An experimental test of limits to tree establishment in Arctic tundra

1 Five treeline species had low seed germination rates and low survivorship and growth of seedlings when transplanted into Alaskan tundra. Seed germination of all species increased with experimental warming, suggesting that the present treeline may in part result from unsuccessful recruitment under cold conditions.
2 Growth, biomass and survivorship of seedlings of treeline species transplanted into tundra were largely unaffected by experimental warming. However, transplanted seedlings of three species ( Betula papyrifera , Picea glauca and Populus tremuloides ) grew more when below‐ground competition with the extant community was reduced. All three measures of transplant performance were greater in shrub tundra than in the less productive tussock or heath tundra. Establishment of trees in tundra may thus be prevented by low resource availability and competition.
3 Two species ( Alnus crispa and Populus balsamifera ) had low seed germination and survivorship of germinated seeds; transplants of these species did not respond to the manipulations and lost biomass following transplanting into tundra. Isolated populations of these two species north of the present treeline in arctic Alaska probably became established during mid‐Holocene warming rather than in recent times.
4 Of all the species studied here, Picea glauca was the most likely to invade intact upland tundra. Its seeds had the highest germination rates and it was the only species whose seedlings survived subsequently. Furthermore, transplanted seedlings of Picea glauca had relatively high survivorship and positive growth in tundra, especially in treatments that increased air temperature or nutrient availability, two factors likely to increase with climate warming.     

Reproduction and seedling establishment of Picea glauca across the northernmost forest‐tundra region in Canada

The northern boundary of boreal forest and the ranges of tree species are expected to shift northward in response to climate warming, which will result in a decrease in the albedo of areas currently covered by tundra vegetation, an increase in terrestrial carbon sequestration, and an alteration of biodiversity in the current Low Arctic. Central to the prediction of forest expansion is an increase in the reproductive capacity and establishment of individual trees. We assessed cone production, seed viability, and transplanted seedling success of Picea glauca (Moench.) Voss. (white spruce) in the early 1990s and again in the late 2000s at four forest stand sites and eight tree island sites (clonal populations beyond present treeline) in the Mackenzie Delta region of the Northwest Territories, Canada. Over the past 20 years, average temperatures in this region have increased by 0.9 °C. This area has the northernmost forest‐tundra ecotone in North America and is one of the few circumpolar regions where the northern limit of conifer trees reaches the Arctic Ocean. We found that cone production and seed viability did not change between the two periods of examination and that both variables decreased northward across the forest‐tundra ecotone. Nevertheless, white spruce individuals at the northern limit of the forest‐tundra ecotone produced viable seeds. Furthermore, transplanted seedlings were able to survive in the northernmost sites for 15 years, but there were no signs of natural regeneration. These results indicate that if climatic conditions continue to ameliorate, reproductive output will likely increase, but seedling establishment and forest expansion within the forest‐tundra of this region is unlikely to occur without the availability of suitable recruitment sites. Processes that affect the availability of recruitment sites are likely to be important elsewhere in the circumpolar ecotone, and should be incorporated into models and predictions of climate change and its effects on the northern forest‐tundra ecotone.     

Constraints on treeline advance in a warming climate: a test of the reproduction limitation hypothesis

Aims Several mechanistic hypotheses have been developed to explain the existence of alpine treelines worldwide. The reproduction limitation hypothesis (RLH) postulates that reproductive processes such as seed production and dispersal are restricted in treeline environments, thereby limiting the establishment of new individuals in advance of extant treelines. Despite its popularity, no study has tested this hypothesis in a comprehensive fashion. In this experiment, we attempted to answer whether there are enough viable seeds being dispersed beyond treeline for sexually mediated treeline advance, and what the implications of climate change might be on these processes.Methods We established 30 plots across two aspects (north vs. south) and three elevational habitats (forest, treeline and tundra) in a white spruce (Picea glauca) boreal forest-alpine tundra ecotone in southwest Yukon, Canada. In each plot, tree characteristics, seed production and predispersal damage were measured. Additionally, eight dispersal trays were positioned in each plot to measure seed rain, and germination trials with and without predation exclosures were constructed in a subset of plots to quantify dispersal and germination success.Important findings Results were highly variable both temporally and spatially. In 2014, a mast year, 69% of adult trees produced cones compared to 0.4% in the following year. Higher density of trees in forest plots compared to treeline and tundra resulted in greater seed production at lower elevations. Across all plots, 88% of seeds were damaged before dispersal or were not viable. Treeline plots had significantly greater predispersal damage. Seed rain was greater in south-facing plots than north-facing plots. Less than 2% of seeds produced on the landscape were dispersed into Tundra plots, located 50 m above treeline. There was a net movement of seeds from the north-facing slope to the south at our study site, likely due to prevailing winds during the dispersal period. Germination counts were more than double on north-facing slopes and one-third higher inside exclosures. Cumulatively, the results provide some evidence for the RLH. Collectively, the high amount of predispersal damage and non-viable seeds, variability associated with dispersal and significant seed predation can functionally influence treeline dynamics. These findings suggest that global treeline distribution models, which rely largely on temperature, may not be entirely accurate for predicting treeline advance—at least at finer temporal scales. Many stochastic factors need to align temporally for successful advance, which is likely to result in a lag of many decades between the period of temperature amelioration and an increased number of trees beyond extant treelines.     

