Elevated CO2 interacts with nutrient inputs to restructure plant communities in phosphorus-limited grasslands.

Globally pervasive increases in atmospheric CO2 and nitrogen (N) deposition could have substantial effects on plant communities, either directly or mediated by their interactions with soil nutrient limitation. While the direct consequences of N enrichment on plant communities are well documented, potential interactions with rising CO2 and globally widespread phosphorus (P) limitation remain poorly understood. We investigated the consequences of simultaneous elevated CO2 (eCO2 ) and N and P additions on grassland biodiversity, community and functional composition in P-limited grasslands. We exposed soil-turf monoliths from limestone and acidic grasslands that have received >25 years of N additions (3.5 and 14 g m-2 year-1 ) and 11 (limestone) or 25 (acidic) years of P additions (3.5 g m-2 year-1 ) to eCO2 (600 ppm) for 3 years. Across both grasslands, eCO2 , N and P additions significantly changed community composition. Limestone communities were more responsive to eCO2 and saw significant functional shifts resulting from eCO2 -nutrient interactions. Here, legume cover tripled in response to combined eCO2 and P additions, and combined eCO2 and N treatments shifted functional dominance from grasses to sedges. We suggest that eCO2 may disproportionately benefit P acquisition by sedges by subsidising the carbon cost of locally intense root exudation at the expense of co-occurring grasses. In contrast, the functional composition of the acidic grassland was insensitive to eCO2 and its interactions with nutrient additions. Greater diversity of P-acquisition strategies in the limestone grassland, combined with a more functionally even and diverse community, may contribute to the stronger responses compared to the acidic grassland. Our work suggests we may see large changes in the composition and biodiversity of P-limited grasslands in response to eCO2 and its interactions with nutrient loading, particularly where these contain a high diversity of P-acquisition strategies or developmentally young soils with sufficient bioavailable mineral P.

Sub-arctic mosses and lichens show idiosyncratic responses to combinations of winter heatwaves, freezing and nitrogen deposition.

Arctic ecosystems are increasingly exposed to extreme climatic events throughout the year, which can affect species performance. Cryptogams (bryophytes and lichens) provide important ecosystem services in polar ecosystems but may be physiologically affected or killed by extreme events. Through field and laboratory manipulations, we compared physiological responses of seven dominant sub-Arctic cryptogams (three bryophytes, four lichens) to single events and factorial combinations of mid-winter heatwave (6°C for 7 days), re-freezing, snow removal and summer nitrogen addition. We aimed to identify which mosses and lichens are vulnerable to these abiotic extremes and if combinations would exacerbate physiological responses. Combinations of extremes resulted in stronger species responses but included idiosyncratic species-specific responses. Species that remained dormant during winter (March), irrespective of extremes, showed little physiological response during summer (August). However, winter physiological activity, and response to winter extremes, was not consistently associated with summer physiological impacts. Winter extremes affect cryptogam physiology, but summer responses appear mild, and lichens affect the photobiont more than the mycobiont. Accounting for Arctic cryptogam response to multiple climatic extremes in ecosystem functioning and modelling will require a better understanding of their winter eco-physiology and repair capabilities.

Bioclimatic atlas of the terrestrial Arctic.

The Arctic is the region on Earth that is warming at the fastest rate. In addition to rising means of temperature-related variables, Arctic ecosystems are affected by increasingly frequent extreme weather events causing disturbance to Arctic ecosystems. Here, we introduce a new dataset of bioclimatic indices relevant for investigating the changes of Arctic terrestrial ecosystems. The dataset, called ARCLIM, consists of several climate and event-type indices for the northern high-latitude land areas > 45°N. The indices are calculated from the hourly ERA5-Land reanalysis data for 1950-2021 in a spatial grid of 0.1 degree (~9 km) resolution. The indices are provided in three subsets: (1) the annual values during 1950-2021; (2) the average conditions for the 1991-2020 climatology; and (3) temporal trends over 1951-2021. The 72-year time series of various climate and event-type indices draws a comprehensive picture of the occurrence and recurrence of extreme weather events and climate variability of the changing Arctic bioclimate.

