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.     

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.     

Subarctic and alpine tree line dynamics during the last 400 years in north‐western and central Canada

Aim The objectives of this study were to: (1) identify episodes of establishment and mortality of young and mature trees at several sites at the alpine tree line in the western Northwest Territories and the latitudinal tree line in northern Manitoba; (2) infer changes in the structure and location of the tree line from patterns of establishment; (3) evaluate any relationship between these changes and climate; and (4) investigate sources of variability between sampling sites and study areas. Location Taiga Cordillera of the western Mackenzie Mountains in the Northwest Territories, and the western Hudson Bay Lowlands in northern Manitoba, Canada. Methods Recent tree line dynamics were examined at six climatically similar sites: three in the western Mackenzie Mountains and three around Churchill, Manitoba. Dendroecological techniques were employed to construct static age distributions of species present at each site. Static age structures, residuals from modelled age distributions, and reconstructions of dynamic stand density were used to identify patterns of establishment and mortality and to compare these to changes in climate. Results Tree line locations advanced and stand density increased during the early‐to‐mid‐20th century around Churchill, although responses were not uniform across sites or species. Results were less conclusive in the Mackenzie Mountains, although the tree line probably advanced during the late 18th century, and stand infilling occurred during the mid‐20th century. Correlation analyses with temperature suggest that conditions during establishment and particularly during recruitment are crucial for controlling tree line dynamics. Main conclusions Tree line advance and stand infilling have continued to the present at Churchill, while the tree line has stagnated in the western Mackenzie Mountains. The results of this study indicate that site‐ and species‐specific responses play a large role in determining the tree line response at multiple scales, illustrating the complexity of tree line dynamics in the context of a changing climate.     

Primary succession of subarctic vegetation and soil on the fast‐rising coast of eastern Hudson Bay,Canada

Aim The objectives of the study are: (1) to evaluate the dynamics of the maritime tree line and forest limit of white spruce, Picea glauca, within the dual framework of primary succession induced by the rapid post‐glacial land emergence on the eastern coast of Hudson Bay and the impacts of recent and past climate changes; and (2) to determine the time lapse between land emergence and seedling, tree, and forest establishment in the context of the primary chronosequence occurring on rising, well‐drained sandy beaches and terraces. Location The study area was located on the eastern coast of Hudson Bay (56°20′ N, 76°32′ W) in northern Québec, Canada. Methods We evaluated the colonization dynamics of white spruce as seedlings, tree‐line trees and primary‐forest trees at eight sites distributed along a 200‐km latitudinal gradient based on a mean land emergence rate of 1.2 m century^?1. A 30‐m wide by 140–300‐m long quadrat was positioned at random at the centre of each site. The elevation above sea level, position and age of all individuals of spruce present in the quadrat areas were determined, and the soils of each chronosequence were described. Results The main stages of primary succession along the emerging coast were common to all the sites, regardless of latitude, but occurred at different elevations above sea level (a.s.l.). White spruce seedlings colonized near‐shore beaches 2 m a.s.l., whereas the tree line and forest limit tended to form only at about 3–4 m and 4–8 m a.s.l., corresponding approximately to 180–825 years and 310–1615 years after land emersion, respectively. White spruce establishment at the tree line occurred about 50 years ago. Climatic conditions at this time were probably more favourable to tree colonization than when the species established at the forest limit. Soil formation was influenced primarily by distance from the seashore and elevation above sea level, with podzolization being accelerated by white spruce cover. Main conclusions The current tree‐line and forest‐limit positions on the rising coast of eastern Hudson Bay correspond to ecological limits established during the course of primary succession within a context of changing climatic conditions. The recent establishment of trees at the tree line and forest limit at relatively old coastal sites is associated with warmer conditions over the last 100 years. Although white spruce was present nearby, coastal sites were devoid of trees before the 20th century.     

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.     