Recent climate warming forces contrasting growth responses of white spruce at treeline in Alaska through temperature thresholds

Northern and high‐latitude alpine treelines are generally thought to be limited by available warmth. Most studies of tree‐growth–climate interaction at treeline as well as climate reconstructions using dendrochronology report positive growth response of treeline trees to warmer temperatures. However, population‐wide responses of treeline trees to climate remain largely unexamined. We systematically sampled 1558 white spruce at 13 treeline sites in the Brooks Range and Alaska Range. Our findings of both positive and negative growth responses to climate warming at treeline challenge the widespread assumption that arctic treeline trees grow better with warming climate. High mean temperatures in July decreased the growth of 40% of white spruce at treeline areas in Alaska, whereas warm springs enhance growth of additional 36% of trees and 24% show no significant correlation with climate. Even though these opposing growth responses are present in all sampled sites, their relative proportion varies between sites and there is no overall clear relationship between growth response and landscape position within a site. Growth increases and decreases appear in our sample above specific temperature index values (temperature thresholds), which occurred more frequently in the late 20th century. Contrary to previous findings, temperature explained more variability in radial growth after 1950. Without accounting for these opposite responses and temperature thresholds, climate reconstructions based on ring width will miscalibrate past climate, and biogeochemical and dynamic vegetation models will overestimate carbon uptake and treeline advance under future warming scenarios.     

Limited decadal growth of mountain birch saplings has minor impact on surrounding tundra vegetation

Temperatures over the Arctic region are increasing at three times the rate of the global average. Consequently, Arctic vegetation is changing and trees are encroaching into the tundra. In this study, we examine the establishment and growth of mountain birch (Betula pubescens ssp. tortuosa), which forms the treeline in subarctic Europe, and its impact on community composition across the treeline ecotone nearby Abisko, Sweden. Birch advancement along elevational gradients was studied by comparing data collected in 2016 with data collected 10 and 15 years previously. Species identity, cover, and phylogenetic relatedness were used to assess the impact of birch encroachment on community composition. Our results show that birch occurrence above the treeline did not affect plant community composition, probably owing to the observed lack of significant growth due to herbivore browsing, nitrogen limitation, or a reduction in snow cover. Independent of birch performance, the tundra community structure shifted toward a novel community dissimilar from the forest plant community found below the treeline. Taken together, our findings are explained by species‐specific responses to climate change, rather than by a linear forest advance. Future treeline advancements are likely more restricted than previously expected.     

The interacting effects of temperature,ground disturbance,and herbivory on seedling establishment: implications for treeline advance with climate warming

Thermal control of treeline position is mediated by local environmental and ecological factors, making trends in treeline migration difficult to extrapolate geographically. We investigated the ecological dynamics of conifer establishment at treeline in the Mealy Mountains (Labrador, Canada) and the potential for its expansion with climate warming. Available seedbed and tree seedling emergence in the treeline ecotone were monitored, and seeds and seedlings of Picea mariana were planted along an elevational gradient from open-canopy forest through tree islands to alpine tundra. Experimental treatments included passive warming of daytime air, ground disturbance, and vertebrate herbivore exclosures. Responses in seed germination and seedling growth, damage, and mortality were monitored over two growing seasons, and re-surveyed after 5 years. While no tree seedlings were observed growing naturally above the treeline, planted seeds were able to germinate, develop and overwinter, and persist for 4 years in all habitats examined. Disturbance of the seedbed was important for seedling emergence in the forest and tree islands. While temperature enhancement alone had little impact on emergence, even moderate temperature increases had significantly disproportionate effects on emergence of seedlings in the alpine habitat when combined with soil disturbance, indicating that future climate warming could lead to treeline advance if viable seed and suitable substrate for recruitment are available. The positive effect of excluding herbivores suggests that herbivory may be an important filter modifying future species distribution. While seedbed conditions and herbivory would control the rate of individual species advance, the results indicate potential upslope migration of the treeline in the Mealy Mountains, with consequent loss of alpine ecosystems.     

