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Plants maintain beneficial mutualistic relationships with the mycobiont communities found in their rhizosphere, leading to an increase in plant productivity and health. In nutrient-depleted substrates like mine tailings, mycobiont inoculation is often recommended to help restore a successful plant cover. Our 15-week greenhouse experiment aimed to assess the individual effects of a dark septate endophyte (Phialocephala fortinii C.J.K. Wang & H.E. Wilcox #4 (KX611529)) and two Helotiales strains (Rhizoscyphus ericae (D.J. Read) W.Y. Zhuang & Korf #22 (EU221877) and Meliniomyces sp. #1 (KT275679)) on the growth of plane-leaved willow (Salix planifolia Pursh) cuttings on sterilized and unsterilized waste rock. Rhizoscyphus ericae increased shoot biomass of cuttings on sterilized waste rock, whereas Meliniomyces sp. had a positive effect on cuttings grown on unsterilized waste rock. However, P. fortinii strain had no effect on the survival rate, shoot production, and biomass production of S. planifolia cuttings. This study demonstrates that controlled inoculation with ecologically well-adapted mycobionts could promote plant establishment and productivity on abandoned waste rock and could be an efficient and integrated biotechnological approach for ecological restoration of Canadian mining boreal ecosystems.

Arctic shrubs reduce surface albedo in winter when branches protrude above the snow. To calculate the albedo of those mixed surfaces, the branch area index (BAI) of Arctic shrubs needs to be known. Moreover, an exposed-vegetation function is required to determine the BAI for protruding branches only. This study used a structural analysis of 30Betula glandulosashrubs, sampled near Umiujaq, northern Quebec, to (i) establish an allometric relationship between shrub height and BAI and (ii) determine a specific exposed-vegetation function for Arctic shrubs. The spectral albedo (400–1080 nm) of mixed surfaces was then simulated with the equations derived from this study and validated with in situ measured spectra. Shrubs were sampled from two sites, one along the coast and the other in a nearby valley. The shrub height–BAI relationship varied between both sites. BAI values of shrubs growing in the wind-sheltered valley were 30%–50% lower. The exposed-vegetation function obtained here differed from the linear functions found in the literature. The linear functions strongly overestimated the BAI of exposed branches. Albedo was well simulated with an accuracy of 3% when using an allometric relationship adapted to the environmental conditions of our study site. However, simulated albedo values were consistently higher than field measurements, probably because radiation absorbed by impurities and buried branches was neglected in the model. We conclude that specific exposed-vegetation and allometric equations need to be implemented in models to accurately simulate the albedo of mixed snow–shrub surfaces.

Les changements environnementaux influencent la végétation nordique, mais également les herbivores qui s’en nourrissent. Comme ces changements s’accentueront, un état de la situation au Nunavik (Québec, Canada) devient nécessaire. Nous discutons ici de l’influence qu’exercent les herbivores vertébrés sur les communautés végétales, et décrivons l’état des populations des deux principales espèces d’herbivores de cette guilde au Nunavik. Réparti en deux troupeaux, le caribou migrateur (Rangifer tarandus) est un élément structurant des écosystèmes du Nunavik. À forte densité, il peut réduire la productivité végétale dans son aire d’estivage et limiter la croissance des espèces consommées. Espèce introduite dans les années 1960, le boeuf musqué (Ovibos moschatus) colonise maintenant l’essentiel des côtes de la région. Quoiqu’il soit connu pour limiter l’expansion arbustive ailleurs dans l’Arctique, l’influence du bovidé au Nunavik demeure marginale jusqu’à maintenant. Sauf pour la bernache du Canada (Branta canadensis) qui niche en grandes colonies, l’influence des autres herbivores vertébrés sur les communautés végétales sera vraisemblablement localisée. L’augmentation du couvert arbustif procure des ressources alimentaires et un abri à diverses espèces qui pourraient voir leur aire de répartition se déplacer vers le nord. Les préférences alimentaires de ces herbivores pourraient moduler la réponse des espèces végétales aux changements climatiques., Environmental changes influence northern plant communities and the populations of herbivores that feed on them. Given the predicted increased rates of change in the north, it is essential to understand the situation in Nunavik (Québec, Canada). This review outlines the influence of vertebrate herbivores on plant communities and describes the population status of the two main members of this guild in the region. The migratory caribou (Rangifer tarandus) is a keystone species, and two herds are present. At high densities, it can reduce plant productivity in its summer range and limit the growth of the species consumed. The muskox (Ovibos moschatus), which was introduced in the 1960s, has now colonized most of the region’s coastal area. Although it may limit shrub expansion elsewhere in the Arctic, to date, its influence in Nunavik remains marginal. Except for the Canada goose (Branta canadensis), which nests in large colonies, the impact of other vertebrate herbivores on plant communities is likely localized. Increases in shrub cover will provide food and shelter for various herbivores that may experience a northward shift in range. Finally, the food preferences of herbivores present in Nunavik is expected to influence the response of plant species to climate change.

