Projected future climatic forcing on the global distribution of vegetation types.

Most emissions scenarios suggest temperature and precipitation regimes will change dramatically across the globe over the next 500 years. These changes will have large impacts on the biosphere, with species forced to migrate to follow their preferred environmental conditions, therefore moving and fragmenting ecosystems. However, most projections of the impacts of climate change only reach 2100, limiting our understanding of the temporal scope of climate impacts, and potentially impeding suitable adaptive action. To address this data gap, we model future climate change every 20 years from 2000 to 2500 CE, under different CO2 emissions scenarios, using a general circulation model. We then apply a biome model to these modelled climate futures, to investigate shifts in climatic forcing on vegetation worldwide, the feasibility of the migration required to enact these modelled vegetation changes, and potential overlap with human land use based on modern-day anthromes. Under a business-as-usual scenario, up to 40% of terrestrial area is expected to be suited to a different biome by 2500. Cold-adapted biomes, particularly boreal forest and dry tundra, are predicted to experience the greatest losses of suitable area. Without mitigation, these changes could have severe consequences both for global biodiversity and the provision of ecosystem services. This article is part of the theme issue 'Ecological novelty and planetary stewardship: biodiversity dynamics in a transforming biosphere'.

Analysis of taiga and tundra lake browning trends from 2002 to 2021 using MODIS data.

Lakes in taiga and tundra regions may be silently undergoing changes due to global warming. One of those changes is browning in lake color. The browning interacts with the carbon cycle, ecosystem dynamics, and water quality in freshwater systems. However, spatiotemporal variabilities of browning in these regions have not been well documented. Using MODIS remote sensing reflectance at near ultraviolet wavelengths from 2002 to 2021 on the Google Earth Engine platform, we quantified long-term browning trends across 7616 lakes (larger than 10 km2) in taiga and tundra biomes. These lakes showed an overall decreased trend in browning (Theil-Sen Slope = 0.00015), with ∼36% of these lakes showing browning trends, and ∼1% of these lakes showing statistically significant (p-value <0.05) browning trends. The browning trends more likely occurred in small lakes in high latitude, low ground ice content regions, where air temperature increased and precipitation decreased. While temperature is projected to increase in response to climate change, our results provide one means to understand how biogeochemical cycles and ecological dynamics respond to climate change.

N-fertilization and disturbance exert long-lasting complex legacies on subarctic ecosystems.

Subarctic ecosystems are subjected to increasing nitrogen (N) enrichment and disturbances that induce particularly strong effects on plant communities when occurring in combination. There is little experimental evidence on the longevity of these effects. We applied N-fertilization (40 kg urea-N ha-1 year-1 for 4 years) and disturbance (removal of vegetation and organic soil layer on one occasion) in two plant communities in a subarctic forest-tundra ecotone in northern Finland. Within the first four years, N-fertilization and disturbance increased the share of deciduous dwarf shrubs and graminoids at the expense of evergreen dwarf shrubs. Individual treatments intensified the other's effect resulting in the strongest increase in graminoids under combined N-fertilization and disturbance. The re-analysis of the plant communities 15 years after cessation of N-fertilization showed an even higher share of graminoids. 18 years after disturbance, the total vascular plant abundance was still substantially lower and the share of graminoids higher. At the same point, the plant community composition was the same under disturbance as under combined N-fertilization and disturbance, indicating that multiple perturbations no longer reinforced the other's effect. Yet, complex interactions between N-fertilization and disturbance were still detected in the soil. We found higher organic N under disturbance and lower microbial N under combined N-fertilization and disturbance, which suggests a lower bioavailability of N sources for soil microorganisms. Our findings support that the effects of enhanced nutrients and disturbance on subarctic vegetation persist over decadal timescales. However, they also highlight the complexity of plant-soil interactions that drive subarctic ecosystem responses to multiple perturbations across varying timescales.

Bacterial and fungal communities in sub-Arctic tundra heaths are shaped by contrasting snow accumulation and nutrient availability.

