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.

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.

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.

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.

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.

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.

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.

Parasitic plant litter input: a novel indirect mechanism influencing plant community structure.

Parasitic plants have major impacts on plant community structure through their direct negative influence on host productivity and competitive ability. However, the possibility that these parasites may also have indirect impacts on community structure (via the mechanism of nutrient-rich litter input) while long hypothesized, has remained unsupported until now. Using the hemiparasite Rhinanthus minor, we established experimental grassland mesocosms to quantify the impacts of Rhinanthus litter and parasitism across two soil fertility levels. We measured the biomass and tissue nutrient concentration of three functional groups within these communities to determine their physiological response to resource abstraction and litter input by the parasite. We show that Rhinanthus alters the biomass and nutrient status of co-occurring plants with contrasting effects on different functional groups via the mechanism of nutrient-rich litter input. Critically, in the case of grass and total community biomass, this partially negates biomass reductions caused directly by parasitism. This demonstrates that the influence of parasitic plant litter on plant community structure can be of equal importance to the much-reported direct impacts of parasitism. We must consider both positive indirect (litter) and negative direct (parasitism) impacts of parasitic plants to understand their role in structuring plant communities.

The role of mosses in carbon uptake and partitioning in arctic vegetation.

The Arctic is already experiencing changes in plant community composition, so understanding the contribution of different vegetation components to carbon (C) cycling is essential in order to accurately quantify ecosystem C balance. Mosses contribute substantially to biomass, but their impact on carbon use efficiency (CUE) - the proportion of gross primary productivity (GPP) incorporated into growth - and aboveground versus belowground C partitioning is poorly known. We used (13) C pulse-labelling to trace assimilated C in mosses (Sphagnum sect. Acutifolia and Pleurozium schreberi) and in dwarf shrub-P. schreberi vegetation in sub-Arctic Finland. Based on (13) C pools and fluxes, we quantified the contribution of mosses to GPP, CUE and partitioning. Mosses incorporated 20 ± 9% of total ecosystem GPP into biomass. CUE of Sphagnum was 68-71%, that of P. schreberi was 62-81% and that of dwarf shrub-P. schreberi vegetation was 58-74%. Incorporation of C belowground was 10 ± 2% of GPP, while vascular plants alone incorporated 15 ± 4% of their fixed C belowground. We have demonstrated that mosses strongly influence C uptake and retention in Arctic dwarf shrub vegetation. They increase CUE, and the fraction of GPP partitioned aboveground. Arctic C models must include mosses to accurately represent ecosystem C dynamics.

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.

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.

Impacts of winter icing events on the growth, phenology and physiology of sub-arctic dwarf shrubs.

The Arctic is experiencing the greatest climate change in winter, including increases in freeze-thaw cycles that can result in ice encasement of vegetation. Ice encasement can expose plants to hypoxia and greater temperature extremes, but currently the impacts of icing on plants in the field remain little understood. With this in mind, a unique field manipulation experiment was established in heathland in northern Sweden with ice encasement simulated in early March 2008, 2009 and 2010 until natural thaw each spring. In the following summers we assessed the impacts on flowering, bud phenology, shoot growth and mortality and leaf damage (measured by chlorophyll fluorescence and electrolyte leakage) of the three dominant dwarf shrub species Empetrum nigrum, Vaccinium vitis-idaea (both evergreen) and Vaccinium myrtillus (deciduous). Two consecutive winters of icing decreased V. vitis-idaea flowering by 57%, while flowering of V. myrtillus and E. nigrum remained unaffected. Vaccinium myrtillus showed earlier budburst but shoot growth for all species was unchanged. Shoot mortality of V. myrtillus and V. vitis-idaea increased after the first year (by 70 and 165%, respectively) and again for V. myrtillus following the third year (by 67%), while E. nigrum shoot mortality remained unaffected, as were chlorophyll fluorescence and electrolyte leakage in all species. Overall, the sub-arctic heathland was relatively tolerant to icing, but the considerable shoot mortality of V. myrtillus contrasting with the general tolerance of E. nigrum suggests plant community structure in the longer term could change if winters continue to see a greater frequency of icing events.

Ecosystem response to climatic change: the importance of the cold season.