The volatile fraction of the leaves of Betula glandulosa Michx. has been investigated for its secondary metabolite composition by GC/MS and GC/FID. The rapid expansion of this shrub species in subarctic landscapes, like the ones found in Nunavik (Northern Québec, Canada), highly impacts ecosystem dynamics. Yet, despite its abundance, few phytochemical investigations have yet been conducted on this species. In this study, we present the first phytochemical investigation of the volatile metabolites of B. glandulosa leaves. Although no essential oil was isolated, volatile compounds were extracted from the hydrosol by steam distillation. The main metabolites observed were linalool (14.6-19.0 %), C

Climate warming is expected to positively alter upward and poleward treelines which are controlled by low temperature and a short growing season. Despite the importance of treelines as a bioassay of climate change, a global field assessment and posterior forecasting of tree growth at annual scales is lacking. Using annually resolved tree-ring data located across Eurasia and the Americas, we quantified and modeled the relationship between temperature and radial growth at treeline during the 20th century. We then tested whether this temperature–growth association will remain stable during the 21st century using a forward model under two climate scenarios (RCP 4.5 and 8.5). During the 20th century, growth enhancements were common in most sites, and temperature and growth showed positive trends. Interestingly, the relationship between temperature and growth trends was contingent on tree age suggesting biogeographic patterns in treeline growth are contingent on local factors besides climate warming. Simulations forecast temperature–growth decoupling during the 21st century. The growing season at treeline is projected to lengthen and growth rates would increase and become less dependent on temperature rise. These forecasts illustrate how growth may decouple from climate warming in cold regions and near the margins of tree existence. Such projected temperature–growth decoupling could impact ecosystem processes in mountain and polar biomes, with feedbacks on climate warming. climate warming, forest limit, growth model, mountain ecosystems, tree rings acceptedVersion

Significance Two defining features of climate change in the Arctic are the rapid decline of sea ice and “shrubification” of the tundra. While previous studies have inferred warming-related linkages between the two, these have been limited to a few locations. Our Pan-Arctic analysis of shrub growth chronologies reveals two important insights. Tundra shrub growth dynamics are associated with sea ice decline throughout the Arctic; however, while shrubs from most locations increased their growth, more than one-third showed evidence of declining growth in response to warming and drying associated with sea ice loss. These results highlight pronounced growth response heterogeneity across the tundra biome that will have important implications for tundra productivity and vegetation–climate feedback., Arctic sea ice extent (SIE) is declining at an accelerating rate with a wide range of ecological consequences. However, determining sea ice effects on tundra vegetation remains a challenge. In this study, we examined the universality or lack thereof in tundra shrub growth responses to changes in SIE and summer climate across the Pan-Arctic, taking advantage of 23 tundra shrub-ring chronologies from 19 widely distributed sites (56°N to 83°N). We show a clear divergence in shrub growth responses to SIE that began in the mid-1990s, with 39% of the chronologies showing declines and 57% showing increases in radial growth (decreasers and increasers, respectively). Structural equation models revealed that declining SIE was associated with rising air temperature and precipitation for increasers and with increasingly dry conditions for decreasers. Decreasers tended to be from areas of the Arctic with lower summer precipitation and their growth decline was related to decreases in the standardized precipitation evapotranspiration index. Our findings suggest that moisture limitation, associated with declining SIE, might inhibit the positive effects of warming on shrub growth over a considerable part of the terrestrial Arctic, thereby complicating predictions of vegetation change and future tundra productivity.