Climate change is affecting winter snow conditions significantly in northern ecosystems but the effects of the changing conditions for soil microbial communities are not well-understood. We utilized naturally occurring differences in snow accumulation to understand how the wintertime subnivean conditions shape bacterial and fungal communities in dwarf shrub-dominated sub-Arctic Fennoscandian tundra sampled in mid-winter, early, and late growing season. Phospholipid fatty acid (PLFA) and quantitative PCR analyses indicated that fungal abundance was higher in windswept tundra heaths with low snow accumulation and lower nutrient availability. This was associated with clear differences in the microbial community structure throughout the season. Members of Clavaria spp. and Sebacinales were especially dominant in the windswept heaths. Bacterial biomass proxies were higher in the snow-accumulating tundra heaths in the late growing season but there were only minor differences in the biomass or community structure in winter. Bacterial communities were dominated by members of Alphaproteobacteria, Actinomycetota, and Acidobacteriota and were less affected by the snow conditions than the fungal communities. The results suggest that small-scale spatial patterns in snow accumulation leading to a mosaic of differing tundra heath vegetation shapes bacterial and fungal communities as well as soil carbon and nutrient availability.

Root-associated fungi and acquisitive root traits facilitate permafrost nitrogen uptake from long-term experimentally warmed tundra.

Root-associated fungi (RAF) and root traits regulate plant acquisition of nitrogen (N), which is limiting to growth in Arctic ecosystems. With anthropogenic warming, a new N source from thawing permafrost has the potential to change vegetation composition and increase productivity, influencing climate feedbacks. Yet, the impact of warming on tundra plant root traits, RAF, and access to permafrost N is uncertain. We investigated the relationships between RAF, species-specific root traits, and uptake of N from the permafrost boundary by tundra plants experimentally warmed for nearly three decades at Toolik Lake, Alaska. Warming increased acquisitive root traits of nonmycorrhizal and mycorrhizal plants. RAF community composition of ericoid (ERM) but not ectomycorrhizal (ECM) shrubs was impacted by warming and correlated with root traits. RAF taxa in the dark septate endophyte, ERM, and ECM guilds strongly correlated with permafrost N uptake for ECM and ERM shrubs. Overall, a greater proportion of variation in permafrost N uptake was related to root traits than RAF. Our findings suggest that warming Arctic ecosystems will result in interactions between roots, RAF, and newly thawed permafrost that may strongly impact feedbacks to the climate system through mechanisms of carbon and N cycling.

Long-term changes in the daytime growing season carbon dioxide exchange following increased temperature and snow cover in arctic tundra.

Increasing temperatures and winter precipitation can influence the carbon (C) exchange rates in arctic ecosystems. Feedbacks can be both positive and negative, but the net effects are unclear and expected to vary strongly across the Arctic. There is a lack of understanding of the combined effects of increased summer warming and winter precipitation on the C balance in these ecosystems. Here we assess the short-term (1-3 years) and long-term (5-8 years) effects of increased snow depth (snow fences) (on average + 70 cm) and warming (open top chambers; 1-3°C increase) and the combination in a factorial design on all key components of the daytime carbon dioxide (CO2 ) fluxes in a wide-spread heath tundra ecosystem in West Greenland. The warming treatment increased ecosystem respiration (ER) on a short- and long-term basis, while gross ecosystem photosynthesis (GEP) was only increased in the long term. Despite the difference in the timing of responses of ER and GEP to the warming treatment, the net ecosystem exchange (NEE) of CO2 was unaffected in the short term and in the long term. Although the structural equation model (SEM) indicates a direct relationship between seasonal accumulated snow depth and ER and GEP, there were no significant effects of the snow addition treatment on ER or GEP measured over the summer period. The combination of warming and snow addition turned the plots into net daytime CO2 sources during the growing season. Interestingly, despite no significant changes in air temperature during the snow-free time during the experiment, control plots as well as warming plots revealed significantly higher ER and GEP in the long term compared to the short term. This was in line with the satellite-derived time-integrated normalized difference vegetation index of the study area, suggesting that more factors than air temperature are drivers for changes in arctic tundra ecosystems.

Drivers of contemporary and future changes in Arctic seasonal transition dates for a tundra site in coastal Greenland.

