Taking the beat of the Arctic: are lemming population cycles changing due to winter climate?

Reports of fading vole and lemming population cycles and persisting low populations in some parts of the Arctic have raised concerns about the spread of these fundamental changes to tundra food web dynamics. By compiling 24 unique time series of lemming population fluctuations across the circumpolar region, we show that virtually all populations displayed alternating periods of cyclic/non-cyclic fluctuations over the past four decades. Cyclic patterns were detected 55% of the time (n = 649 years pooled across sites) with a median periodicity of 3.7 years, and non-cyclic periods were not more frequent in recent years. Overall, there was an indication for a negative effect of warm spells occurring during the snow onset period of the preceding year on lemming abundance. However, winter duration or early winter climatic conditions did not differ on average between cyclic and non-cyclic periods. Analysis of the time series shows that there is presently no Arctic-wide collapse of lemming cycles, even though cycles have been sporadic at most sites during the last decades. Although non-stationary dynamics appears a common feature of lemming populations also in the past, continued warming in early winter may decrease the frequency of periodic irruptions with negative consequences for 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.

Predator-mediated interactions through changes in predator home range size can lead to local prey exclusion.

The strength of indirect biotic interactions is difficult to quantify in the wild and can alter community composition. To investigate whether the presence of a prey species affects the population growth rate of another prey species, we quantified predator-mediated interaction strength using a multi-prey mechanistic model of predation and a population matrix model. Models were parametrized using behavioural, demographic and experimental data from a vertebrate community that includes the arctic fox (Vulpes lagopus), a predator feeding on lemmings and eggs of various species such as sandpipers and geese. We show that the positive effects of the goose colony on sandpiper nesting success (due to reduction of search time for sandpiper nests) were outweighed by the negative effect of an increase in fox density. The fox numerical response was driven by changes in home range size. As a result, the net interaction from the presence of geese was negative and could lead to local exclusion of sandpipers. Our study provides a rare empirically based model that integrates mechanistic multi-species functional responses and behavioural processes underlying the predator numerical response. This is an important step forward in our ability to quantify the consequences of predation for community structure and dynamics.

Distinct Growth Responses of Tundra Soil Bacteria to Short-Term and Long-Term Warming.

Increases in Arctic temperatures have thawed permafrost and accelerated tundra soil microbial activity, releasing greenhouse gases that amplify climate warming. Warming over time has also accelerated shrub encroachment in the tundra, altering plant input abundance and quality, and causing further changes to soil microbial processes. To better understand the effects of increased temperature and the accumulated effects of climate change on soil bacterial activity, we quantified the growth responses of individual bacterial taxa to short-term warming (3 months) and long-term warming (29 years) in moist acidic tussock tundra. Intact soil was assayed in the field for 30 days using 18O-labeled water, from which taxon-specific rates of 18O incorporation into DNA were estimated as a proxy for growth. Experimental treatments warmed the soil by approximately 1.5°C. Short-term warming increased average relative growth rates across the assemblage by 36%, and this increase was attributable to emergent growing taxa not detected in other treatments that doubled the diversity of growing bacteria. However, long-term warming increased average relative growth rates by 151%, and this was largely attributable to taxa that co-occurred in the ambient temperature controls. There was also coherence in relative growth rates within broad taxonomic levels with orders tending to have similar growth rates in all treatments. Growth responses tended to be neutral in short-term warming and positive in long-term warming for most taxa and phylogenetic groups co-occurring across treatments regardless of phylogeny. Taken together, growing bacteria responded distinctly to short-term and long-term warming, and taxa growing in each treatment exhibited deep phylogenetic organization. IMPORTANCE Soil carbon stocks in the tundra and underlying permafrost have become increasingly vulnerable to microbial decomposition due to climate change. The microbial responses to Arctic warming must be understood in order to predict the effects of future microbial activity on carbon balance in a warming Arctic. In response to our warming treatments, tundra soil bacteria grew faster, consistent with increased rates of decomposition and carbon flux to the atmosphere. Our findings suggest that bacterial growth rates may continue to increase in the coming decades as faster growth is driven by the accumulated effects of long-term warming. Observed phylogenetic organization of bacterial growth rates may also permit taxonomy-based predictions of bacterial responses to climate change and inclusion into ecosystem models.

