and later snowmelt delayed canopy green-up. The fleece treatment did not affect shoot growth and biomass in any shrub species despite decreasing leaf frost damage in Empetrum nigrum. Snow addition decreased frost damage and stimulated growth of Vaccinium vitis-idaea by c. 50%, while decreasing Betula nana growth (p < .1). All of these effects were consistent the mostly barren circles and surrounding heath. Synthesis. In cryoturbated arctic tundra, growth of Vaccinium vitis-idaea may substantially increase when a thicker snow cover delays snowmelt, whereas in longer term, warmer winters and springs may favour E. nigrum instead. This may affect shrub community composition and cover, with potentially far-reaching effects on arctic ecosystem functioning via its effects on cryoturbation, carbon cycling and trophic cascading. Our results highlight the importance of disentangling effects of winter and spring climate change timing and nature, as spring conditions are a crucial factor in determining the impact of winter warming on plant performance.

Root phenology is important in controlling carbon and nutrient fluxes in terrestrial ecosystems, yet, remains largely unexplored, especially in the Arctic. We compared below- and aboveground phenology and ending of the growing season in two contrasting vegetation types of subarctic tundra: heath and meadow, and their response to experimental warming in autumn. Root phenology was measured in-situ with minirhizotrons and compared with aboveground phenology assessed with repeat digital photography. The end of the growing season, both below- and aboveground, was similar in meadow and heath and the belowground growing season ended later than aboveground in the two vegetation types. Root growth was higher and less equally distributed over time in meadow compared to heath. The warming treatment increased air and soil temperature by 0.5 °C and slightly increased aboveground greenness, but did not affect root growth or prolong the below- and aboveground growing season in either of the vegetation types. These results imply that vegetation types differ in root dynamics and suggest that other factors than temperature control autumnal root growth in these ecosystems. Further investigations of root phenology will help to identify those drivers, in which including responses of functionally contrasting vegetation types will help to estimate how climate change affects belowground processes and their roles in ecosystem function.

As global climate change advances, shifts in winter precipitation are becoming more common in high latitude ecosystems, resulting in less insulating snow cover and deeper soil frost. Long-term alterations to soil frost can impact on ecosystem processes such as decomposition, microbial activity and vegetation dynamics. In this study we utilized the longest running, well-characterized soil frost manipulation experiment in a boreal forest. We measured nematode family composition and feeding group abundances at four different soil layer depths from plots that had been subjected to deep soil frost for one and 11 years. The overall abundance of nematodes and the different feeding groups were unaffected by deep soil frost. However, a higher Maturity Index was weakly associated with deep soil frost (indicative of lower nutrient enrichment and more persister nematode (i.e., K-strategist) families), likely due to the loss of nutrients and reduced inputs from inhibited decomposition. Multivariate and regression analyses showed that most nematode families were weakly associated with dominant understory plant species and strongly associated with soil organic matter (SOM). This is probably the result of higher resource availability in the control plots, which is favorable to the nematode community. These results indicate that the nematode community was more strongly driven by the long-term indirect effects of deep soil frost on SOM as opposed to the direct effects. Our findings highlight that the indirect effects of altered winter precipitation and soil frost patterns may be more important than direct winter climate effects. Further, such indirect effects on SOM and the plant community that may affect the nematode community can only be seen in long-term experiments. Finally, given the critical role nematodes play in soil food webs and carbon and nutrient cycling, our results demonstrate the necessity of considering the response of nematodes to global climate change in boreal forest soils.

Since the publication of the original article, the authors noticed some errors in reference citation had been introduced throughout the paper. The following text contains excerpts from the original article and how they should appear with correct referencing. The publisher apologises for any inconvenience this has caused readers.