Cytochrome and alternative pathway respiration in white spruce (Picea glauca) roots. Effects of growth and measurement temperature

Interactions between growth temperature and measurement temperature were examined for their effects on white spruce [ Picea glauca (Moench) Voss] root respiration. Total dark respiration rates increased with measurement temperature and were unaffected by growth temperature. Partitioning of respiratory electron flow between the cytochrome and alternative pathways was also unaffected by growth temperature. The proportion of respiration mediated by the alternative pathway was constant at measurement temperatures between 4°C and 18°C, but was increased at higher temperatures. Changes in alternative pathway activity were paralleled by changes in capacity, and the alternative pathway was almost fully engaged at all temperatures. Roots grown at low temperature displayed higher carbohydrate levels than roots grown at higher temperatures, but respiration rate was unaffected. Spruce root respiration did not appear to acclimate to growth temperature, and the alternative pathway was not preferentially engaged at low temperature.     

Effects of triadimefon and osmotic stress on plasma membrane composition and ATPase activity in white spruce (Picea glauca) needles

White spruce [ Picea glauca (Moench) Voss.] seedlings were grown in solution culture and treated with 20 mg I^-1 triadimefon [1-(chlorophenoxy)-3,3-dimethyl-1-(1,2,4-triazol-1-yl)-2-butanol] for 4 weeks and then subjected to osmotic stress with polyethylene glycol 3350. Water potentials and electrolyte leakage were measured in control and triadimefon-treated seedlings before and after the plants were subjected to osmotic stress. The plasma membranes were isolated from needles to study their lipid composition and the activity of plasma-membrane bound ATPase. Triadimefon treatment reduced water potentials and increased leakage of electrolytes from seedlings. However, when the seedlings were exposed to osmotic stress, triadimefon-treated plants maintained higher water potentials and leaked less electrolytes compared with the control plants. Both triadimefon and osmotic stress treatments inhibited the activity of plasma membrane-bound ATPase and altered the composition of free sterols in the plasma membranes. Triadimefon-treated plants contained traces of campesterol, which was not present in control. Osmotic stress drastically reduced phospholipid:protein and sterol:protein ratios and increased sterol:phospholipid ratios in the plasma membranes     

Conifer seedling recruitment across a gradient from forest to alpine tundra: effects of species,provenance, and site

Background: Seedling germination and survival is a critical control on forest ecosystem boundaries, such as at the alpine–treeline ecotone. In addition, while it is known that species respond individualistically to the same suite of environmental drivers, the potential additional effect of local adaptation on seedling success has not been evaluated.

Aims: To determine whether local adaptation may influence the position and movement of forest ecosystem boundaries, we quantified conifer seedling recruitment in common gardens across a subalpine forest to alpine tundra gradient at Niwot Ridge, Colorado, USA.

Methods: We studied Pinus flexilis and Picea engelmannii grown from seed collected locally at High (3400 m a.s.l.) and Low (3060 m a.s.l.) elevations. We monitored emergence and survival of seeds sown directly into plots and survival of seedlings germinated indoors and transplanted after snowmelt.

Results: Emergence and survival through the first growing season was greater for P. flexilis than P. engelmannii and for Low compared with High provenances. Yet survival through the second growing season was similar for both species and provenances. Seedling emergence and survival tended to be greatest in the subalpine forest and lowest in the alpine tundra. Survival was greater for transplants than for field-germinated seedlings.

Conclusions: These results suggest that survival through the first few weeks is critical to the establishment of natural germinants. In addition, even small distances between seed sources can have a significant effect on early demographic performance – a factor that has rarely been considered in previous studies of tree recruitment and species range shifts.     