Arctic rooting depth distribution influences modelled carbon emissions but cannot be inferred from aboveground vegetation type.

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.

Ecosystem and soil respiration radiocarbon detects old carbon release as a fingerprint of warming and permafrost destabilization with climate change.

The permafrost region has accumulated organic carbon in cold and waterlogged soils over thousands of years and now contains three times as much carbon as the atmosphere. Global warming is degrading permafrost with the potential to accelerate climate change as increased microbial decomposition releases soil carbon as greenhouse gases. A 19-year time series of soil and ecosystem respiration radiocarbon from Alaska provides long-term insight into changing permafrost soil carbon dynamics in a warmer world. Nine per cent of ecosystem respiration and 23% of soil respiration observations had radiocarbon values more than 50‰ lower than the atmospheric value. Furthermore, the overall trend of ecosystem and soil respiration radiocarbon values through time decreased more than atmospheric radiocarbon values did, indicating that old carbon degradation was enhanced. Boosted regression tree analyses showed that temperature and moisture environmental variables had the largest relative influence on lower radiocarbon values. This suggested that old carbon degradation was controlled by warming/permafrost thaw and soil drying together, as waterlogged soil conditions could protect soil carbon from microbial decomposition even when thawed. Overall, changing conditions increasingly favoured the release of old carbon, which is a definitive fingerprint of an accelerating feedback to climate change as a consequence of warming and permafrost destabilization. This article is part of the Theo Murphy meeting issue 'Radiocarbon in the Anthropocene'.

Cross-scale regulation of seasonal microclimate by vegetation and snow in the Arctic tundra.

Climate warming is inducing widespread vegetation changes in Arctic tundra ecosystems, with the potential to alter carbon and nutrient dynamics between vegetation and soils. Yet, we lack a detailed understanding of how variation in vegetation and topography influences fine-scale temperatures ("microclimate") that mediate these dynamics, and at what resolution vegetation needs to be sampled to capture these effects. We monitored microclimate at 90 plots across a tundra landscape in western Greenland. Our stratified random study design covered gradients of topography and vegetation, while nested plots (0.8-100 m2 ) enabled comparison across different sampling resolutions. We used Bayesian mixed-effect models to quantify the direct influence of plot-level topography, moisture and vegetation on soil, near-surface and canopy-level temperatures (-6, 2, and 15 cm). During the growing season, colder soils were predicted by shrub cover (-0.24°C per 10% increase), bryophyte cover (-0.35°C per 10% increase), and vegetation height (-0.17°C per 1 cm increase). The same three factors also predicted the magnitude of differences between soil and above-ground temperatures, indicating warmer soils at low cover/height, but colder soils under closed/taller canopies. These findings were consistent across plot sizes, suggesting that spatial predictions of microclimate may be possible at the operational scales of satellite products. During winter, snow cover (+0.75°C per 10 snow-covered days) was the key predictor of soil microclimate. Topography and moisture explained little variation in the measured temperatures. Our results not only underline the close connection of vegetation and snow with microclimate in the Arctic tundra but also point to the need for more studies disentangling their complex interplay across tundra environments and seasons. Future shifts in vegetation cover and height will likely mediate the impact of atmospheric warming on the tundra soil environment, with potential implications for below-ground organisms and ecosystem functioning.

Assessing dynamic vegetation model parameter uncertainty across Alaskan arctic tundra plant communities.