Climate change is impacting surficial geomorphic processes, especially in sensitive areas such as the sub-Arctic. One of the most common examples involves landslides, which often develop in glacio-isostatically raised marine clays in northeastern Canada. One of these sites, an expansive area of complex landslide terrain located at the mouth of the Great Whale River in Nunavik, has already been studied due to its age and morphology. We present new data, based on the multidisciplinary research including geomorphic, dendrochronological, and hydroclimatological analyses, allowing us to determine how contemporary climate change has affected landslide reactivation during the last 80 years. Our research included collecting 60 cores from Picea glauca trees, growing on the marginal zone of a landslide deposit, as well as from a reference site. The tilted trees formed eccentric growth-ring patterns, which provided us with reliable dates on the landslide events. In addition to these dendrochronological data, we studied these landslides using repeated aerial photography, which showed changes in river channel constrictions in the period 1969–2019. Based on the eccentricity index of the tree ring data, we recognized disturbance events due to landslides. We compared these data with the hydroclimatological conditions and found clearly visible correlations between heavy rainfall and discharge (>95th percentile) of the Great Whale River. The increased landslide activity over the past several years can be linked to an increase in extreme summertime rainfall events. Increased landslide activity poses a real threat, through its input of large amounts of fine-grained sediment to the river, causing it to narrow.

Lichens are an important component of biodiversity in northern ecosystems and are involved in diverse ecological processes. They contribute to nutrient availability through nitrogen fixation, are a substantial part of caribou winter diet, and influence global climate by increasing land surface albedo. Over the last decades, increased primary productivity in northern ecosystems, mainly associated with the expansion of shrub species, has led to a decline of lichen-dominated areas. We evaluated the impacts of shrubs on lichens by comparing lichen communities in the open environment and underneath dwarf birch (Betula glandulosa) canopy in Nunavik, Canada. Our results showed a decrease in abundance, richness and evenness and a shift in community composition between open areas and understory. These changes were mainly induced by the presence of a shrub canopy rather than by its characteristics, because shrub height and canopy closure had little effect. Richness and evenness dropped from shrub edge to shrub center, suggesting that the intensity of the decline was positively correlated to the time spent under the shrub canopy. Important changes in lichen communities are therefore expected to occur with further shrub expansion and may have substantial unfavorable implications for global climate and ecosystem functioning.

Species in cold-limited biomes are expected to expand their distribution ranges in response to climate warming. For plants, range shifts can only occur via successful recruitment beyond their current distribution limit. However, many environmental and ecological filters can act upon recruitment and establishment, thereby potentially limiting the expected climate-driven shifts. In this study, we investigate potential mechanical and chemical constraints that vegetation above the tall shrubline in alpine and Arctic tundra could impose upon the successful establishment of willow species in the Canadian Western Arctic. We collected willow seeds from an alpine and an Arctic shrubline and conducted germination trials to test (1) for seedbed preferences among three natural and one experimentally scarified seedbeds, and (2) for vulnerability to allelopathic chemicals produced by ericaceous dwarf shrub species. We found that germination was almost four times higher on manually exposed bare ground than on intact, herbaceous vegetation. Seeds of two willow species, Salix arctica and Salix pulchra, were not affected by leaf extracts from dwarf shrubs, Cassiope tetragona and Vaccinum uliginosum, but the germination of Salix richardsonii was reduced by as much as 24% in the presence of chemicals from C. tetragona. Our results suggest that biotic interactions could limit the predicted expansion of tall shrubs in the tundra by interfering with germination. Seemingly species-specific responses highlight the need for replicated studies across a wider range of species combinations. Potential range shifts may not occur as a uniform translocation of the shrubline, but could change the composition of the plant community by filtering out certain species.

Widespread shrub expansion has been observed at the circumpolar scale in response to climate change. This phenomenon could be inhibited by large herbivores through selective browsing that reduces shrub growth. Our objective was to assess the dynamics of shrub species in the Deception Bay area, in Nunavik (Quebec, Canada), a region in the summer range of the Riviere-aux-Feuilles migratory caribou herd. We surveyed abundance and stem mortality of all shrub species along systematically located transects in a 54 km2 area. Our results showed that shrubs covered 11.5% of the area, and were dominated by prostrate Betula glandulosa (8.7%). Stem mortality of established individuals was greater for Salix planifolia (45.6%) and Salix glauca (46.5%) than for B. glandulosa (9.3%). Using dendrochronological analysis, we determined the age structure of the two dominant erect shrub species in ten 10 m × 10 m plots. We recorded a high number of B. glandulosa and, to a lesser extent, of S. planifolia recruits (71.0 and 4.6 per 100 m2, respectively) established after 1999, when the Riviere-aux-Feuilles herd was abundant. Thus, our results do not support the hypothesis that large herbivores counteracted the positive effects of climate change on shrub recruitment. However, since the structure of the recruits are prostrate rather than erect, as is usually observed with shrub expansion in the warming Arctic, it is likely that herbivory, and/or, the harsh climate, is constraining habitat change.