高山林线变化的更新受限机制研究进展

全球林线位置对气候变暖的响应表现为上升、无变化或下降等截然不同趋势,表明影响林线位置及动态的因子十分复杂,除了较普遍认为的低温调控机制外,还存在其它控制林线位置变化的机制。林线向上迁移开始于种子向林线以上的传播及幼苗在林线以上的定居,这些过程中的限制因子均会影响林线的位移,因此研究更新过程及其限制因子对理解高山林线对气候变化的响应具有重要的科学意义。主要从种子和幼苗两个关键阶段综述高山林线森林更新的研究进展。在种子阶段,夏季积温不足导致种子产量和活力下降,风速过低和浓密灌丛限制种子向林线以上传播,近地表的霜冻/水分胁迫和灌木释放的化感物质会阻碍种子在林线以上萌发。在幼苗阶段,除冬季低温外,生长季内较大的温度日振幅和偶然出现的冻害事件也是导致幼苗死亡的重要原因,而低温环境下的强烈光照引起的低温光抑制会显著降低生长季的光合作用;土壤低温、由土壤温度昼夜变化引起的冻举事件、夏季土壤干旱可能会导致幼苗光合作用下降和死亡率上升;积雪太浅会导致生长季早期幼苗水分供应的严重缺乏,但积雪太深会导致幼苗感染真菌的可能性增加;浓密的灌木和草本植物以及植食动物的啃食也会降低林线以上的幼苗存活率。气候变暖对林线幼苗定居的影响复杂且具有很大不确定性,需要进一步研究气候变暖导致的环境因子变化对林线更新各关键阶段的影响。未来气候变暖无疑会导致生长季起始日提前,结束日推迟,这很可能会增加生长季期间尤其是早期的低温冻害事件,对高山林线树种幼苗的存活具有重要影响。在未来研究中,需要找出定义生长季冻害事件的温度阈值,利用长期气象观测数据分析增温背景下生长季早期冻害事件特征的变化趋势,并进一步开展野外模拟增温实验以深刻理解林线树种的种子萌发和幼苗定居与生长季冻害事件的关系,加强对不同地区林线树种的繁殖策略研究,这将有助于人们进一步理解不同区域林线的形成机制并预测未来气候变化条件下林线的动态变化趋势。     

Early effects of triadimefon on water relations, sterol composition and plasma membrane ATPase in white spruce (Picea glauca) seedlings

Seedlings of white spruce ( Picea glauca [Moench] Voss.) were treated with triadimefon solution applied to the soil, and their early responses studied from 12 h to 7 days after treatment. Transpiration rates declined and respiration rates increased immediately after the commencement of triadimefon treatment. Photosynthetic rates declined less than transpiration rates, resulting in an increase in water use efficiency, whereas root and shoot water potentials remained unchanged during the first 5 days of triadimefon treatment. Triadimefon decreased root hydraulic conductivity and inhibited the activity of the plasma membrane ATPase. In addition, triadimefon-treated roots drastically increased the ratios between free sterols and sterol esters and decreased the ratios between sterol esters and acylated sterol glycosides.     

Critical periods for impact of climate warming on early seedling establishment in subarctic tundra

Climate warming is expected to shift bioclimatic zones and plant species distribution. Yet, few studies have explored whether seedling establishment is a possible bottleneck for future migration and population resilience. We test how warming affects the early stages of seedling establishment in 10 plant species in subarctic tundra. To zoom into the life phases where the effects of warming actually take place, we used a novel approach of breaking down the whole‐season warming effect into full factorial combination of early‐, mid‐, and late‐season warming periods. Seeds were sown in containers placed under field conditions in subarctic heath and were exposed to 3 °C elevation of surface temperature and 30% addition of summer precipitation relative to ambient. Heating was achieved with Free Air Temperature Increase systems. Whole‐season heating reduced germination and establishment, significantly in four out of 10 species. The whole‐season warming effect originated from additive effects of individual periods, although some of the periods had disproportionally stronger influence. Early‐germinating species were susceptible to warming; the critical phases were early summer for germination and mid summer for seedling survival. Graminoids, which emerged later, were less susceptible although some negative effects during late summer were observed. Some species with intermediate germination time were affected by all periods of warming. Addition of water generally could not mitigate the negative effects of whole‐season heating, but at individual species level both strengthening and amelioration of these negative effects were observed. We conclude that summer warming is likely to constrain seedling recruitment in open micro sites, which is a common seed regeneration niche in tundra ecosystem. Importantly, we described both significant temporal and species‐specific variation in the sensitivity of seedling establishment to warming which needs to be taken into consideration when modelling population dynamics and vegetation transitions in a warmer climate.     