As the Arctic region moves into uncharted territory under a warming climate, it is important to refine the terrestrial biosphere models (TBMs) that help us understand and predict change. One fundamental uncertainty in TBMs relates to model parameters, configuration variables internal to the model whose value can be estimated from data. We incorporate a version of the Terrestrial Ecosystem Model (TEM) developed for arctic ecosystems into the Predictive Ecosystem Analyzer (PEcAn) framework. PEcAn treats model parameters as probability distributions, estimates parameters based on a synthesis of available field data, and then quantifies both model sensitivity and uncertainty to a given parameter or suite of parameters. We examined how variation in 21 parameters in the equation for gross primary production influenced model sensitivity and uncertainty in terms of two carbon fluxes (net primary productivity and heterotrophic respiration) and two carbon (C) pools (vegetation C and soil C). We set up different parameterizations of TEM across a range of tundra types (tussock tundra, heath tundra, wet sedge tundra, and shrub tundra) in northern Alaska, along a latitudinal transect extending from the coastal plain near Utqiagvik to the southern foothills of the Brooks Range, to the Seward Peninsula. TEM was most sensitive to parameters related to the temperature regulation of photosynthesis. Model uncertainty was mostly due to parameters related to leaf area, temperature regulation of photosynthesis, and the stomatal responses to ambient light conditions. Our analysis also showed that sensitivity and uncertainty to a given parameter varied spatially. At some sites, model sensitivity and uncertainty tended to be connected to a wider range of parameters, underlining the importance of assessing tundra community processes across environmental gradients or geographic locations. Generally, across sites, the flux of net primary productivity (NPP) and pool of vegetation C had about equal uncertainty, while heterotrophic respiration had higher uncertainty than the pool of soil C. Our study illustrates the complexity inherent in evaluating parameter uncertainty across highly heterogeneous arctic tundra plant communities. It also provides a framework for iteratively testing how newly collected field data related to key parameters may result in more effective forecasting of Arctic change.

Model responses to CO2 and warming are underestimated without explicit representation of Arctic small-mammal grazing.

We use a simple model of coupled carbon and nitrogen cycles in terrestrial ecosystems to examine how "explicitly representing grazers" vs. "having grazer effects implicitly aggregated in with other biogeochemical processes in the model" alters predicted responses to elevated carbon dioxide and warming. The aggregated approach can affect model predictions because grazer-mediated processes can respond differently to changes in climate compared with the processes with which they are typically aggregated. We use small-mammal grazers in a tundra as an example and find that the typical three-to-four-year cycling frequency is too fast for the effects of cycle peaks and troughs to be fully manifested in the ecosystem biogeochemistry. We conclude that implicitly aggregating the effects of small-mammal grazers with other processes results in an underestimation of ecosystem response to climate change, relative to estimations in which the grazer effects are explicitly represented. The magnitude of this underestimation increases with grazer density. We therefore recommend that grazing effects be incorporated explicitly when applying models of ecosystem response to global change.

Isotopic niche partitioning and individual specialization in an Arctic raptor guild.

Intra- and inter-specific resource partitioning within predator communities is a fundamental component of trophic ecology, and one proposed mechanism for how populations partition resources is through individual niche variation. The Niche Variation Hypothesis (NVH) predicts that inter-individual trait variation leads to functional trade-offs in foraging efficiency, resulting in populations composed of individual dietary specialists. The degree to which niche specialization persists within a population is plastic and responsive to fluctuating resource availability. We quantified niche overlap and tested the NVH within an Arctic raptor guild, focusing on three species that employ different foraging strategies: golden eagles (generalists); gyrfalcons (facultative specialists); and rough-legged hawks (specialists). Tundra ecosystems exhibit cyclic populations of arvicoline rodents (lemmings and voles), providing a unique system in which to examine predator diet in response to interannual fluctuations in resource availability. Using blood δ13C and δ15N values from 189 raptor nestlings on Alaska's Seward Peninsula (2014-2019), we calculated isotopic niche width and used Bayesian stable isotope mixing models (BSIMMs) to characterize individual specialization and test the NVH. Nest-level specialization estimated from stable isotopes was strongly correlated with indices of specialization based on camera trap data. We observed a high degree of isotopic niche overlap between the three species and gyrfalcons displayed a positive relationship between individual specialization and population niche width on an interannual basis consistent with the NVH. Our findings suggest plasticity in niche specialization may reduce intra- and inter-specific resource competition under dynamic ecological conditions.