A circumpolar increase in shrub growth and cover has been underway in Arctic and subarctic ecosystems for the last few decades, but there is considerable spatial heterogeneity in this shrubification process. Although topography, hydrology, and edaphic factors are known to influence shrubification patterns, a better understanding of the landscape‐scale factors driving this phenomenon is needed to accurately predict its impacts on ecosystem function. In this study, we generated land cover change models in order to identify variables driving shrub cover increase near Umiujaq (Quebec, Canada). Using land cover maps from 1990/1994 and 2010, we modeled observed changes using two contrasting conceptual approaches: binomial modeling of transitions to shrub dominance and multinomial modeling of all land cover transitions. Models were used to generate spatially explicit predictions of transition to shrub dominance in the near future as well as long‐term predictions of the abundance of different land cover types. Model predictions were validated using both field data and current Landsat‐derived trends of normalized difference vegetation index (NDVI) increase in the region in order to assess their consistency with observed patterns of change. We found that both variables related to topography and to vegetation were useful in modeling land cover changes occurring near Umiujaq. Shrubs tended to preferentially colonize low‐elevation areas and moderate slopes, while their cover was more likely to increase in the vicinity of existing shrub patches. Deterministic realizations of the spatially explicit models of land cover change had a good predictive capability, although they performed better at predicting the proportion of different cover types than at predicting the precise location of the changes. Binomial models performed as well as multinomial models, indicating that neglecting land cover changes other than shrubification does not result in decreased prediction accuracy. The predicted probabilities of shrub increase in the region were consistent with patterns of change inferred from field data, but only partly supported by recent local increases in NDVI. Our findings increase the current understanding of the factors driving shrubification, while warranting further research on its impacts on ecosystem function and on the link between land cover changes and shifts in remotely sensed vegetation indices.

Warmer summer temperatures and enhanced soil fertility increase shrub growth in tundra ecosystems, and these factors have likely contributed to shrub expansion at the circumpolar scale over the last decades. Conversely, large herbivores have the potential to counteract the positive impacts of climate change on shrub growth. Indeed, by stripping the leaves, herbivores have the potential to control the growth of shrub species and, consequently, limit their expansion. To disentangle the impacts of climate change and herbivory on Betula glandulosa Michx., we conducted a 5-year factorial experiment near Deception Bay, Nunavik, Canada, in which we simulated warmer temperatures, increased nitrogen availability and three caribou browsing intensities during the growing season. At the end of the experiment, we harvested the above-ground biomass of B. glandulosa and conducted dendrochronological analyses on stems. Fertilised plots under ambient temperature had 34% greater shrub biomass than plots assigned to the combined treatment of nitrogen addition and warmer temperatures. Browsing intensity had no effect on final biomass. Nitrogen addition increased radial growth (18%–33%; 3 years out of 5). Overall, browsing had a cumulative negative impact on B. glandulosa radial growth during the 5-year experiment. While browsing had no effect in the first year of the experiment, moderate browsing (leaves stripped on 25% of available shoots) decreased radial growth by 27% at year 2, 32% at year 4 and 27% at year 5. Heavy browsing (leaves stripped on 75% of available shoots annually) decreased radial growth by c. 27% at year 2, 37% at year 3, 50% at year 4 and 48% at year 5. We did not observe significant interactions between browsing, temperature and nitrogen availability. Synthesis. Our results clearly showed that caribou browsing may limit the growth of B. glandulosa, and thus can potentially limit its expansion. Herbivory should thus be considered when predicting tundra vegetation changes in the Arctic, at least in areas with high herbivore densities.

Many tundra plant species rely more heavily on clonal propagation than on sexual reproduction. However, if reproduction bottlenecks are alleviated, shifts in the balance between these strategies can occur. To better understand the colonization and expansion dynamics of clonal dwarf shrubs in a rapidly changing environment, we monitored a crowberry population (Empetrum hermaphroditum) on a sand dune system in subarctic Quebec. Our objectives were to quantify survival, recruitment, and growth and to determine whether performance varied across a topographic and successional gradient. In 2012, we resurveyed a 6-ha plot where all individuals had been counted in 2007. Along the successional gradient, we measured vegetation cover, soil characteristics, shoot elongation, and seed germination. Over the 2007–2012 period, the population continued to experience abundant recruitment, fast growth, and low mortality, resulting in a 40% increase in population size and a 244 m2 increase in cover. Performance patt...

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.