Changes in high arctic tundra plant reproduction in response to long‐term experimental warming

We provide new information on changes in tundra plant sexual reproduction in response to long‐term (12 years) experimental warming in the High Arctic. Open‐top chambers (OTCs) were used to increase growing season temperatures by 1–2 °C across a range of vascular plant communities. The warming enhanced reproductive effort and success in most species; shrubs and graminoids appeared to be more responsive than forbs. We found that the measured effects of warming on sexual reproduction were more consistently positive and to a greater degree in polar oasis compared with polar semidesert vascular plant communities. Our findings support predictions that long‐term warming in the High Arctic will likely enhance sexual reproduction in tundra plants, which could lead to an increase in plant cover. Greater abundance of vegetation has implications for primary consumers – via increased forage availability, and the global carbon budget – as a function of changes in permafrost and vegetation acting as a carbon sink. Enhanced sexual reproduction in Arctic vascular plants may lead to increased genetic variability of offspring, and consequently improved chances of survival in a changing environment. Our findings also indicate that with future warming, polar oases may play an important role as a seed source to the surrounding polar desert landscape.     

Spatio-temporal variability and environmental controls of methane fluxes at the forest–tundra ecotone in the Fennoscandian mountains

We report on temporal and spatial variability in net methane (CH₄) fluxes measured during the thaw period of 1999 and 2000 at three study sites along a c. 8° latitudinal gradient in the Fennoscandian mountain range and across the mountain birch‐tundra ecotone. All of the sites studied here were underlain by well‐drained mesic soils. In addition, we conducted warming experiments in the field to simulate future climate change. Our results show significant CH₄ uptake at mesic sites spanning the forest‐tundra ecotone: on average 0.031 and 0.0065 mg CH₄ m^?2 h^?1 during the 1999 and 2000 thaw periods, respectively, in Abisko (Sweden), and 0.019 and 0.032 mg CH₄ m^?2 h^?1 during 2000 in Dovrefjell and Joatka (Norway), respectively. These values were both temporally and spatially highly variable, and multiple regression analysis of data from Abisko showed no consistent relationship with soil‐moisture status and temperature. Also, there was no consistent difference in CH₄ fluxes between forest and tundra plots; our data, therefore, provide no support for the hypothesis that conversion of tundra to mountain birch forest, or vice versa, would result in a systematic change in the magnitude or direction of net CH₄ fluxes in this region. Experimental warming treatments were associated with a 2.4 °C increase in soil temperatures (5 cm depth) in 1999 in Abisko, but no consistent soil warming was noted at any of the three field locations during 2000. In spite of this, there were significant treatment effects, principally early during the thaw period, with increased CH₄ uptake compared with control (ambient) plots. These results suggest that direct effects of air warming on vegetation processes (e.g. transpiration, root exudation and nutrient assimilation) can influence CH₄ fluxes even in predominantly methanotrophic environments. We conclude that net CH₄ oxidation is significant in these cold, mesic soils and could be strengthened in an environmental change scenario involving a combination of (i) an increase in the length of the thaw period and (ii) increased mean temperatures during this period in combination with decreased soil‐moisture content.     

Episodic recruitment of the seedling banks in balsam fir and white spruce

• Premise of the study: In ecosystems where seed production is low and masting years are sporadic, or with species that have short-lived seeds, regeneration is assured by seedling banks rather than seed banks. Seedling establishment and survival play a critical role in determining the composition of these plant communities by supplying new individuals for their maintenance. Seedling emergence and mortality were investigated to test the hypothesis that recruitment into the seedling bank is periodic. • Materials and methods: Seed production and seedling emergence and survival was monitored during 1994-2007 in balsam fir (Abies balsamea) and white spruce (Picea glauca) in four pristine stands of the boreal forest of Quebec, Canada. Measurements were collected twice per month by sampling one permanent plot of 20 × 20 m per stand. • Key results: Seed-rain abundance reached 9 × 10³ seeds m⁻² year⁻¹, and was characterized by synchronous sequences of low and high seed production. New seedlings appeared only during the year following a seed production of at least 1 × 10³ and 1.5 × 10² seeds m⁻² year⁻¹ for balsam fir and white spruce, respectively. Seedlings emerged in July and survived 34-52 d on average, with balsam fir showing a longer lifespan and lower mortality, although 85–99% of seedlings died before completing one year of life. • Conclusions: The emergence of young seedlings was coupled with massive seed rains, which allowed synchronous replenishment of the seedling banks among stands and species, and generated different cohorts, yielding a discontinuous age structure.