1. Climate change is a world-wide threat to biodiversity and ecosystem structure, functioning and services. To understand the underlying drivers and mechanisms, and to predict the consequences for nature and people, we urgently need better understanding of the direction and magnitude of climate change impacts across the soil–plant–atmosphere continuum. An increasing number of climate change studies are creating new opportunities for meaningful and high-quality generalizations and improved process understanding. However, significant challenges exist related to data availability and/or compatibility across studies, compromising opportunities for data re-use, synthesis and upscaling. Many of these challenges relate to a lack of an established ‘best practice’ for measuring key impacts and responses. This restrains our current understanding of complex processes and mechanisms in terrestrial ecosystems related to climate change. 2. To overcome these challenges, we collected best-practice methods emerging from major ecological research networks and experiments, as synthesized by 115 experts from across a wide range of scientific disciplines. Our handbook contains guidance on the selection of response variables for different purposes, protocols for standardized measurements of 66 such response variables and advice on data management. Specifically, we recommend a minimum subset of variables that should be collected in all climate change studies to allow data re-use and synthesis, and give guidance on additional variables critical for different types of synthesis and upscaling. The goal of this community effort is to facilitate awareness of the importance and broader application of standardized methods to promote data re-use, availability, compatibility and transparency. We envision improved research practices that will increase returns on investments in individual research projects, facilitate second-order research outputs and create opportunities for collaboration across scientific communities. Ultimately, this should significantly improve the quality and impact of the science, which is required to fulfil society's best practice, coordinated experiments, data management and documentation, ecosystem, experimental macroecology, methodology, open science, vegetation This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. © 2019 The Authors. Methods in Ecology and Evolution published by John Wiley & Sons Ltd on behalf of British Ecological Society.

Climate-warming-induced permafrost thaw exposes large amounts of carbon and nitrogen in soil at considerable depths, below the seasonally thawing active layer. The extent to which plant roots can reach and interact with these hitherto detached, deep carbon and nitrogen stores remains unknown. We aimed to quantify how permafrost thaw affects root dynamics across soil depths and plant functional types compared with above-ground abundance, and potential consequences for plant-soil interactions. A decade of experimental permafrost thaw strongly increased total root length and growth in the active layer, and deep roots invaded the newly thawed permafrost underneath. Root litter input to soil across all depths was 10 times greater with permafrost thaw. Root growth timing was unaffected by experimental permafrost thaw but peaked later in deeper soil, reflecting the seasonally receding thaw front. Deep-rooting species could sequester

1. Despite time-lags and nonlinearity in ecological processes, the majority of our knowledge about ecosystem responses to long-term changes in climate originates from relatively short-term experime ...

Despite their critical role in every terrestrial ecosystem, fine root production and mortality have not been widely compared among systems due to the practical difficulties of belowground research. ...

Decomposition of plant litter is a key control over carbon (C) storage in the soil. The biochemistry of the litter being produced, the environment in which the decomposition is taking place, and the community composition and metabolism of the decomposer organisms exert a combined influence over decomposition rates. As deciduous shrubs and trees are expanding into tundra ecosystems as a result of regional climate warming, this change in vegetation represents a change in litter input to tundra soils and a change in the environment in which litter decomposes. To test the importance of litter biochemistry and environment in determining litter mass loss, we reciprocally transplanted litter between heath (Empetrum nigrum), shrub (Betula nana), and forest (Betula pubescens) at a sub‐Arctic treeline in Sweden. As expansion of shrubs and trees promotes deeper snow, we also used a snow fence experiment in a tundra heath environment to understand the importance of snow depth, relative to other factors, in the decomposition of litter. Our results show that B. pubescens and B. nana leaf litter decomposed at faster rates than E. nigrum litter across all environments, while all litter species decomposed at faster rates in the forest and shrub environments than in the tundra heath. The effect of increased snow on decomposition was minimal, leading us to conclude that microbial activity over summer in the productive forest and shrub vegetation is driving increased mass loss compared to the heath. Using B. pubescens and E. nigrum litter, we demonstrate that degradation of carbohydrate‐C is a significant driver of mass loss in the forest. This pathway was less prominent in the heath, which is consistent with observations that tundra soils typically have high concentrations of “labile” C. This experiment suggests that further expansion of shrubs and trees may stimulate the loss of undecomposed carbohydrate C in the tundra.