We assessed the response of Salix richardsonii, a deciduous shrub, to climate change by determining the combination of climatic factors that regulated its growth over the past half-century. We tested whether increasing arctic temperatures promote shrub growth and increased cover. We analyzed fifty-four stems (out of seventy sampled) from S. richardsonii shrubs near the Walker Bay research station in Nunavut, Canada (68°21′ N, 108°05′ W) and surveyed shrub cover in 1996 and 2010. We measured annual growth rings, removed the age-related pattern, and used a response function analysis to explore the climate–growth relationship. The standardized chronology was positively associated with mean July temperature, corroborating other evidence that summer temperature is an important driver of shrub radial growth. Basal area increment revealed a long-term increase in radial growth, although it has stabilized this century. Surveys showed no significant increase in shrub cover at Walker Bay from 1996 to 2010. Our results support a growing body of evidence that increased shrub growth does not necessarily translate into a prolonged increase in shrub cover. Instead, we conclude that the heterogeneity of the arctic shrub response to climate change may be associated with variation in the proximate factors limiting recruitment such as water table saturation and herbivory.

The relative importance of sexual reproduction and clonal growth for the long-term maintenance of crowberry populations is poorly understood. Our objective was to characterize reproductive output, recruitment and clonal growth in different populations along a fire chronosequence at the treeline. Our research hypothesis was that sexual reproduction would be more important in younger sites, but that clonal growth would be prevalent in older sites. We quantified cover and reproductive output and reconstructed the establishment chronology of the crowberry populations. The positive relationship between crowberry cover and site age is mainly attributable to the presence of a tree cover at older sites. Fruit production, lower in forested sites, and seed viability were not related to site age. Recruitment of individuals appeared to be relatively similar between sites, while the number of clonal individuals increased with the time elapsed since the last fire. Our results partially support our research hypothesis. While clonal growth is more important for population maintenance at older sites, the species keeps investing in sexual reproduction, even when dominant.

International audience; The microbial respiration of ancient permafrost carbon represents a positive feedback to climate warming. However, warming-induced shrub expansion in circumpolar latitudes may partly offset these emissions, due to greater biomass and litter inputs than that of primary tundra vegetation. Quantifying this carbon sink is challenging as the simultaneous mineralization of ancient carbon renders the attribution of changes in soil carbon stocks uncertain. We measured the contribution of shrubs to the terrestrial carbon reservoir in a Low-Arctic region where ancient carbon stocks are among the lowest in the Arctic. The study site near the eastern shore of Hudson Bay is experiencing rapid Betula glandulosa Michx. expansion throughout lichen tundra. We find that the terrestrial carbon stocks (i.e. soil and vegetation) under a cover of low to medium-size shrubs is increased by 3.9±1.3 kg m −2 , regardless of shrub cover age. Along water tracks, taller shrubs and the transition to moss understories provide an even greater increase in terrestrial carbon (6.5±3.5 kg m −2). Using published maps of vegetation change from 1994 to 2010, we estimate that the carbon sink associated to shrub expansion in our study area (5.228 km 2) has been 2.4±0.8 Gg or 29±9 g m −2 yr −1. Extrapolating this result to the Arctic requires additional studies in representative environments.

Expansion of shrub species is widely reported in northern regions, although its extent varies across the landscape. In subarctic Quebec (Canada), where dwarf birch (Betula glandulosa Michx) is the main species responsible for shrub expansion, little is known about the causes and consequences of this phenomenon. This study aims to identify the drivers of dwarf birch densification heterogeneity at the landscape level and to evaluate its influence on other shrub species. We used model selection with Akaike's information criterion to rank ecologically relevant models including topographic, historical and edaphic variables. The influence of dwarf birch densification was evaluated through regression analysis. We found that the best model explaining the heterogeneity in dwarf birch densification in western Nunavik includes factors related to both historical conditions (initial shrub cover, time elapsed since last wildfire) and topography (type of environment). Among these factors, only the initial shrub...