Divergent shrub-cover responses driven by climate, wildfire, and permafrost interactions in Arctic tundra ecosystems.

The expansion of shrubs across the Arctic tundra may fundamentally modify land-atmosphere interactions. However, it remains unclear how shrub expansion pattern is linked with key environmental drivers, such as climate change and fire disturbance. Here we used 40+ years of high-resolution (~1.0 m) aerial and satellite imagery to estimate shrub-cover change in 114 study sites across four burned and unburned upland (ice-poor) and lowland (ice-rich) tundra ecosystems in northern Alaska. Validated with data from four additional upland and lowland tundra fires, our results reveal that summer precipitation was the most important climatic driver (r = 0.67, p < 0.001), responsible for 30.8% of shrub expansion in the upland tundra between 1971 and 2016. Shrub expansion in the uplands was largely enhanced by wildfire (p < 0.001) and it exhibited positive correlation with fire severity (r = 0.83, p < 0.001). Three decades after fire disturbance, the upland shrub cover increased by 1077.2 ± 83.6 m2  ha-1 , ~7 times the amount identified in adjacent unburned upland tundra (155.1 ± 55.4 m2  ha-1 ). In contrast, shrub cover markedly decreased in lowland tundra after fire disturbance, which triggered thermokarst-associated water impounding and resulted in 52.4% loss of shrub cover over three decades. No correlation was found between lowland shrub cover with fire severity (r = 0.01). Mean summer air temperature (MSAT) was the principal factor driving lowland shrub-cover dynamics between 1951 and 2007. Warmer MSAT facilitated shrub expansion in unburned lowlands (r = 0.78, p < 0.001), but accelerated shrub-cover losses in burned lowlands (r = -0.82, p < 0.001). These results highlight divergent pathways of shrub-cover responses to fire disturbance and climate change, depending on near-surface permafrost and drainage conditions. Our study offers new insights into the land-atmosphere interactions as climate warming and burning intensify in high latitudes.

Alleviation of nutrient co-limitation induces regime shifts in post-fire community composition and productivity in Arctic tundra.

Recent unprecedented fires in the Arctic during the past two decades have indicated a pressing need to understand the long-term ecological impacts of fire in this biome. Anecdotal evidence suggests that tundra fires can induce regime shifts that change tussock tundra to more shrub-dominated ecosystems. However, the ecological mechanisms regulating these shifts are poorly understood, but are hypothesized to involve changes to nutrient availability in this nutrient limited system. Here we conducted a 4-year two-factorial (control: C, nitrogen along: N+ , phosphorus alone: P+ , nitrogen and phosphorus combined: NP+ ) fertilization experiment in both unburned and burned tundra to test this hypothesis after a decade of post-fire recovery. A decade after fire, the burned site exhibited an increase in soil nitrogen and phosphorus availability and a transition toward taller, more productive, and more deciduous vegetation. This shift in vegetation structure, composition, and function was induced at the unburned site through the addition of both NP+ and the alleviation of their co-limitation. Both burned and unburned tundra responded similarly to fertilizer treatments by increasing leaf area index, greenness, and canopy height in NP+ treatments, and exhibited no significant response in individual N+ or P+ treatments. These results point to a greater need to understand coupled carbon, nitrogen, and phosphorus cycles in this system, and suggest that post-fire regime shifts are regulated by the alleviation of nitrogen and phosphorus co-limitation in Arctic tundra.

Will borealization of Arctic tundra herbivore communities be driven by climate warming or vegetation change?