Arctic greening and browning: Challenges and a cascade of complexities.

Arctic greening (the increase in plant biomass and productivity at high latitudes) is one of the clearest large-scale vegetation changes seen in recent decades. However, despite being the subject of considerable research effort, our understanding of this phenomenon is far from complete. Challenges around remote sensing, process based understanding, and the spatial and temporal heterogeneity of greening-including the opposite process of Arctic browning-challenges our ability to model and predict Arctic vegetation change and its biogeochemical consequences.

The missing pieces for better future predictions in subarctic ecosystems: A Torneträsk case study.

Arctic and subarctic ecosystems are experiencing substantial changes in hydrology, vegetation, permafrost conditions, and carbon cycling, in response to climatic change and other anthropogenic drivers, and these changes are likely to continue over this century. The total magnitude of these changes results from multiple interactions among these drivers. Field measurements can address the overall responses to different changing drivers, but are less capable of quantifying the interactions among them. Currently, a comprehensive assessment of the drivers of ecosystem changes, and the magnitude of their direct and indirect impacts on subarctic ecosystems, is missing. The Torneträsk area, in the Swedish subarctic, has an unrivalled history of environmental observation over 100 years, and is one of the most studied sites in the Arctic. In this study, we summarize and rank the drivers of ecosystem change in the Torneträsk area, and propose research priorities identified, by expert assessment, to improve predictions of ecosystem changes. The research priorities identified include understanding impacts on ecosystems brought on by altered frequency and intensity of winter warming events, evapotranspiration rates, rainfall, duration of snow cover and lake-ice, changed soil moisture, and droughts. This case study can help us understand the ongoing ecosystem changes occurring in the Torneträsk area, and contribute to improve predictions of future ecosystem changes at a larger scale. This understanding will provide the basis for the future mitigation and adaptation plans needed in a changing climate.

Niche differentiation and plasticity in soil phosphorus acquisition among co-occurring plants.

How species coexist despite competing for the same resources that are in limited supply is central to our understanding of the controls on biodiversity1,2. Resource partitioning may facilitate coexistence, as co-occurring species use different sources of the same limiting resource3,4. In plant communities, however, direct evidence for partitioning of the commonly limiting nutrient, phosphorus (P), has remained scarce due to the challenges of quantifying P acquisition from its different chemical forms present in soil5. To address this, we used 33P to directly trace P uptake from DNA, orthophosphate and calcium phosphate into monocultures and mixed communities of plants growing in grassland soil. We show that co-occurring plants acquire P from these important organic and mineral sources in different proportions, and that differences in P source use are consistent with the species' root adaptations for P acquisition. Furthermore, the net benefit arising from niche plasticity (the gain in P uptake for a species in a mixed community compared to monoculture) correlates with species abundance in the wild, suggesting that niche plasticity for P is a driver of community structure. This evidence for P resource partitioning and niche plasticity may explain the high levels of biodiversity frequently found in P-limited ecosystems worldwide6,7.

The Regulation of Plant Secondary Metabolism in Response to Abiotic Stress: Interactions Between Heat Shock and Elevated CO2.

Future climate change is set to have an impact on the physiological performance of global vegetation. Increasing temperature and atmospheric CO2 concentration will affect plant growth, net primary productivity, photosynthetic capability, and other biochemical functions that are essential for normal metabolic function. Alongside the primary metabolic function effects of plant growth and development, the effect of stress on plant secondary metabolism from both biotic and abiotic sources will be impacted by changes in future climate. Using an untargeted metabolomic fingerprinting approach alongside emissions measurements, we investigate for the first time how elevated atmospheric CO2 and temperature both independently and interactively impact on plant secondary metabolism through resource allocation, with a resulting "trade-off" between secondary metabolic processes in Salix spp. and in particular, isoprene biosynthesis. Although it has been previously reported that isoprene is suppressed in times of elevated CO2, and that isoprene emissions increase as a response to short-term heat shock, no study has investigated the interactive effects at the metabolic level. We have demonstrated that at a metabolic level isoprene is still being produced during periods of both elevated CO2 and temperature, and that ultimately temperature has the greater effect. With global temperature and atmospheric CO2 concentrations rising as a result of anthropogenic activity, it is imperative to understand the interactions between atmospheric processes and global vegetation, especially given that global isoprene emissions have the potential to contribute to atmospheric warming mitigation.