Pathways of tundra encroachment by trees and tall shrubs in the western Brooks Range of Alaska

Climate change is expected to increase woody vegetation abundance in the Arctic, yet the magnitude, spatial pattern and pathways of change remain uncertain. We compared historical orthophotos photos (1952 and 1979) with high-resolution satellite imagery (2015) to examine six decades of change in abundance of white spruce Picea glauca and tall shrubs (Salix spp., Alnus spp.) near the Agashashok River in northwest Alaska. We established ~3000 random points within our ~5500 ha study area for classification into nine cover types. To examine physiographic controls on tree abundance, we fit multinomial log-linear models with predictors derived from a digital elevation model and with arctic tundra, alpine tundra and ‘tree’ as levels of a categorical response variable. Between 1952 and 2015, points classified as arctic and alpine tundra decreased by 31% and 15%, respectively. Meanwhile, tall shrubs increased by 86%, trees mixed with tall shrubs increased by 385% and forest increased by 84%. Tundra with tall shrubs rarely transitioned to forest. The best multinomial model explained 71% of variation in cover and included elevation, slope and an interaction between slope and ‘northness’. Treeline was defined as the elevation where the probability of tree presence equaled that of tundra. Mean treeline elevation in 2015 was 202 m, corresponding with a June–August mean air temperature > 11°C, which is > 4°C warmer than the 6–7°C isotherm associated with global treeline elevations. Our results show dramatic increases in the abundance of trees and tall shrubs, question the universality of air temperature as a predictor of treeline elevation and suggest two mutually exclusive pathways of vegetation change, because tundra that gained tall shrubs rarely transitioned to forest. Conversion of tundra to tall shrubs and forest has important and potentially contrasting implications for carbon cycling, surface energy exchange and wildlife habitat in the Arctic.     

Snow distribution, soil temperature and late winter CO2 efflux from soils near the Arctic treeline in northwest Alaska

The Arctic treeline is advancing in many areas and changes in carbon (C) cycling are anticipated. Differences in CO₂ exchange between adjacent forest and tundra are not well known and contrasting conclusions have been drawn about the effects of forest advance on ecosystem C stocks. Measurements of CO₂ exchange in tundra and adjacent forest showed the forest was a greater C sink during the growing season in northern Canada. There is, however, reason to expect that forests lose more C than tundra during the wintertime, as forests may accumulate and retain more snow. Deeper snow insulates the soil and warmer soils should lead to greater rates of belowground respiration and CO₂ efflux. In this study, I tested the hypotheses that forests maintain a deeper snowpack, have warmer soils and lose more C during winter than adjacent tundra near the Arctic treeline in northwest Alaska. Measurements of snow depth, soil temperature and CO₂ efflux were made at five forest and two treeline sites in late winter of three consecutive years. Snow depth and soil temperature were greater in forest than treeline sites, particularly in years with higher snowfall. There was a close exponential correlation between soil temperature and CO₂ efflux across sites and years. The temperature-efflux model was driven using hourly soil temperatures from all the sites to provide a first approximation of the difference in winter C loss between treeline and forest sites. Results showed that greater wintertime C loss from forests could offset greater summertime C gain.     

Shrub–tree interactions and environmental changes drive treeline dynamics in the Subarctic