Climate change has had a significant impact on the seasonal transition dates of Arctic tundra ecosystems, causing diverse variations between distinct land surface classes. However, the combined effect of multiple controls as well as their individual effects on these dates remains unclear at various scales and across diverse land surface classes. Here we quantified spatiotemporal variations of three seasonal transition dates (start of spring, maximum normalized difference vegetation index (NDVImax ) day, end of fall) for five dominating land surface classes in the ice-free Greenland. Using a distributed snow model, structural equation modeling, and a random forest model, based on ground observations and remote sensing data, we assessed the indirect and direct effects of climate, snow, and terrain on seasonal transition dates. We then presented new projections of likely changes in seasonal transition dates under six future climate scenarios. The coupled climate, snow cover, and terrain conditions explained up to 61% of seasonal transition dates across different land surface classes. Snow ending day played a crucial role in the start of spring and timing of NDVImax . A warmer June and a decline in wind could advance the NDVImax day. Increased precipitation and temperature during July-August are the most important for delaying the end of fall. We projected that a 1-4.5°C increase in temperature and a 5%-20% increase in precipitation would lengthen the spring-to-fall period for all five land surface classes by 2050, thus the current order of spring-to-fall lengths for the five land surface classes could undergo notable changes. Tall shrubs and fens would have a longer spring-to-fall period under the warmest and wettest scenario, suggesting a competitive advantage for these vegetation communities. This study's results illustrate controls on seasonal transition dates and portend potential changes in vegetation composition in the Arctic under climate change.

Rapid migration of Mongolian oak into the southern Asian boreal forest.

The migration of trees induced by climatic warming has been observed at many alpine treelines and boreal-tundra ecotones, but the migration of temperate trees into southern boreal forest remains less well documented. We conducted a field investigation across an ecotone of temperate and boreal forests in northern Greater Khingan Mountains of northeast China. Our analysis demonstrates that Mongolian oak (Quercus mongolica), an important temperate tree species, has migrated rapidly into southern boreal forest in synchrony with significant climatic warming over the past century. The average rate of migration is estimated to be 12.0 ± 1.0 km decade-1 , being slightly slower than the movement of isotherms (14.7 ± 6.4 km decade-1 ). The migration rate of Mongolian oak is the highest observed among migratory temperate trees (average rate 4.0 ± 1.0 km decade-1 ) and significantly higher than the rates of tree migration at boreal-tundra ecotones (0.9 ± 0.4 km decade-1 ) and alpine treelines (0.004 ± 0.003 km decade-1 ). Compared with the coexisting dominant boreal tree species, Dahurian larch (Larix gmelinii), temperate Mongolian oak is observed to have significantly lower capacity for light acquisition, comparable water-use efficiency but stronger capacity to utilize nutrients especially the most limiting nutrient, nitrogen. In the context of climatic warming, and in addition to a high seed dispersal capacity and potential thermal niche differences, the advantage of nutrient utilization, reflected by foliar elementomes and stable nitrogen isotope ratios, is also likely a key mechanism for Mongolian oak to coexist with Dahurian larch and facilitate its migration toward boreal forest. These findings highlight a rapid deborealization of southern Asian boreal forest in response to climatic warming.

Cascading effects of earthworm invasion increase graminoid density and rodent grazing intensities.

Human-mediated dispersal of non-native earthworms can cause substantial changes to the functioning and composition of ecosystems previously earthworm-free. Some of these earthworm species have the potential to "geoengineer" soils and increase plant nitrogen (N) uptake. Yet the possible consequences of increased plant N concentrations on rodent grazing remains poorly understood. In this study, we present findings from a common garden experiment with two tundra communities, meadow (forb dominated) and heath (shrub dominated), half of them subjected to 4 years of earthworm presence (Lumbricus spp. and Aporrectodea spp.). Within four summers, our earthworm treatment changed plant community composition by increasing graminoid density by, on average, 94% in the heath vegetation and by 49% in the meadow. Rodent winter grazing was more intense on plants growing in soils with earthworms, an effect that coincided with higher N concentrations in plants, indicating a higher palatability. Even though earthworms reduced soil moisture, plant community productivity, as indicated by vegetation greenness (normalized difference vegetation index), was not negatively impacted. We conclude that earthworm-induced changes in plant composition and trophic interactions may fundamentally alter the functioning of tundra ecosystems.

Limiting resources for soil microbial growth in climate change simulation treatments in the subarctic.