Summary NDVI analysis and repeated aerial photographs have revealed significant shrub expansion in many subarctic regions. While the recent increase in temperature is usually considered to be the main driver of this phenomenon at regional scales, very little is known about the local heterogeneity of shrub responses across the landscape. In this study, we aim to identify the climatic factors controlling the growth of the largely distributed shrub species Betula glandulosa in three types of environments (terrace, hilltop and snowbed). We also aim to evaluate the relationship between B. glandulosa growth and the NDVI data for the Boniface River region, in north-western Quebec, where the study took place. In the field, we harvested 180 B. glandulosa individuals (20 per site, 3 sites per type of environment). We constructed specific growth-ring width chronologies and mean axial growth rate chronologies for each site and used them for dendroclimatic analysis (response functions). We also used linear regressions to evaluate the relative influence of dwarf birch growth on the NDVI trend. We found a sharp increase in B. glandulosa radial growth in the 1990s followed by a sharp decreasing trend from 2002 on terraces and hilltops, while growth remained high in snowbeds. Betula glandulosa growth was positively correlated with summer temperatures on terraces and hilltops, whereas winter precipitation promoted growth on snowbeds. The NDVI trend was largely correlated to B. glandulosa growth on terraces and hilltops for the period between 1986 and 2002 (71–80% explained variance). Synthesis. Our results suggest that topography plays a major role in B. glandulosa growth and therefore in shrub community dynamics. Because terraces and hilltops represent 70% of the land surface, the sharp B. glandulosa growth increase at these sites promoted an important overall expansion of the shrub community in the region. However, the decline in B. glandulosa growth observed after 2002 suggests that the expansion could be slowed down in the near future, therefore limiting shrub growth contribution to the regional NDVI signal.

Aim Alterations to the abundance and distribution of plants, especially shrubs, have been predicted by warming experiments and confirmed by remote-sensing analyses and land-based observations in the Low Arctic. In the High Arctic, analyses of satellite images suggest a recent increase in the normalized difference vegetation index (NDVI) and warming experiments have shown a positive correlation between temperature and shrub cover, but little is known about the responses of natural systems. In this context, we investigated populations of Arctic willow (Salix arctica Pall.) to assess the potential contribution of this structuring species to High Arctic shrubification. Location Seven polar semidesert sites, including two glacier forelands, in High Arctic Canada and Greenland. Methods We evaluated the size distribution and establishment age of Arctic willow populations through field measurements and dendrochronological analysis. We also assessed the cover of Arctic willow at each site and compared current data with historical records. Results Age determination of over 200 large individuals established by seed suggests that colonization events were associated with the melting of permanent snow and glacial ice. Size distributions showed that recent recruitment was more frequent on glacier forelands, even though seedling establishment was common in other sites. Comparing our measurements with previous studies at the same sites revealed that significant increases in cover over the past few decades occurred only on glacier forelands. Main conclusions Our results suggest that the population structure and dynamics of Arctic willow in the High Arctic are influenced by site-specific factors. Rapid glacier retreat is increasing the extent of the glacier forelands, which are currently experiencing relatively rapid plant colonization. Some sheltered sites with greater snow accumulation will probably experience increased shrub colonization under current and predicted climate change. It appears, however, that shrub recruitment and growth are still strongly limited by water availability, and the reduction in permanent snow cover may exacerbate this stress.

Over the last few decades, shrub species have expanded rapidly in open tundra environments due to climate change. Previous experimental studies in this environment have shown that nutrient addition and, to a lesser extent, warming, had positive effects on shrub growth. However, the response of shrub species in open forested ecosystems such as lichen woodland is still largely unknown. The main objective of this study was to evaluate the performance of Betula glandulosa (Michx., dwarf birch) seedlings subjected to direct (warmer temperature, increased precipitation) and indirect (increased nutrient availability) effects of climate change in a lichen woodland (25 % tree cover). The study took place 10 km south of the subarctic treeline in western Quebec (Canada). At the end of the second growth season, seedling leaf, woody stem and root biomass along with leaf area had increased significantly in response to nutrient addition. Moreover, seedlings exposed to nutrient addition had greater nitrogen, phosphorus and potassium concentrations in their leaves. Warming treatment also had a significant but weaker impact on leaf and woody stem biomass, while increase in precipitation had only a slight impact on seedling root biomass. Our results indicate that B. glandulosa response to simulated changes in the abiotic environment is similar to that observed in open tundra, suggesting that this species could also become more widespread in the forested ecosystems of the forest–tundra ecotone.

Recent climate changes have increased the primary productivity of many Arctic and subarctic regions. Erected shrub has been shown to increase in abundance over the last decades in northern regions in response to warmer climate. At the same time, caribou herds are declining throughout the circumboreal regions. Based on observation of heavy browsing on shrubs at Deception Bay (Nunavik, Canada), we hypothesized that the densification of shrubs observed in nearby locations did not occur at our study site despite of observed warming because of a recent peak of the Riviere-aux-Feuilles caribou herd. To assess shrub cover changes, we compared a 1972 mosaic of aerial photos to a 2010 satellite image over a 5 km2 area, divided into 56 grids of 100 30 m × 30 m cells. Most cells (n = 4,502) did not show any changes in the cover of shrubs but those who did were as likely to increase as to decrease. The relative cover of shrubs in cells who changed was not higher in 2010 (6.1 ± 0.2 %) than in 1972 (7.3 ± 0.4 %). More than 70 % of birch and willow had more than 50 % of their shoot browsed, suggesting that caribou may limit shrub expansion at this site. We cannot rule out that abiotic factors also contribute to the inertia in shrub cover. Increases in shrub abundance reported in Nunavik and elsewhere were located closer to the tree line or in discontinuous permafrost, whereas our site is characterized by herbaceous arctic tundra, continuous permafrost and relatively low annual precipitation.