Poleward shifts in species distributions are expected and frequently observed with a warming climate. In Arctic ecosystems, the strong warming trends are associated with increasing greenness and shrubification. Vertebrate herbivores have the potential to limit greening and shrub advance and expansion on the tundra, posing the question of whether changes in herbivore communities could partly mediate the impacts of climate warming on Arctic tundra. Therefore, future changes in the herbivore community in the Arctic tundra will depend on whether the community tracks the changing climates directly (i.e. occurs in response to temperature) or indirectly, in response to vegetation changes (which can be modified by trophic interactions). In this study, we used biogeographic and remotely sensed data to quantify spatial variation in vertebrate herbivore communities across the boreal forest and Arctic tundra biomes. We then tested whether present-day herbivore community structure is determined primarily by temperature or vegetation. We demonstrate that vertebrate herbivore communities are significantly more diverse in the boreal forest than in the Arctic tundra in terms of species richness, phylogenetic diversity and functional diversity. A clear shift in community structure was observed at the biome boundary, with stronger northward declines in diversity in the Arctic tundra. Interestingly, important functional traits characterizing the role of herbivores in limiting tundra vegetation change, such as body mass and woody plant feeding, did not show threshold changes across the biome boundary. Temperature was a more important determinant of herbivore community structure across these biomes than vegetation productivity or woody plant cover. Thus, our study does not support the premise that herbivore-driven limitation of Arctic tundra shrubification or greening would limit herbivore community change in the tundra. Instead, borealization of tundra herbivore communities is likely to result from the direct effect of climate warming.

Pengzhenrongella sicca gen. nov., sp. nov., a new member of suborder Micrococcineae isolated from High Arctic tundra soil.

A yellow bacterial strain, designated LRZ-2T, was isolated from High Arctic tundra near the settlement Ny-Ålesund in the Svalbard Archipelago, Norway. The cells were Gram-stain-positive, aerobic and non-sporulating. Phylogenetic analysis based on 16S rRNA gene sequence comparisons revealed that strain LRZ-2T represented a novel member of the suborder Micrococcineae. Its nearest phylogenetic neighbours were the members of the genus Luteimicrobium, with 16S rRNA gene sequence similarity of 95.3-96.9 %. The average nucleotide identity and digital DNA-DNA hybridization values between the genomes of strain LRZ-2T and its closely related strains were 77.4-74.3 % and 21.4-19.6 %, respectively. The DNA G+C content was 72.4 mol%. The peptidoglycan type of the isolate was A4ß with an interpeptide bridge comprising l-ornithine and d-glutamic acid. The predominant menaquinone was MK-9 (H4) and the major fatty acids were anteiso-C15 : 0, C16 : 0, anteiso-C15 : 1 A, anteiso-C17 : 0 and iso-C15 : 0. The polar lipids were diphosphatidylglycerol, phosphatidylinositol, phosphatidylinositol mannoside, phosphatidylinositol dimannoside, unidentified phosphoglycolipid, four unidentified phospholipids and two unidentified polar lipids. Strain LRZ-2T showed a 16S rRNA gene signature pattern consisting of nucleotides at positions 120 (A), 131-231 (C-G), 196 (C), 342-347 (C-G), 444-490 (A-U), 580-761 (C-G), 602-636 (C-G), 670-736 (A-U), 822-878 (G-C), 823-877 (G-C), 826-874 (C-G), 827 (U), 843 (C), 950-1231 (U-A), 1047-1210 (G-C), 1109 (C), 1145 (G), 1309-1328 (G-C), 1361 (G) and 1383 (C), which clearly distinguished it from all genera previously reported in the suborder Micrococcineae. On the basis of the phylogenetic, phenotypic and chemotaxonomic data, strain LRZ-2T is considered to represent a novel species of a new genus, for which the name Pengzhenrongella sicca gen. nov., sp. nov. is proposed. The type strain of Pengzhenrongella sicca is LRZ-2T (=CCTCC AB 2012163T=DSM 100332T).