Treelines have drawn persistent research interest as they can respond markedly to climate. However, the mechanisms that determine tree seedling recruitment and the response of the forest‐tundra ecotone to environmental changes remain poorly understood. We hypothesise that treeline tree seedling performance depends on the interplay between climatic and soil nutritional changes and facilitative and competitive interactions between trees and shrubs. We conducted a seedling transplantation experiment with Betula pubescens at a subarctic treeline, in northern Sweden, which followed a full factorial design with four treatment factors relating to environmental regimes of stress and resource availability: site (forest vs treeline); temperature (+/? passive warming); shrub presence (+/?Vaccinium myrtillus removal); and nutrient availability (+/? NPK addition). During three growing seasons we assessed the establishment and performance of Betula. The experimental manipulations caused highly significant effects on seedling performance. Although Vaccinium enhanced seedling survival and reduced the effects of excessive solar radiation and insect herbivory, the seedlings growing with the shrub had a poorer performance by the end of the experimental period. Also, seedlings in the forest had a poorer performance than those at the treeline. Betula seedlings showed a very pronounced and positive response to passive warming and to nutrient addition, but such effects were more evident at the treeline site and often interacted with the presence of Vaccinium. This experiment shows that shrub–tree interactions are important drivers of subarctic treeline dynamics and that they vary with time and space. Facilitation, competition, herbivory and environmental changes at the tree seedling stage act as important filters in structuring the forest–tundra ecotone. We demonstrate that changes in this ecotone cannot be simply predicted from changing temperature patterns alone, and that complex interactions need to be considered, not only between shrubs and trees, but also with herbivores and between warming and soil nutrient availability.     

DEMOGRAPHIC ASPECTS OF COEXISTENCE IN ENGELMANN SPRUCE AND SUBALPINE FIR

The mechanisms for the maintenance of coexistence of Engelmann spruce and subalpine fir in subalpine forests of the Colorado Front Range were examined by comparing age, size, and spatial distributions of spruce and fir in two adjacent, previously logged sites of differing moisture availability. Adult tree ages were calculated from stem cores, while seedling ages were calculated from a multiple regression equation based on diameter, height, and number of branch whorls. Tree size was measured by height and diameter; spatial distributions were described by Morisita's index of dispersion. Cumulative age and size distributions were significantly different in the two species, with greater longevity and a larger overall size in spruce than fir. Both species showed a significant linear relationship between size and age, while fir showed a faster height growth rate than spruce. The linear relationship beween age and size was much closer in seedlings than in adults. Seedling spatial distribution was highly clumped in both species, but mature trees showed little or no clumping. Because both species are mainly wind dispersed, the greater clumping in spruce than in fir seedlings suggests that spruce have more specific establishment requirements than fir. Colonization patterns indicated that spruce seedlings were primarily found in forest gaps or associated with fir canopy trees, while fir seedlings were more commonly found in the forest, associated with either spruce or fir canopy trees. Tree density, growth rates, and mortality rates were higher in the wet site, with spruce showing the largest between site differences. These data suggest a new hypothesis for coexistence stating that Engelmann spruce and subalpine fir are maintained as codominants because the greater longevity and size of spruce is balanced by the faster height growth and more flexible seedling establishment requirements of fir.     

Variation in carbohydrate source–sink relations of forest and treeline white spruce in southern, interior and northern Alaska

Two opposing hypotheses have been presented to explain reduced tree growth at the treeline, compared with growth in lower elevation or lower latitude forests: the carbon source and sink limitation hypotheses. The former states that treeline trees have an unfavorable carbon balance and cannot support growth of the magnitude observed at lower elevations or latitudes, while the latter argues that treeline trees have an adequate carbon supply, but that cold temperatures directly limit growth. In this study, we examined the relative importance of source and sink limitation in forest and treeline white spruce (Picea glauca) in three mountain ranges from southern to northern Alaska. We related seasonal changes in needle nonstructural carbohydrate (NSC) content with branch extension growth, an approach we argue is more powerful than using needle NSC concentration. Branch extension growth in the southernmost Chugach Mountains was much greater than in the White Mountains and the Brooks Range. Trees in the Chugach Mountains showed a greater seasonal decline in needle NSC content than trees in the other mountain ranges, and the seasonal change in NSC was correlated with site-level branch growth across mountain ranges. There was no evidence of a consistent difference in branch growth between the forest and treeline sites, which differ in elevation by approximately 100 m. Our results point to a continuum between source and sink limitation of growth, with high-elevation trees in northern and interior Alaska showing greater evidence of sink limitation, and those in southern Alaska showing greater potential for source limitation.     