Arctic browning: Impacts of extreme climatic events on heathland ecosystem CO2 fluxes.

Extreme climatic events are among the drivers of recent declines in plant biomass and productivity observed across Arctic ecosystems, known as "Arctic browning." These events can cause landscape-scale vegetation damage and so are likely to have major impacts on ecosystem CO2 balance. However, there is little understanding of the impacts on CO2 fluxes, especially across the growing season. Furthermore, while widespread shoot mortality is commonly observed with browning events, recent observations show that shoot stress responses are also common, and manifest as high levels of persistent anthocyanin pigmentation. Whether or how this response impacts ecosystem CO2 fluxes is not known. To address these research needs, a growing season assessment of browning impacts following frost drought and extreme winter warming (both extreme climatic events) on the key ecosystem CO2 fluxes Net Ecosystem Exchange (NEE), Gross Primary Productivity (GPP), ecosystem respiration (Reco ) and soil respiration (Rsoil ) was carried out in widespread sub-Arctic dwarf shrub heathland, incorporating both mortality and stress responses. Browning (mortality and stress responses combined) caused considerable site-level reductions in GPP and NEE (of up to 44%), with greatest impacts occurring at early and late season. Furthermore, impacts on CO2 fluxes associated with stress often equalled or exceeded those resulting from vegetation mortality. This demonstrates that extreme events can have major impacts on ecosystem CO2 balance, considerably reducing the carbon sink capacity of the ecosystem, even where vegetation is not killed. Structural Equation Modelling and additional measurements, including decomposition rates and leaf respiration, provided further insight into mechanisms underlying impacts of mortality and stress on CO2 fluxes. The scale of reductions in ecosystem CO2 uptake highlights the need for a process-based understanding of Arctic browning in order to predict how vegetation and CO2 balance will respond to continuing climate change.

Impact of Multiple Ecological Stressors on a Sub-Arctic Ecosystem: No Interaction Between Extreme Winter Warming Events, Nitrogen Addition and Grazing.

Climate change is one of many ongoing human-induced environmental changes, but few studies consider interactive effects between multiple anthropogenic disturbances. In coastal sub-arctic heathland, we quantified the impact of a factorial design simulating extreme winter warming (WW) events (7 days at 6-7°C) combined with episodic summer nitrogen (+N) depositions (5 kg N ha-1) on plant winter physiology, plant community composition and ecosystem CO2 fluxes of an Empetrum nigrum dominated heathland during 3 consecutive years in northern Norway. We expected that the +N would exacerbate any stress effects caused by the WW treatment. During WW events, ecosystem respiration doubled, leaf respiration declined (-58%), efficiency of Photosystem II (Fv/Fm) increased (between 26 and 88%), while cell membrane fatty acids showed strong compositional changes as a result of the warming and freezing. In particular, longer fatty acid chains increased as a result of WW events, and eicosadienoic acid (C20:2) was lower when plants were exposed to the combination of WW and +N. A larval outbreak of geometrid moths (Epirrita autumnata and Operophtera brumata) following the first WW led to a near-complete leaf defoliation of the dominant dwarf shrubs E. nigrum (-87%) and Vaccinium myrtillus (-81%) across all experimental plots. Leaf emergence timing, plant biomass or composition, NDVI and growing season ecosystem CO2 fluxes were unresponsive to the WW and +N treatments. The limited plant community response reflected the relative mild winter freezing temperatures (-6.6°C to -11.8°C) recorded after the WW events, and that the grazing pressure probably overshadowed any potential treatment effects. The grazing pressure and WW both induce damage to the evergreen shrubs and their combination should therefore be even stronger. In addition, +N could have exacerbated the impact of both extreme events, but the ecosystem responses did not support this. Therefore, our results indicate that these sub-arctic Empetrum-dominated ecosystems are highly resilient and that their responses may be limited to the event with the strongest impact.