Nitrate is an important nitrogen source for Arctic tundra plants.

Plant nitrogen (N) use is a key component of the N cycle in terrestrial ecosystems. The supply of N to plants affects community species composition and ecosystem processes such as photosynthesis and carbon (C) accumulation. However, the availabilities and relative importance of different N forms to plants are not well understood. While nitrate (NO3-) is a major N form used by plants worldwide, it is discounted as a N source for Arctic tundra plants because of extremely low NO3- concentrations in Arctic tundra soils, undetectable soil nitrification, and plant-tissue NO3- that is typically below detection limits. Here we reexamine NO3- use by tundra plants using a sensitive denitrifier method to analyze plant-tissue NO3- Soil-derived NO3- was detected in tundra plant tissues, and tundra plants took up soil NO3- at comparable rates to plants from relatively NO3--rich ecosystems in other biomes. Nitrate assimilation determined by 15N enrichments of leaf NO3- relative to soil NO3- accounted for 4 to 52% (as estimated by a Bayesian isotope-mixing model) of species-specific total leaf N of Alaskan tundra plants. Our finding that in situ soil NO3- availability for tundra plants is high has important implications for Arctic ecosystems, not only in determining species compositions, but also in determining the loss of N from soils via leaching and denitrification. Plant N uptake and soil N losses can strongly influence C uptake and accumulation in tundra soils. Accordingly, this evidence of NO3- availability in tundra soils is crucial for predicting C storage in tundra.

Snow melt stimulates ecosystem respiration in Arctic ecosystems.

Cold seasons in Arctic ecosystems are increasingly important to the annual carbon balance of these vulnerable ecosystems. Arctic winters are largely harsh and inaccessible leading historic data gaps during that time. Until recently, cold seasons have been assumed to have negligible impacts on the annual carbon balance but as data coverage increases and the Arctic warms, the cold season has been shown to account for over half of annual methane (CH4 ) emissions and can offset summer photosynthetic carbon dioxide (CO2 ) uptake. Freeze-thaw cycle dynamics play a critical role in controlling cold season CO2 and CH4 loss, but the relationship has not been extensively studied. Here, we analyze freeze-thaw processes through in situ CO2 and CH4 fluxes in conjunction with soil cores for physical structure and porewater samples for redox biogeochemistry. We find a movement of water toward freezing fronts in soil cores, leaving air spaces in soils, which allows for rapid infiltration of oxygen-rich snow melt in spring as shown by oxidized iron in porewater. The snow melt period coincides with rising ecosystem respiration and can offset up to 41% of the summer CO2 uptake. Our study highlights this important seasonal process and shows spring greenhouse gas emissions are largely due to production from respiration instead of only bursts of stored gases. Further warming is projected to result in increases of snowpack and deeper thaws, which could increase this ecosystem respiration dominate snow melt period causing larger greenhouse gas losses during spring.

Is subarctic forest advance able to keep pace with climate change?

Recent climate warming and scenarios for further warming have led to expectations of rapid movement of ecological boundaries. Here we focus on the circumarctic forest-tundra ecotone (FTE), which represents an important bioclimatic zone with feedbacks from forest advance and corresponding tundra disappearance (up to 50% loss predicted this century) driving widespread ecological and climatic changes. We address FTE advance and climate history relations over the 20th century, using FTE response data from 151 sites across the circumarctic area and site-specific climate data. Specifically, we investigate spatial uniformity of FTE advance, statistical associations with 20th century climate trends, and whether advance rates match climate change velocities (CCVs). Study sites diverged into four regions (Eastern Canada; Central and Western Canada and Alaska; Siberia; and Western Eurasia) based on their climate history, although all were characterized by similar qualitative patterns of behaviour (with about half of the sites showing advancing behaviour). The main associations between climate trend variables and behaviour indicate the importance of precipitation rather than temperature for both qualitative and quantitative behaviours, and the importance of non-growing season as well as growing season months. Poleward latitudinal advance rates differed significantly among regions, being smallest in Eastern Canada (~10 m/year) and largest in Western Eurasia (~100 m/year). These rates were 1-2 orders of magnitude smaller than expected if vegetation distribution remained in equilibrium with climate. The many biotic and abiotic factors influencing FTE behaviour make poleward advance rates matching predicted 21st century CCVs (~103 -104  m/year) unlikely. The lack of empirical evidence for swift forest relocation and the discrepancy between CCV and FTE response contradict equilibrium model-based assumptions and warrant caution when assessing global-change-related biotic and abiotic implications, including land-atmosphere feedbacks and carbon sequestration.