Winter climate and snow cover are the important drivers of plant community development in polar regions. However, the impacts of changing winter climate and associated changes in snow regime have received much less attention than changes during summer. Here, we synthesize the results from studies on the impacts of extreme winter weather events on polar heathland and lichen communities. Dwarf shrubs, mosses and soil arthropods were negatively impacted by extreme warming events while lichens showed variable responses to changes in extreme winter weather events. Snow mould formation underneath the snow may contribute to spatial heterogeneity in plant growth, arthropod communities and carbon cycling. Winter snow cover and depth will drive the reported impacts of winter climate change and add to spatial patterns in vegetation heterogeneity. The challenges ahead lie in obtaining better predictions on the snow patterns across the landscape and how these will be altered due to winter climate change.

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.

Impacts of extreme winter warming events on plant physiology in a sub-Arctic heath community.

Insulation provided by snow cover and tolerance of freezing by physiological acclimation allows Arctic plants to survive cold winter temperatures. However, both the protection mechanisms may be lost with winter climate change, especially during extreme winter warming events where loss of snow cover from snow melt results in exposure of plants to warm temperatures and then returning extreme cold in the absence of insulating snow. These events cause considerable damage to Arctic plants, but physiological responses behind such damage remain unknown. Here, we report simulations of extreme winter warming events using infrared heating lamps and soil warming cables in a sub-Arctic heathland. During these events, we measured maximum quantum yield of photosystem II (PSII), photosynthesis, respiration, bud swelling and associated bud carbohydrate changes and lipid peroxidation to identify physiological responses during and after the winter warming events in three dwarf shrub species: Empetrum hermaphroditum, Vaccinium vitis-idaea and Vaccinium myrtillus. Winter warming increased maximum quantum yield of PSII, and photosynthesis was initiated for E. hermaphroditum and V. vitis-idaea. Bud swelling, bud carbohydrate decreases and lipid peroxidation were largest for E. hermaphroditum, whereas V. myrtillus and V. vitis-idaea showed no or less strong responses. Increased physiological activity and bud swelling suggest that sub-Arctic plants can initiate spring-like development in response to a short winter warming event. Lipid peroxidation suggests that plants experience increased winter stress. The observed differences between species in physiological responses are broadly consistent with interspecific differences in damage seen in previous studies, with E. hermaphroditum and V. myrtillus tending to be most sensitive. This suggests that initiation of spring-like development may be a major driver in the damage caused by winter warming events that are predicted to become more frequent in some regions of the Arctic and that may ultimately drive plant community shifts.

Transition zones between vegetation patches in a heterogeneous Arctic landscape: how plant growth and photosynthesis change with abundance at small scales.

Arctic vegetation is often highly heterogeneous, consisting of mosaics of vegetation patches. Vegetation transition zones at the boundaries between patches can therefore make up a significant proportion of the landscape, yet performance of plants in transition zones, compared with their main patches, has not been previously investigated. Not only does this limit our understanding of plant productivity in highly heterogeneous landscapes, this knowledge is also needed to improve our understanding of the controls on Arctic ecosystem productivity, and improve modelling uncertainty in estimates of landscape C fluxes. We investigated changes in annual growth and photosynthetic capacity (A(max)) of two widely distributed dwarf shrubs (Empetrum hermaphroditum and Vaccinium uliginosum) in these species' own patches where they dominate, in transition zones at their patch edges and vegetation beyond these in vegetation where the species had lowest abundance. Maximum shoot growth occurred not in the study species' own patches but in patches where both species mixed with a taller dwarf shrub Betula nana. Here shoot growth was 23 and 25% higher for E. hermaphroditum and V. uliginosum, respectively, compared to within their own patches. However, A(max) was maintained at similar levels at all sites, with a slight tendency toward increased rates in vegetation towards more nutrient-rich, wetter areas compared to the species' own patches, despite these more nutrient-rich areas having greater shading. There was only a very minor tendency towards decreased A(max) in vegetation towards drier, more exposed areas compared to the species' own patches. We show that, despite large changes in abundance and growth, A(max) is not highly variable at small scales, meaning that variation in A(max) is unlikely to be a large source of error when modelling whole-landscape C fluxes using measurements made within patches.