Reproduction as a bottleneck to treeline advance across the circumarctic forest tundra ecotone

The fundamental niche of many species is shifting with climate change, especially in sub‐arctic ecosystems with pronounced recent warming. Ongoing warming in sub‐arctic regions should lessen environmental constraints on tree growth and reproduction, leading to increased success of trees colonising tundra. Nevertheless, variable responses of treeline ecotones have been documented in association with warming temperatures. One explanation for time lags between increasingly favourable environmental conditions and treeline ecotone movement is reproductive limitations caused by low seed availability. Our objective was to assess the reproductive constraints of the dominant tree species at the treeline ecotone in the circumpolar north. We sampled reproductive structures of trees (cones and catkins) and stand attributes across circumarctic treeline ecotones. We used generalized linear mixed models to estimate the sensitivity of seed production and the availability of viable seed to regional climate, stand structure, and species‐specific characteristics. Both seed production and viability of available seed were strongly driven by specific, sequential seasonal climatic conditions, but in different ways. Seed production was greatest when growing seasons with more growing degree days coincided with years with high precipitation. Two consecutive years with more growing degree days and low precipitation resulted in low seed production. Seasonal climate effects on the viability of available seed depended on the physical characteristics of the reproductive structures. Large‐coned and ‐seeded species take more time to develop mature embryos and were therefore more sensitive to increases in growing degree days in the year of flowering and embryo development. Our findings suggest that both moisture stress and abbreviated growing seasons can have a notable negative influence on the production and viability of available seed at treeline. Our synthesis revealed that constraints on predispersal reproduction within the treeline ecotone might create a considerable time lag for range expansion of tree populations into tundra ecosystems.     

Multi-scale Analysis of Environmental Conditions and Conifer Seedling Distribution Across the Treeline Ecotone of Northern Manitoba, Canada

Treeline represents not only an important physiognomic boundary but also an important transition between disjunct mesoclimates and environmental limitations on establishment of tree species. The circumboreal treeline is controlled by some still to be understood physiological mechanism dependent on air temperatures, though younger life stages are increasingly influenced by numerous other biotic and abiotic factors at finer spatial and temporal scales. The goal of this study was to evaluate environmental and reproductive characteristics across treeline around Churchill, Manitoba, and to determine which factors are most important for successful seedling establishment by tamarack, white spruce and black spruce. We examined mid-winter snowpack, soil characteristics, seed viability, seedling establishment, and dominant vegetation at sites within forest and at treeline. Growing season was longer at treeline due to less snow accumulation, though soil temperatures were more variable throughout the year when compared with forest. Conifer seed germination was greater than 88% for most of the region and total seedling density was relatively consistent between sites. Seedlings were negatively associated with other plants within the forest, but low stature vegetation seemed to facilitate establishment at treeline. The associations between seedling establishment and habitat availability observed at several sites suggest that treeline advance in the Churchill area could be contingent on the facilitative effects of plants at and beyond treeline. The results of this study support the premise that fine-scale biotic and abiotic patterns and processes such as snowpack and facilitation by neighboring vegetation certainly cannot be overlooked in analyses of patterns at treeline in a changing Subarctic.     

The nature of spatial transitions in the Arctic

Aim Describe the spatial and temporal properties of transitions in the Arctic and develop a conceptual understanding of the nature of these spatial transitions in the face of directional environmental change. Location Arctic tundra ecosystems of the North Slope of Alaska and the tundra‐forest region of the Seward Peninsula, Alaska Methods We synthesize information from numerous studies on tundra and treeline ecosystems in an effort to document the spatial changes that occur across four arctic transitions. These transitions are: (i) the transition between High‐Arctic and Low‐Arctic systems, (ii) the transition between moist non‐acidic tundra (MNT) and moist acidic tundra (MAT, also referred to as tussock tundra), (iii) the transition between tussock tundra and shrub tundra, (iv) the transition between tundra and forested systems. By documenting the nature of these spatial transitions, in terms of their environmental controls and vegetation patterns, we develop a conceptual model of temporal dynamics of arctic ecotones in response to environmental change. Results Our observations suggest that each transition is sensitive to a unique combination of controlling factors. The transition between High and Low Arctic is sensitive primarily to climate, whereas the MNT/MAT transition is also controlled by soil parent material, permafrost and hydrology. The tussock/shrub tundra transition appears to be responsive to several factors, including climate, topography and hydrology. Finally, the tundra/forest boundary responds primarily to climate and to climatically associated changes in permafrost. There were also important differences in the demography and distribution of the dominant plant species across the four vegetation transitions. The shrubs that characterize the tussock/shrub transition can achieve dominance potentially within a decade, whereas spruce trees often require several decades to centuries to achieve dominance within tundra, and Sphagnum moss colonization of non‐acidic sites at the MNT/MAT boundary may require centuries to millennia of soil development. Main conclusions We suggest that vegetation will respond most rapidly to climatic change when (i) the vegetation transition correlates more strongly with climate than with other environmental variables, (ii) dominant species exhibit gradual changes in abundance across spatial transitions, and/or (iii) the dominant species have demographic properties that allow rapid increases in abundance following climatic shifts. All three of these properties characterize the transition between tussock tundra and low shrub tundra. It is therefore not surprising that of the four transitions studied this is the one that appears to be responding most rapidly to climatic warming.     