Foraging deeply: Depth-specific plant nitrogen uptake in response to climate-induced N-release and permafrost thaw in the High Arctic.

Warming in the Arctic accelerates top-soil decomposition and deep-soil permafrost thaw. This may lead to an increase in plant-available nutrients throughout the active layer soil and near the permafrost thaw front. For nitrogen (N) limited high arctic plants, increased N availability may enhance growth and alter community composition, importantly affecting the ecosystem carbon balance. However, the extent to which plants can take advantage of this newly available N may be constrained by the following three factors: vertical distribution of N within the soil profile, timing of N-release, and competition with other plants and microorganisms. Therefore, we investigated species- and depth-specific plant N uptake in a high arctic tundra, northeastern Greenland. Using stable isotopic labelling (15 N-NH4 + ), we simulated autumn N-release at three depths within the active layer: top (10 cm), mid (45 cm) and deep-soil near the permafrost thaw front (90 cm). We measured plant species-specific N uptake immediately after N-release (autumn) and after 1 year, and assessed depth-specific microbial N uptake and resource partitioning between above- and below-ground plant parts, microorganisms and soil. We found that high arctic plants actively foraged for N past the peak growing season, notably the graminoid Kobresia myosuroides. While most plant species (Carex rupestris, Dryas octopetala, K. myosuroides) preferred top-soil N, the shrub Salix arctica also effectively acquired N from deeper soil layers. All plants were able to obtain N from the permafrost thaw front, both in autumn and during the following growing season, demonstrating the importance of permafrost-released N as a new N source for arctic plants. Finally, microbial N uptake markedly declined with depth, hence, plant access to deep-soil N pools is a competitive strength. In conclusion, plant species-specific competitive advantages with respect to both time- and depth-specific N-release may dictate short- and long-term plant community changes in the Arctic and consequently, larger-scale climate feedbacks.

Direct and indirect effects of regional and local climatic factors on trophic interactions in the Arctic tundra.

Climate change can impact ecosystems by reshaping the dynamics of resource exploitation for predators and their prey. Alterations of these pathways could be especially intense in ecosystems characterized by a simple trophic structure and rapid warming trends, such as in the Arctic. However, quantifying the multiple direct and indirect pathways through which climate change is likely to alter trophic interactions and their relative strength remains a challenge. Here, we aim to identify direct and indirect causal mechanisms driven by climate affecting predator-prey interactions of species sharing a tundra food web. We based our study on relationships between one Arctic predator (Arctic fox) and its two main prey - lemmings (preferred prey) and snow geese (alternate prey) - which are exposed to variable local and regional climatic factors across years. We used a combination of models mapping multiple causal links among key variables derived from a long-term dataset (21 years). We obtained several possible scenarios linking regional climate factors (Arctic oscillations) and local temperature and precipitation to the breeding of species. Our results suggest that both regional and local climate factors have direct and indirect impacts on the breeding of foxes and geese. Local climate showed a positive causal link with goose nesting success, while both regional and local climate displayed contrasted effects on the proportion of fox breeding. We found no impact of climate on lemming abundance. We observed positive relationships between lemming, fox and goose reproduction highlighting numerical and functional responses of fox to the variability of lemming abundance. Our study measures causal links and strength of interactions in a food web, quantifying both numerical response of a predator and apparent interactions between its two main prey. These results improve our understanding of the complex effects of climate on predator-prey interactions and our capacity to anticipate food web response to ongoing climate change.