The microbial use of resources to sustain life and reproduce influences for example, decomposition and plant nutrient provisioning. The study of "limiting factors" has shed light on the interaction between plants and their environment. Here, we investigated whether carbon (C), nitrogen (N), or phosphorus (P) was limiting for soil microorganisms in a subarctic tundra heath, and how changes in resource availability associated with climate change affected this. We studied samples in which changes in resource availability due to climate warming were simulated by the addition of birch litter and/or inorganic N. To these soils, we supplied factorial C (as glucose), N (as NH4 NO3 ), and P (as KH2 PO4 /K2 HPO4 ) additions ("limiting factor assays," LFA), to determine the limiting factors. The combination of C and P induced large growth responses in all soils and, combined with a systematic tendency for growth increases by C, this suggested that total microbial growth was primarily limited by C and secondarily by P. The C limitation was alleviated by the field litter treatment and strengthened by N fertilization. The microbial growth response to the LFA-C and LFA-P addition was strongest in the field-treatment that combined litter and N addition. We also found that bacteria were closer to P limitation than fungi. Our results suggest that, under a climate change scenario, increased C availability resulting from Arctic greening, treeline advance, and shrubification will reduce the microbial C limitation, while increased N availability resulting from warming will intensify the microbial C limitation. Our results also suggest that the synchronous increase of both C and N availability might lead to a progressive P limitation of microbial growth, primarily driven by bacteria being closer to P limitation. These shifts in microbial resource limitation might lead to a microbial targeting of the limiting element from organic matter, and also trigger competition for nutrients between plants and microorganisms, thus modulating the productivity of the ecosystem.

Wayfinding across ocean and tundra: what traditional cultures teach us about navigation.

Research on human navigation by psychologists and neuroscientists has come mainly from a limited range of environments and participants inhabiting western countries. By contrast, numerous anthropological accounts illustrate the diverse ways in which cultures adapt to their surrounding environment to navigate. Here, we provide an overview of these studies and relate them to cognitive science research. The diversity of cues in traditional navigation is much higher and multimodal compared with navigation experiments in the laboratory. It typically involves an integrated system of methods, drawing on a detailed understanding of the environmental cues, specific tools, and forms part of a broader cultural system. We highlight recent methodological developments for measuring navigation skill and modelling behaviour that will aid future research into how culture and environment shape human navigation.

Lichen-associated microbial members are prevalent in the snow microbiome of a sub-arctic alpine tundra.

Snow is the largest component of the cryosphere, with its cover and distribution rapidly decreasing over the last decade due to climate warming. It is imperative to characterize the snow (nival) microbial communities to better understand the role of microorganisms inhabiting these rapidly changing environments. Here, we investigated the core nival microbiome, the cultivable microbial members, and the microbial functional diversity of the remote Uapishka mountain range, a massif of alpine sub-arctic tundra and boreal forest. Snow samples were taken over a two-month interval along an altitude gradient with varying degree of anthropogenic traffic and vegetation cover. The core snow alpine tundra/boreal microbiome, which was present across all samples, constituted of Acetobacterales, Rhizobiales and Acidobacteriales bacterial orders, and of Mycosphaerellales and Lecanorales fungal orders, with the dominant fungal taxa being associated with lichens. The snow samples had low active functional diversity, with Richness values ranging from 0 to 19.5. The culture-based viable microbial enumeration ranged from 0 to 8.05 × 103 CFUs/mL. We isolated and whole-genome sequenced five microorganisms which included three fungi, one alga, and one potentially novel bacterium of the Lichenihabitans genus; all of which appear to be part of lichen-associated taxonomic clades.

Growth form and leaf habit drive contrasting effects of Arctic amplification in long-lived woody species.