Understanding the drivers of extensive plant damage in boreal and Arctic ecosystems: Insights from field surveys in the aftermath of damage.

The exact cause of population dieback in nature is often challenging to identify retrospectively. Plant research in northern regions has in recent decades been largely focussed on the opposite trend, namely increasing populations and higher productivity. However, a recent unexpected decline in remotely-sensed estimates of terrestrial Arctic primary productivity suggests that warmer northern lands do not necessarily result in higher productivity. As large-scale plant dieback may become more frequent at high northern latitudes with increasing frequency of extreme events, understanding the drivers of plant dieback is especially urgent. Here, we report on recent extensive damage to dominant, short, perennial heath and tundra plant populations in boreal and Arctic Norway, and assess the potential drivers of this damage. In the High-Arctic archipelago of Svalbard, we recorded that 8-50% of Cassiope tetragona and Dryas octopetala shoots were dead, and that the ratios of dead shoots increased from 2014 to 2015. In boreal Norway, 38-63% of Calluna vulgaris shoots were dead, while Vaccinium myrtillus had damage to 91% of shoots in forested sites, but was healthy in non-forested sites. Analyses of numerous sources of environmental information clearly point towards a winter climate-related reason for damage to three of these four species. In Svalbard, the winters of 2011/12 and 2014/15 were documented to be unusually severe, i.e. insulation from ambient temperature fluctuation by snow was largely absent, and ground-ice enforced additional stress. In boreal Norway, the 2013/14 winter had a long period with very little snow combined with extremely low precipitation rates, something which resulted in frost drought of uncovered Calluna plants. However, extensive outbreaks of a leaf-defoliating geometrid moth were identified as the driver of Vaccinium mortality. These results suggest that weather and biotic extreme events potentially have strong impacts on the vegetation state of northern lands.

Tight coupling of leaf area index to canopy nitrogen and phosphorus across heterogeneous tallgrass prairie communities.

Nitrogen (N) and phosphorus (P) are limiting nutrients for many plant communities worldwide. Foliar N and P along with leaf area are among the most important controls on photosynthesis and hence productivity. However, foliar N and P are typically assessed as species level traits, whereas productivity is often measured at the community scale. Here, we compared the community-level traits of leaf area index (LAI) to total foliar nitrogen (TFN) and total foliar phosphorus (TFP) across nearly three orders of magnitude LAI in grazed and ungrazed tallgrass prairie in north-eastern Kansas, USA. LAI was strongly correlated with both TFN and TFP across communities, and also within plant functional types (grass, forb, woody, and sedge) and grazing treatments (bison or cattle, and ungrazed). Across almost the entire range of LAI values and contrasting communities, TFN:TFP ratios indicated co-limitation by N and P in almost all communities; this may further indicate a community scale trend of an optimal N and P allocation per unit leaf area for growth. Previously, results from the arctic showed similar tight relationships between LAI:TFN, suggesting N is supplied to canopies to maximize photosynthesis per unit leaf area. This tight coupling between LAI, N, and P in tallgrass prairie suggests a process of optimal allocation of N and P, wherein LAI remains similarly constrained by N and P despite differences in species composition, grazing, and canopy density.

Nitrogen accumulation and partitioning in a High Arctic tundra ecosystem from extreme atmospheric N deposition events.

Arctic ecosystems are threatened by pollution from recently detected extreme atmospheric nitrogen (N) deposition events in which up to 90% of the annual N deposition can occur in just a few days. We undertook the first assessment of the fate of N from extreme deposition in High Arctic tundra and are presenting the results from the whole ecosystem (15)N labelling experiment. In 2010, we simulated N depositions at rates of 0, 0.04, 0.4 and 1.2 g Nm(-2)yr(-1), applied as (15)NH4(15)NO3 in Svalbard (79(°)N), during the summer. Separate applications of (15)NO3(-) and (15)NH4(+) were also made to determine the importance of N form in their retention. More than 95% of the total (15)N applied was recovered after one growing season (~90% after two), demonstrating a considerable capacity of Arctic tundra to retain N from these deposition events. Important sinks for the deposited N, regardless of its application rate or form, were non-vascular plants>vascular plants>organic soil>litter>mineral soil, suggesting that non-vascular plants could be the primary component of this ecosystem to undergo measurable changes due to N enrichment from extreme deposition events. Substantial retention of N by soil microbial biomass (70% and 39% of (15)N in organic and mineral horizon, respectively) during the initial partitioning demonstrated their capacity to act as effective buffers for N leaching. Between the two N forms, vascular plants (Salix polaris) in particular showed difference in their N recovery, incorporating four times greater (15)NO3(-) than (15)NH4(+), suggesting deposition rich in nitrate will impact them more. Overall, these findings show that despite the deposition rates being extreme in statistical terms, biologically they do not exceed the capacity of tundra to sequester pollutant N during the growing season. Therefore, current and future extreme events may represent a major source of eutrophication.