Les changements climatiques peuvent avoir un impact sur les écosystèmes au travers des modifications de la dynamique d'exploitation des ressources par les prédateurs et leurs proies. Dans le cas de l'Arctique, caractérisée par un réseau trophique simple et une sensibilité marquée au réchauffement climatique, l'altération de ces relations trophiques pourrait être particulièrement importante. Cependant, la quantification des nombreux liens directs et indirects à travers lesquels les changements climatiques peuvent affecter les interactions trophiques demeure un défi. Notre objectif est d'identifier les mécanismes causaux directs et indirects, sous-tendus par le climat, affectant les interactions prédateur-proie au sein d'un réseau trophique au cœur de la toundra. Notre étude se base sur les relations entre un prédateur (renard arctique) et ses deux proies principales -le lemming (proie préférée) et la grande oie des neiges (proie alternative)- et qui subissent un accroissement des précipitations et des températures au travers des années. Nous avons utilisé une combinaison de modèles illustrant les liens causaux multiples entre les variables clés issues d'une base de données à long-terme (21 ans). Nous avons obtenu plusieurs scénarios possibles reliant les facteurs climatiques régionaux (Oscillation Arctique) et les températures et précipitations locales à la reproduction de nos 3 espèces. Nos résultats suggèrent que les facteurs climatiques régionaux et locaux présentent des impacts directs et indirects sur la reproduction du renard arctique et de l'oie des neiges. Le climat local présente un lien causal positif avec le succès de nidification de l'oie, alors que le climat local et régional démontrent un effet contrasté sur la proportion de renard en reproduction. Aucune relation entre les facteurs climatiques et l'abondance des lemmings n'a été trouvée. Nous avons observé des liens causaux positifs entre la reproduction du lemming, du renard et de l'oie, mettant en évidence les réponses numériques et fonctionnelles du renard arctique face à la variabilité de l'abondance de lemming. Notre étude est une des premières à mesurer les liens causaux et les forces d'interaction entre les espèces partageant un même réseau trophique, quantifiant ainsi la réponse numérique du prédateur et les interactions apparentes entre ses proies principales. Ces résultats améliorent notre compréhension des effets complexes du climat sur les interactions prédateur-proie et notre capacité à anticiper la réponse des réseaux trophiques aux changements climatiques en cours.

A mechanism of expansion: Arctic deciduous shrubs capitalize on warming-induced nutrient availability.

Warming-induced nutrient enrichment in the Arctic may lead to shifts in leaf-level physiological properties and processes with potential consequences for plant community dynamics and ecosystem function. To explore the physiological responses of Arctic tundra vegetation to increasing nutrient availability, we examined how a set of leaf nutrient and physiological characteristics of eight plant species (representing four plant functional groups) respond to a gradient of experimental nitrogen (N) and phosphorus (P) enrichment. Specifically, we examined a set of chlorophyll fluorescence measures related to photosynthetic efficiency, performance and stress, and two leaf nutrient traits (leaf %C and %N), across an experimental nutrient gradient at the Arctic Long Term Ecological Research site, located in the northern foothills of the Brooks Range, Alaska. In addition, we explicitly assessed the direct relationships between chlorophyll fluorescence and leaf %N. We found significant differences in physiological and nutrient traits between species and plant functional groups, and we found that species within one functional group (deciduous shrubs) have significantly greater leaf %N at high levels of nutrient addition. In addition, we found positive, saturating relationships between leaf %N and chlorophyll fluorescence measures across all species. Our results highlight species-specific differences in leaf nutrient traits and physiology in this ecosystem. In particular, the effects of a gradient of nutrient enrichment were most prominent in deciduous plant species, the plant functional group known to be increasing in relative abundance with warming in this ecosystem.