The decomposition of large stocks of soil organic carbon in thawing permafrost might depend on more than climate change-induced temperature increases: indirect effects of thawing via altered bacterial community structure (BCS) or rooting patterns are largely unexplored. We used a 10-year in situ permafrost thaw experiment and aerobic incubations to investigate alterations in BCS and potential respiration at different depths, and the extent to which they are related with each other and with root density. Active layer and permafrost BCS strongly differed, and the BCS in formerly frozen soils (below the natural thawfront) converged under induced deep thaw to strongly resemble the active layer BCS, possibly as a result of colonization by overlying microorganisms. Overall, respiration rates decreased with depth and soils showed lower potential respiration when subjected to deeper thaw, which we attributed to gradual labile carbon pool depletion. Despite deeper rooting under induced deep thaw, root density measurements did not improve soil chemistry-based models of potential respiration. However, BCS explained an additional unique portion of variation in respiration, particularly when accounting for differences in organic matter content. Our results suggest that by measuring bacterial community composition, we can improve both our understanding and the modeling of the permafrost carbon feedback. A correction to this article has been published. DOI: 10.1038/s41396-019-0384-1

1. Earlier snowmelt at high latitudes advances above-ground plant phenology, thereby affecting water, nutrient and carbon cycles. Despite the key role of fine roots in these ecosystem processes, phenological responses to earlier snowmelt have never been assessed below-ground.2. We experimentally advanced snowmelt in two contrasting plant community types (heath and meadow) in northern Sweden and measured above- and below-ground phenology (leaf-out, flowering and fine root growth). We expected earlier snowmelt to advance both above- and belowground phenology, and shrub-dominated heath to be more responsive than meadow.3. Snow melted on average 9 days earlier in the manipulated plots than in controls, and soil temperatures were on average 0,9 °C higher during the snowmelt period of 3 weeks. This resulted in small advances in above-ground phenology, but contrary to our expectations, root phenology was unresponsive, with root growth generally starting before leaf-out. These responses to the snowmelt treatment were similar in both plant community types, despite strong differences in dominating plant functional types and root properties, such as root length and turnover.4. The lack of a response in root phenology, despite warmer soil temperatures and aboveground phenological advances, adds evidence that above-ground plant responses might not be directly translated to below-ground plant responses, and that our understanding of factors driving below-ground phenology is still limited, although of major importance for water, nutrient and carbon cycling 1.Earlier snowmelt at high latitudes advances aboveground plant phenology, thereby affecting water, nutrient and carbon cycles. Despite the key role of fine roots in these ecosystem processes, phenological responses to earlier snowmelt have never been assessed belowground.2.We experimentally advanced snowmelt in two contrasting plant community types (heath and meadow) in northern Sweden and measured above- and belowground phenology (leaf-out, flowering and fine root growth). We expected earlier snowmelt to advance both above- and belowground phenology, and shrub-dominated heath to be more responsive than meadow.3.Snow melted on average nine days earlier in the manipulated plots than in controls, and soil temperatures were on average 0.9 °C higher during the snowmelt period of three weeks. This resulted in small advances in aboveground phenology, but contrary to our expectations, root phenology was unresponsive, with root growth generally starting before leaf-out. These responses to the snowmelt treatment were similar in both plant community types, despite strong differences in dominating plant functional types and root properties, such as root length and turnover.4.The lack of a response in root phenology, despite warmer soil temperatures and aboveground phenological advances, adds evidence that aboveground plant responses might not be directly translated to belowground plant responses, and that our understanding of factors driving belowground phenology is still limited, although of major importance for water, nutrient and carbon cycling.