Current global change is inducing heterogeneous warming trends worldwide, with faster rates at higher latitudes in the Northern Hemisphere. Consequently, tundra vegetation is experiencing an increase in growth rate and uneven but expanding distribution. Yet, the drivers of this heterogeneity in woody species responses are still unclear. Here, applying a retrospective approach and focusing on long-term responses, we aim to get insight into growth trends and climate sensitivity of long-lived woody species belonging to different functional types with contrasting growth forms and leaf habits (shrub vs. tree and deciduous vs. evergreen). A total of 530 samples from 7 species (common juniper, dwarf birch, woolly willow, Norway spruce, lodgepole pine, rowan, and downy birch) were collected in 10 sites across Iceland. We modelled growth trends and contrasted yearly ring-width measurements, filtering in high- and low-frequency components, with precipitation, land- and sea-surface temperature records (1967-2018). Shrubs and trees showed divergent growth trends, with shrubs closely tracking the recent warming, whereas trees, especially broadleaved, showed strong fluctuations but no long-term growth trends. Secondary growth, particularly the high-frequency component, was positively correlated with summer temperatures for most of the species. On the contrary, growth responses to sea surface temperature, especially in the low frequency, were highly diverging between growth forms, with a strong positive association for shrubs and a negative for trees. Within comparable vegetation assemblage, long-lived woody species could show contrasting responses to similar climatic conditions. Given the predominant role of oceanic masses in shaping climate patterns in the Arctic and Low Arctic, further investigations are needed to deepen the knowledge on the complex interplay between coastal tundra ecosystems and land-sea surface temperature dynamics.

Ambient Temperature Effects on the Spring and Autumn Somatic Growth Trajectory Show Plasticity in the Photoneuroendocrine Response Pathway in the Tundra Vole.

Seasonal mammals register photoperiodic changes through the photoneuroendocrine system enabling them to time seasonal changes in growth, metabolism, and reproduction. To a varying extent, proximate environmental factors like ambient temperature (Ta) modulate timing of seasonal changes in physiology, conferring adaptive flexibility. While the molecular photoneuroendocrine pathway governing the seasonal responses is well defined, the mechanistic integration of nonphotoperiodic modulatory cues is poorly understood. Here, we explored the interaction between Ta and photoperiod in tundra voles, Microtus oeconomus, a boreal species in which the main impact of photoperiod is on postnatal somatic growth. We demonstrate that postweaning growth potential depends on both gestational and postweaning patterns of photoperiodic exposure, with the highest growth potential seen in voles experiencing short (8 h) gestational and long (16 h) postweaning photoperiods-corresponding to a spring growth program. Modulation by Ta was asymmetric: low Ta (10 °C) enhanced the growth potential of voles gestated on short photoperiods independent of postweaning photoperiod exposure, whereas in voles gestated on long photoperiods, showing a lower autumn-programmed growth potential, the effect of Ta was highly dependent on postweaning photoperiod. Analysis of the primary molecular elements involved in the expression of a neuroendocrine response to photoperiod, thyrotropin beta subunit (tshß) in the pars tuberalis, somatostatin (srif) in the arcuate nucleus, and type 2/3 deiodinase (dio2/dio3) in the mediobasal hypothalamus identified dio2 as the most Ta-sensitive gene across the study, showing increased expression at higher Ta, while higher Ta reduced somatostatin expression. Contrastingly dio3 and tshß were largely insensitive to Ta. Overall, these observations reveal a complex interplay between Ta and photoperiodic control of postnatal growth in M. oeconomus, and suggest that integration of Ta into the control of growth occurs downstream of the primary photoperiodic response cascade revealing potential adaptivity of small herbivores facing rising temperatures at high latitudes.

Recolonization following past persecution questions the importance of persistent snow cover as a range limiting factor for wolverines.

Globally, climate is changing rapidly, which causes shifts in many species' distributions, stressing the need to understand their response to changing environmental conditions to inform conservation and management. Northern latitudes are expected to experience strongest changes in climate, with milder winters and decreasing snow cover. The wolverine (Gulo gulo) is a circumpolar, threatened carnivore distributed in northern tundra, boreal, and subboreal habitats. Previous studies have suggested that wolverine distribution and reproduction are constrained by a strong association with persistent spring snow cover. We assess this hypothesis by relating spatial distribution of 1589 reproductive events, a fitness-related proxy for female reproduction and survival, to snow cover over two decades. Wolverine distribution has increased and number of reproductive events increased 20 times in areas lacking spring snow cover during our study period, despite low monitoring effort where snow is sparse. Thus, the relationship between reproductive events and persistent spring snow cover weakened during this period. These findings show that wolverine reproductive success and hence distribution are less dependent on spring snow cover than expected. This has important implications for projections of future habitat availability, and thus distribution, of this threatened species. Our study also illustrates how past persecution, or other factors, that have restricted species distribution to remote areas can mask actual effects of environmental parameters, whose importance reveals when populations expand beyond previously restricted ranges. Overwhelming evidence shows that climate change is affecting many species and ecological processes, but forecasting potential consequences on a given species requires longitudinal data to revisit hypotheses and reassess the direction and magnitude of climate effects with new data. This is especially important for conservation-oriented management of species inhabiting dynamic systems where environmental factors and human activities interact, a common scenario for many species in different ecosystems around the globe.