Altitudinal variations in seedling and sapling density and age structure of timberline tree species in the Sergyemla Mountains, southeast Tibet

The growth situation of seedlings of treeline species can be used as an indicator for the upper or lower shift of treeline under global climate change, since any advance of a treeline would require seedling survival above the current treeline elevation. We investigated seedling and sapling density of two dominant conifers Abies georgei var. smithii and Sabina saltuaria at lower, middle and upper altitude of two contrasting slopes in Sergyemla Mountain, southeast Tibet. Seedling and sapling age structure of A. georgei var. smithii was analyzed further. Seedling density of both species, as well as sapling density of S. saltuaria, showed a declining trend as elevation increased. Sapling density of A. georgei var. smithii was somewhat higher at middle altitude than other positions. The frequency of old-aged A. georgei var. smithii seedlings/saplings (age >10 year) was the lowest at the highest altitude and tended to increase with the declining elevation. The results indicate that the higher density of seedling and sapling in high-altitude might be attributed to better soil moisture availability and solar insolation than in the mid- and low-altitudes during the growth season. Yet confined by other environmental factors, A. georgei var. smithii seedlings in high-altitude suffer from lower survival rate and lower possibility to grow tall compared with those in mid- and low-altitudes.     

Growth and tillering patterns within tussocks of Eriophorum vaginatum

Growth patterns were investigated for Eriophorum vaginatum tussocks from disturbed and undisturbed tussock tundra at two sites in Alaska. Total basal area of tussocks decreased with increased cryoturbation but mean basal area per tussock did not. Flowering was observed in tussocks of significantly smaller size on disturbed compared to undisturbed tundra. For tussocks with < 10% cover by shrubs and moss, number of tillers per tussock was linearly related to tussock diameter in most disturbed and undisturbed sites. Exceptions occurred in an area that had been bladed with a bulldozer 7 yr before our survey where tiller number increased as the square of diameter and in an area with much frost activity where tiller number was not emulated with diameter. The ratio of daughter tillers to adult tillers decreased with diameter in disturbed tundra, whereas the trend was less pronounced in undisturbed tundra.
Microsuccession in undisturbed tussock tundra was investigated by sampling tussocks with different amounts of shrub and moss cover for number of daughter tillers per adult tiller, weight per tiller, percent nitrogen, and percent phosphorus. Small tussocks without cover by other species and large, partially covered tussocks were not significantly different by any measures, but tussocks that were almost completely buried had significantly (p < 0.05) lower values of tillering index, weight per tiller, and percent phosphorus.     

Response of Betula glandulosa seedlings to simulated increases in nutrient availability,temperature and precipitation in a lichen woodland at the forest–tundra ecotone

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 Québec (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.     

Altitudinal variations in seedling and sapling density and age structure of timberline tree species in the Sergyemla Mountains, southeast Tibet

The growth situation of seedlings of treeline species can be used as an indicator for the upper or lower shift of treeline under global climate change, since any advance of a treeline would require seedling survival above the current treeline elevation. We investigated seedling and sapling density of two dominant conifers Abies georgei var. smithii and Sabina saltuaria at lower, middle and upper altitude of two contrasting slopes in Sergyemla Mountain, southeast Tibet. Seedling and sapling age structure of A. georgei var. smithii was analyzed further. Seedling density of both species, as well as sapling density of S. saltuaria, showed a declining trend as elevation increased. Sapling density of A. georgei var. smithii was somewhat higher at middle altitude than other positions. The frequency of old-aged A. georgei var. smithii seedlings/saplings (age >10 year) was the lowest at the highest altitude and tended to increase with the declining elevation. The results indicate that the higher density of seedling and sapling in high-altitude might be attributed to better soil moisture availability and solar insolation than in the mid- and low-altitudes during the growth season. Yet confined by other environmental factors, A. georgei var. smithii seedlings in high-altitude suffer from lower survival rate and lower possibility to grow tall compared with those in mid- and low-altitudes.     