The influence of vegetation and soil characteristics on active-layer thickness of permafrost soils in boreal forest.

Carbon release from thawing permafrost soils could significantly exacerbate global warming as the active-layer deepens, exposing more carbon to decay. Plant community and soil properties provide a major control on this by influencing the maximum depth of thaw each summer (active-layer thickness; ALT), but a quantitative understanding of the relative importance of plant and soil characteristics, and their interactions in determine ALTs, is currently lacking. To address this, we undertook an extensive survey of multiple vegetation and edaphic characteristics and ALTs across multiple plots in four field sites within boreal forest in the discontinuous permafrost zone (NWT, Canada). Our sites included mature black spruce, burned black spruce and paper birch, allowing us to determine vegetation and edaphic drivers that emerge as the most important and broadly applicable across these key vegetation and disturbance gradients, as well as providing insight into site-specific differences. Across sites, the most important vegetation characteristics limiting thaw (shallower ALTs) were tree leaf area index (LAI), moss layer thickness and understory LAI in that order. Thicker soil organic layers also reduced ALTs, though were less influential than moss thickness. Surface moisture (0-6 cm) promoted increased ALTs, whereas deeper soil moisture (11-16 cm) acted to modify the impact of the vegetation, in particular increasing the importance of understory or tree canopy shading in reducing thaw. These direct and indirect effects of moisture indicate that future changes in precipitation and evapotranspiration may have large influences on ALTs. Our work also suggests that forest fires cause greater ALTs by simultaneously decreasing multiple ecosystem characteristics which otherwise protect permafrost. Given that vegetation and edaphic characteristics have such clear and large influences on ALTs, our data provide a key benchmark against which to evaluate process models used to predict future impacts of climate warming on permafrost degradation and subsequent feedback to climate.

The effects of foundation species on community assembly: a global study on alpine cushion plant communities.

Foundation species can change plant community structure by modulating important ecological processes such as community assembly, yet this topic is poorly understood. In alpine systems, cushion plants commonly act as foundation species by ameliorating local conditions. Here, we analyze diversity patterns of species' assembly within cushions and in adjacent surrounding open substrates (83 sites across five continents) calculating floristic dissimilarity between replicate plots, and using linear models to analyze relationships between microhabitats and species diversity. Floristic dissimilarity did not change across biogeographic regions, but was consistently lower in the cushions than in the open microhabitat. Cushion plants appear to enable recruitment of many relatively stress-intolerant species that otherwise would not establish in these communities, yet the niche space constructed by cushion plants supports a more homogeneous composition of species than the niche space beyond the cushion's influence. As a result, cushion plants support higher α-diversity and a larger species pool, but harbor assemblies with lower ß-diversity than open microhabitats. We conclude that habitats with and without dominant foundation species can strongly differ in the processes that drive species recruitment, and thus the relationship between local and regional species diversity.

Arctic soil microbial diversity in a changing world.

The Arctic region is a unique environment, subject to extreme environmental conditions, shaping life therein and contributing to its sensitivity to environmental change. The Arctic is under increasing environmental pressure from anthropogenic activity and global warming. The unique microbial diversity of Arctic regions, that has a critical role in biogeochemical cycling and in the production of greenhouse gases, will be directly affected by and affect, global changes. This article reviews current knowledge and understanding of microbial taxonomic and functional diversity in Arctic soils, the contributions of microbial diversity to ecosystem processes and their responses to environmental change.