Logging in tropical forests may create large canopy openings. These gaps provide suitable conditions for some opportunistic shrubs and herbs to take advantage of the surge in resources and rapidly colonize disturbed sites. This dense plant cover may limit forest regeneration by interfering with tree seedling establishment, growth, and survival by altering the light and nutrients available to seedlings, modifying herbivore behavior, or a number of other factors. In Kibale National Park (Uganda), old logging sites are mainly covered by dense stands of Acanthus pubescens Engl., which appear to inhibit tree regeneration. We wanted to identify the ecological processes underlying this regeneration collapse. To do so, we designed a factorial experiment to evaluate the influences of herbivory and vegetation cover on the growth and survival of tree seedlings. We compared the survival and growth of transplanted tree seedlings in A. pubescens stands and logged forests, in the presence or absence of the understory vegetation layer (logged forest) or vegetation cover (A. pubescens), and with or without herbivory. We found no evidence to support the hypothesis that herbivory is significantly higher under dense A. pubescens cover. Seedling survival was not influenced by the environment. Seedling growth, however, was positively influenced by the removal of A. pubescens, suggesting that changes in resource availability associated with the presence of A. pubescens, may be important for regeneration. Our results suggest that sustained cutting of A. pubescens cover could foster the growth of established seedlings and could lead to tree regeneration and habitat restoration.

Shrub densification has been widely reported across the circumpolar arctic and subarctic biomes in recent years. Long-term analyses based on dendrochronological techniques applied to shrubs have linked this phenomenon to climate change. However, the multi-stemmed structure of shrubs makes them difficult to sample and therefore leads to non-uniform sampling protocols among shrub ecologists, who will favor either root collars or stems to conduct dendrochronological analyses. Through a comparative study of the use of root collars and stems of Betula glandulosa, a common North American shrub species, we evaluated the relative sensitivity of each plant part to climate variables, and assessed if this sensitivity is consistent across three different types of environments in northwestern Quebec, Canada (terrace, hilltop and snowbed). We found that root collars had greater sensitivity to climate than stems, and that these differences were maintained across the three types of environment. Growth at the root collar was best explained by spring precipitation and summer temperature, whereas stem growth showed weak and inconsistent responses to climate variables. Moreover, sensitivity to climate was not consistent among plant parts, as individuals having climate sensitive root collars did not tend to have climate sensitive stems. These differences in sensitivity of shrub parts to climate highlight the complexity of resource allocation in multi-stemmed plants. Whereas stem initiation and growth are driven by micro-environmental variables such as light availability and competition, root collars integrate the growth of all plant parts instead, rendering them less affected by mechanisms such as competition and more responsive to signals of global change. Although further investigations are required to determine the degree to which these findings are generalizable across the tundra biome, our results indicate that consistency and caution in the choice of plant parts are a key consideration for the success of future dendroclimatological studies on shrubs. This article is protected by copyright. All rights reserved.

Chemical investigation of the methanol extract of the lichen Stereocaulon paschale collected in Nunavik, Canada, led to the isolation and identification of two new dibenzofurans (1 and 3) and 11 known lichen metabolites. The structures of the new compounds were established by analysis of 1D and 2D NMR spectroscopic and high-resolution mass spectrometric data. Herein, the first isolation of ascomatic acid dibenzofuran derivatives (1-3) from a whole lichen organism is reported. In addition, some of the isolated metabolites showed antibacterial activity against the oral pathogens Porphyromonas gingivalis and Streptococcus mutans.