Evolutionary history of the Arctic flora.

The Arctic tundra is a relatively young and new type of biome and is especially sensitive to the impacts of global warming. However, little is known about how the Arctic flora was shaped over time. Here we investigate the origin and evolutionary dynamics of the Arctic flora by sampling 32 angiosperm clades that together encompass 3626 species. We show that dispersal into the Arctic and in situ diversification within the Arctic have similar trends through time, initiating at approximately 10-9 Ma, increasing sharply around 2.6 Ma, and peaking around 1.0-0.7 Ma. Additionally, we discover the existence of a long-term dispersal corridor between the Arctic and western North America. Our results suggest that the initiation and diversification of the Arctic flora might have been jointly driven by progressive landscape and climate changes and sea-level fluctuations since the early Late Miocene. These findings have important conservation implications given rapidly changing climate conditions in the Arctic.

Metaproteomics reveals functional partitioning and vegetational variation among permafrost-affected Arctic soil bacterial communities.

Microbial activity in Arctic soils controls the cycling of significant stores of organic carbon and nutrients. We studied in situ processes in Alaskan soils using original metaproteomic methods in order to relate important heterotrophic functions to microbial taxa and to understand the microbial response to Arctic greening. Major bacterial groups show strong metabolic specialization in organic topsoils. α-/ß-/γ-Proteobacteria specialized in the acquisition of small, soluble compounds, whereas Acidobacteria, Actinobacteria, and other detritosphere groups specialized in the degradation of plant-derived polymers. α-/ß-/γ-Proteobacteria dominated the expression of transporters for common root exudates and limiting nitrogenous compounds, supporting an ecological model of dependence upon plants for carbon and competition with plants for nitrogen. Detritosphere groups specialized in distinct substrates, with Acidobacteria producing the most enzymes for hemicellulose depolymerization. Acidobacteria was the most active group across the three plant ecotypes sampled-the largely nonvascular, lower biomass intertussock and the largely vascular, higher biomass tussock and shrub. Functional partitioning among bacterial groups was stable between plant ecotypes, but certain functions associated with α-/ß-/γ-Proteobacteria were more strongly expressed in higher biomass ecotypes. We show that refined metaproteomic approaches can elucidate soil microbial ecology as well as biogeochemical trajectories of major carbon stocks. IMPORTANCE The Arctic is warming twice as fast as the rest of the planet, and Arctic soils currently store twice as much carbon as the entire atmosphere-two facts that make understanding how Arctic soil microbial communities are responding to climate change particularly urgent. Greening of vegetation cover across the Arctic landscape is one of the most prominent climate-driven shifts in Arctic terrestrial ecology, with potentially profound effects on biogeochemical cycling by the soil microbiome. Here we use metaproteomics to document microbial metabolic functions that drive soil carbon and nutrient cycling processes in an Arctic tundra landscape. We identify functional roles among bacterial taxonomic groups that are largely stable across vegetation types, with certain functions strongly expressed by rhizosphere groups reflecting a community metabolic response to greening.

Plant traits poorly predict winner and loser shrub species in a warming tundra biome.

Climate change is leading to species redistributions. In the tundra biome, shrubs are generally expanding, but not all tundra shrub species will benefit from warming. Winner and loser species, and the characteristics that may determine success or failure, have not yet been fully identified. Here, we investigate whether past abundance changes, current range sizes and projected range shifts derived from species distribution models are related to plant trait values and intraspecific trait variation. We combined 17,921 trait records with observed past and modelled future distributions from 62 tundra shrub species across three continents. We found that species with greater variation in seed mass and specific leaf area had larger projected range shifts, and projected winner species had greater seed mass values. However, trait values and variation were not consistently related to current and projected ranges, nor to past abundance change. Overall, our findings indicate that abundance change and range shifts will not lead to directional modifications in shrub trait composition, since winner and loser species share relatively similar trait spaces.