Climatic and biotic extreme events moderate long-term responses of above- and belowground sub-Arctic heathland communities to climate change.

Climate change impacts are not uniform across the Arctic region because interacting factors causes large variations in local ecosystem change. Extreme climatic events and population cycles of herbivores occur simultaneously against a background of gradual climate warming trends and can redirect ecosystem change along routes that are difficult to predict. Here, we present the results from sub-Arctic heath vegetation and its belowground micro-arthropod community in response to the two main drivers of vegetation damage in this region: extreme winter warming events and subsequent outbreaks of the defoliating autumnal moth caterpillar (Epirrita autumnata). Evergreen dwarf shrub biomass decreased (30%) following extreme winter warming events and again by moth caterpillar grazing. Deciduous shrubs that were previously exposed to an extreme winter warming event were not affected by the moth caterpillar grazing, while those that were not exposed to warming events (control plots) showed reduced (23%) biomass from grazing. Cryptogam cover increased irrespective of grazing or winter warming events. Micro-arthropods declined (46%) following winter warming but did not respond to changes in plant community. Extreme winter warming and caterpillar grazing suppressed the CO2 fluxes of the ecosystem. Evergreen dwarf shrubs are disadvantaged in a future sub-Arctic with more stochastic climatic and biotic events. Given that summer warming may further benefit deciduous over evergreen shrubs, event and trend climate change may both act against evergreen shrubs and the ecosystem functions they provide. This is of particular concern given that Arctic heath vegetation is typically dominated by evergreen shrubs. Other components of the vegetation showed variable responses to abiotic and biotic events, and their interaction indicates that sub-Arctic vegetation response to multiple pressures is not easy to predict from single-factor responses. Therefore, while biotic and climatic events may have clear impacts, more work is needed to understand their net effect on Arctic ecosystems.

Facilitative plant interactions and climate simultaneously drive alpine plant diversity.

Interactions among species determine local-scale diversity, but local interactions are thought to have minor effects at larger scales. However, quantitative comparisons of the importance of biotic interactions relative to other drivers are rarely made at larger scales. Using a data set spanning 78 sites and five continents, we assessed the relative importance of biotic interactions and climate in determining plant diversity in alpine ecosystems dominated by nurse-plant cushion species. Climate variables related with water balance showed the highest correlation with richness at the global scale. Strikingly, although the effect of cushion species on diversity was lower than that of climate, its contribution was still substantial. In particular, cushion species enhanced species richness more in systems with inherently impoverished local diversity. Nurse species appear to act as a 'safety net' sustaining diversity under harsh conditions, demonstrating that climate and species interactions should be integrated when predicting future biodiversity effects of climate change.

Leaf and fine root carbon stocks and turnover are coupled across Arctic ecosystems.

Estimates of vegetation carbon pools and their turnover rates are central to understanding and modelling ecosystem responses to climate change and their feedbacks to climate. In the Arctic, a region containing globally important stores of soil carbon, and where the most rapid climate change is expected over the coming century, plant communities have on average sixfold more biomass below ground than above ground, but knowledge of the root carbon pool sizes and turnover rates is limited. Here, we show that across eight plant communities, there is a significant positive relationship between leaf and fine root turnover rates (r(2) = 0.68, P < 0.05), and that the turnover rates of both leaf (r(2) = 0.63, P < 0.05) and fine root (r(2) = 0.55, P < 0.05) pools are strongly correlated with leaf area index (LAI, leaf area per unit ground area). This coupling of root and leaf dynamics supports the theory of a whole-plant economics spectrum. We also show that the size of the fine root carbon pool initially increases linearly with increasing LAI, and then levels off at LAI = 1 m(2) m(-2), suggesting a functional balance between investment in leaves and fine roots at the whole community scale. These ecological relationships not only demonstrate close links between above and below-ground plant carbon dynamics but also allow plant carbon pool sizes and their turnover rates to be predicted from the single readily quantifiable (and remotely sensed) parameter of LAI, including the possibility of estimating root data from satellites.