Complex interactions between plants and soil microorganisms drive key ecosystem and community properties such as productivity and diversity. In nutrient-poor systems such as sand dunes, plant traits and fungal symbioses related to nutrient acquisition can strongly influence vegetation dynamics. We investigated plant and fungal communities in a relic foredune plain located on an archipelago in Quebec, Canada. We detected distinct communities across the edaphic and successional gradient. Our results showed a clear increase in plant species richness, as well as in the diversity of nutrient-acquisition strategies. We also found a strong correlation between aboveground vegetation and soil fungal communities, and both responded similarly to soil physicochemical properties. Soil pH influenced the composition of plant and fungal communities, and could act as an important environmental filter along this relic foredune plain. The increasing functional diversity in plant nutrient-acquisition strategies across the gradient might favor resource partitioning and facilitation among co-occurring plant species. The coordinated changes in soil microbial and plant communities highlight the importance of aboveground-belowground linkages and positive biotic interactions during ecological succession in nutrient-poor environments.

Global vegetation models predict rapid poleward migration of tundra and boreal forest vegetation in response to climate warming. Local plot and air-photo studies have documented recent changes in high-latitude vegetation composition and structure, consistent with warming trends. To bridge these two scales of inference, we analyzed a 24-year (1986-2010) Landsat time series in a latitudinal transect across the boreal forest-tundra biome boundary in northern Quebec province, Canada. This region has experienced rapid warming during both winter and summer months during the last forty years. Using a per-pixel (30 m) trend analysis, 30% of the observable (cloud-free) land area experienced a significant (p < 0.05) positive trend in the Normalized Difference Vegetation Index (NDVI). However, greening trends were not evenly split among cover types. Low shrub and graminoid tundra contributed preferentially to the greening trend, while forested areas were less likely to show significant trends in NDVI. These trends reflect increasing leaf area, rather than an increase in growing season length, because Landsat data were restricted to peak-summer conditions. The average NDVI trend (0.007/yr) corresponds to a leaf-area index (LAI) increase of ~0.6 based on the regional relationship between LAI and NDVI from the Moderate Resolution Spectroradiometer (MODIS). Across the entire transect, the area-averaged LAI increase was ~0.2 during 1986-2010. A higher area-averaged LAI change (~0.3) within the shrub-tundra portion of the transect represents a 20-60% relative increase in LAI during the last two decades. Our Landsat-based analysis subdivides the overall high-latitude greening trend into changes in peak-summer greenness by cover type. Different responses within and among shrub, graminoid, and tree-dominated cover types in this study indicate important fine-scale heterogeneity in vegetation growth. Although our findings are consistent with community shifts in low-biomass vegetation types over multi-decadal time scales, the response in tundra and forest ecosystems to recent warming was not uniform.

Expansion of deciduous shrub species in open subarctic landscapes has been reported at several locations around the circumpolar region. However, few data are available to evaluate the response of shrub species in forested ecosystems. For this study, we conducted a dendrochronological analysis to compare the establishment and growth of Betula glandulosa Michx. individuals prior to and after an experimental removal of tree cover (1987) in a lichen woodland located at the boreal forest – tundra ecotone. Tree removal was followed by a B. glandulosa recruitment pulse. It is likely that the newly established seedlings benefited more from heavy trampling during tree removal, which destroyed the lichen cover, than from tree removal itself. A strong growth increase after tree removal suggests that the black spruce tree cover suppressed B. glandulosa growth prior to 1987. Moreover, while radial growth for the period prior to tree removal was not a function of the number of growing degree-days during the growing season, the same climatic variable explained 46% of the growth variability in the second period. These results suggest that the presence of a tree cover might limit the response of shrub species to climate change in forested ecosystems.

Tree regeneration at treeline is often inhibited by harsh climatic conditions and ecological constraints such as the presence of a continuous lichen cover. The objective of this study was to verify if recent climate warming and increased caribou (Rangifer tarandus L.) activity, which destroys the lichen cover, could act synergistically to increase black spruce (Picea mariana (Mill.) B.S.P.) regeneration. We collected cones produced in 2006–2007 to compare seed viability with data from 1989 to 1995. We conducted experimental plantations (field and controlled conditions) to determine if germination and seedling emergence were higher in disturbed environments where mineral soil was exposed. We sampled naturally established seedlings to evaluate the relationship between the year of establishment and caribou activity and to compare growth in disturbed and undisturbed environments. We found that seed viability was significantly higher in 2006–2007 compared with 1989–1995. The number of germinated seeds per cone increased by a factor >1000. Germination and seedling establishment were higher in the disturbed environment (mineral soil). Finally, although seedling establishment in areas with continuous lichen cover was rather constant over the last 50 years, 73.5% of the seedlings recorded on bare mineral soil became established in the years following high caribou activity. Our results suggest that climate warming and caribou activity are likely to act synergistically to promote black spruce regeneration at treeline.