Questions Do introduced herbivores and fire explain the mono-dominance of one legume shrub (Adenocarpus viscosus subsp. spartioides) above the tree line on an oceanic island, given the fact that a number of other legume shrub species are potentially present? What drives the observed landscape-scale life–death pattern within the mono-dominant shrub species population? Location The sub-alpine scrub vegetation of La Palma (Canary Islands, Atlantic Ocean). Methods An 11-yr exclosure experiment with sites distributed along an elevation and orientation gradient was used to identify the influence of introduced herbivore pressure on four endemic shrub species and their seedling recruitment. Further, we assessed the population dynamics and spatial pattern of the dominant shrub species A. viscosus subsp. spartioides. Habitat and vitality characteristics were investigated, assessing spatial topographic features and tree ring-based age estimates. Linear mixed models, ANOVAs, linear regression and variation partitioning were used as statistical analysis tools. Results Outside of the exclosures A. viscosus subsp. spartioides was virtually mono-dominant in the study area, even though other shrub species seem better suited in the absence of introduced herbivores. The presence of introduced herbivores significantly reduced seedling recruitment within all target species, except for A. viscosus subsp. spartioides. Mean age of A. viscosus subsp. spartioides increased with elevation, although vitality analyses indicated that the sub-alpine scrub is elevated above its growth optimum. Three out of four investigated shrub species showed differences in growth height depending on elevation and island orientation. Conclusion Introduced herbivores and fire are identified as key disturbances enhancing the occurrence of A. viscosus subsp. spartioides, a commonly less competitive species. However, Genista benehoavensis, a single island endemic shrub species, seems to be better adapted to the harsh climate conditions of the sub-alpine scrub in the absence of introduced herbivores than any other shrub species.

There is compelling evidence from experiments and observations that climate warming prolongs the growing season in arctic regions. Until now, the start, peak, and end of the growing season, which are used to model influences of vegetation on biogeochemical cycles, were commonly quantified using above-ground phenological data. Yet, over 80% of the plant biomass in arctic regions can be below ground, and the timing of root growth affects biogeochemical processes by influencing plant water and nutrient uptake, soil carbon input and microbial activity. We measured timing of above- and below-ground production in three plant communities along an arctic elevation gradient over two growing seasons. Below-ground production peaked later in the season and was more temporally uniform than above-ground production. Most importantly, the growing season continued c. 50% longer below than above ground. Our results strongly suggest that traditional above-ground estimates of phenology in arctic regions, including remotely sensed information, are not as complete a representation of whole-plant production intensity or duration, as studies that include root phenology. We therefore argue for explicit consideration of root phenology in studies of carbon and nutrient cycling, in terrestrial biosphere models, and scenarios of how arctic ecosystems will respond to climate warming.

Novel disturbance regimes (e.g., introduced herbivores and fire) are among the major drivers of degradation in island ecosystems. High-elevation ecosystems (HEEs) on islands might be especially vulnerable to these disturbances due to high endemism. Here, data from an 11-year exclosure experiment in the HEE of La Palma (Canary Islands) are presented where mammalian herbivores have been introduced. We investigate the combined effect of herbivory and fire on total species richness, seedling richness, and seedling establishment on the whole system and a subset of highly endangered species (target species). Total species richness, seedling species richness, and seedling establishment decreased with herbivory. Five out of eight target species were exclusively found inside the exclosures indicating the negative impact of introduced herbivores on endemic high elevation flora. Target species were generally affected more negatively by introduced herbivores and were subject to significantly higher browsing pressure, probably owing to their lack of defense strategies. A natural wildfire that occurred six years before data sampling substantially increased total species richness and seedling richness in both herbivory exclosure and reference conditions. We conclude that species composition of the HEE has been severely altered by the introduction of non-native herbivores, even though fire seems to have a positive effect on this system.

openaire: EC/H2020/819202/EU//SOS.aquaterra As global temperatures continue to rise, a key uncertainty of climate projections is the microbial decomposition of vast organic carbon stocks in thawing permafrost soils. Decomposition rates can accelerate up to fourfold in the presence of plant roots, and this mechanism—termed the rhizosphere priming effect—may be especially relevant to thawing permafrost soils as rising temperatures also stimulate plant productivity in the Arctic. However, priming is currently not explicitly included in any model projections of future carbon losses from the permafrost area. Here, we combine high-resolution spatial and depth-resolved datasets of key plant and permafrost properties with empirical relationships of priming effects from living plants on microbial respiration. We show that rhizosphere priming amplifies overall soil respiration in permafrost-affected ecosystems by ~12%, which translates to a priming-induced absolute loss of ~40 Pg soil carbon from the northern permafrost area by 2100. Our findings highlight the need to include fine-scale ecological interactions in order to accurately predict large-scale greenhouse gas emissions, and suggest even tighter restrictions on the estimated 200 Pg anthropogenic carbon emission budget to keep global warming below 1.5 °C.