Recent advance of white spruce (Picea glauca) in the coastal tundra of the eastern shore of Hudson Bay (Québec, Canada)

Aim The species‐specific response of tree‐line species to climatic forcing is a crucial topic in modelling climate‐driven ecosystem dynamics. In northern Québec, Canada, black spruce (Picea mariana) is the dominant species at the tree line, but white spruce (Picea glauca) also occurs along the maritime coast of Hudson Bay, and is expanding along the coast and on lands that have recently emerged because of isostatic uplift. Here we outline the present distribution, structure, dynamics and recent spread of white spruce from the tree line up to its northernmost position in the shrub tundra along the Hudson Bay coast. We aimed to obtain a minimum date of the arrival of the species in the area and to evaluate its dynamics relative to recent climate changes. Location White spruce populations and individuals were sampled along a latitudinal transect from the tree line to the northernmost individual in the shrub tundra along the Hudson Bay coast and in the Nastapoka archipelago in northern Québec and Nunavut, Canada (56°06′–56°32′ N). Methods White spruce populations were mapped, and the position, dimension, growth form and origin (seed or layering) of every individual recorded. Tree‐ring analyses of living and dead trees allowed an estimation of the population structure, past recruitment, growth trends and growth rate of the species. A macrofossil analysis was performed of the organic horizon of the northernmost white spruce stands and individuals. Radiocarbon dates of white spruce remains and organic matter were obtained. The rate of isostatic uplift was assessed by radiocarbon dating of drifted wood fragments. Results The first recorded establishment of white spruce was almost synchronous at all sites and occurred around ad 1660. Spruce recruitment was rather continuous at the tree line, while it showed a gap in the northern shrub tundra during the first decades of the 19th century. A vigorous, recent establishment of seedlings was observed in the shrub tundra; only wind‐exposed, low krummholz (stunted individuals) did not show any sexual regeneration. A period of suppressed growth occurred from the 1810s to the 1850s in most sites. A growth increase was evident from the second half of the 19th century and peaked in the 1880s and the 20th century. A shift from stunted to tree growth form has occurred since the mid‐19th century. No sample associated with white spruce remains gave a date older than 300 ¹⁴C years bp [calibrated age (cal.) ad 1430–1690]. Main conclusions White spruce probably arrived recently in the coastal tundra of Hudson Bay due to a delayed post‐glacial spread. The arrival of the species probably occurred during the Little Ice Age. The established individuals survived by layering during unfavourable periods, but acted as nuclei for sexual recruitment almost continuously, except in the northernmost and most exposed sites. Warmer periods were marked by strong seedling recruitment and a shift to tree growth form. Unlike white spruce, black spruce showed no evidence of an ongoing change in growth form and sexual recruitment. Ecological requirements and recent history of tree‐line species should be taken into account in order to understand the present dynamics of high‐latitude ecosystems.     

Carbon dynamics at frost-patterned tundra driven by long-term vegetation change rather than by short-term non-growing season warming

Frost-patterned grounds, such as mostly barren frost boils surrounded by denser vegetation, are typical habitat mosaics in tundra. Plant and microbial processes in these habitats may be susceptible to short-term warming outside the growing season, while the areal cover of barren frost boils has decreased during the past decades due to climate warming-induced shrub expansion. The relative importance of such short-term and long-term climate impacts on carbon (C) dynamics remains unknown. We measured ecosystem CO₂ uptake and release (in the field), microbial respiration (in the laboratory), as well as microbial biomass N and soil extractable N in frost boils and the directly adjacent heath in late spring and late summer. These habitats had been experimentally warmed with insulating fleeces from late September until late May for three consecutive years, which allowed us to investigate the direct short-term effects of warming and longer-term, indirect climate effects via vegetation establishment into frost boils. Non-growing season warming increased C uptake at the frost boils in late spring and decreased it in late summer, while the timing and direction of responses was opposite for the heath. Experimental warming had no effects on microbial or ecosystem C release or soil N at either of the habitats. However, C cycling was manifold higher at the heath compared to the frost boils, likely because of a higher SOM stock in the soil. Short-term climate change can thus directly alter ecosystem C uptake at frost-patterned grounds but will most likely not affect microbial C release. We conclude that the C dynamics at frost-patterned grounds under a changing climate depend most strongly on the potential of vegetation to encroach into frost boils in the long-term.