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.     

Similar tree seedling responses to shrubs and to simulated environmental changes at Pyrenean and subarctic treelines

Background: Climate, land-use and disturbance regimes are key drivers of treeline dynamics worldwide, but local and regional spatio-temporal patterns indicate that additional factors play an important role. Some studies suggest that shrub-tree interactions control tree seedling recruitment patterns across the treeline ecotone, but little is known about the generality of this interaction.

Aims: We established an experiment in a treeline ecotone in the central Pyrenees to investigate the role of such interactions and other environmental factors on tree seedling growth and survival. It was based on a similar experiment we completed recently in the subarctic Scandes. By comparing local phenomena between both experiments we assessed the generality of the findings across regions with different biogeographic histories and species characteristics.

Methods: We followed the survival and growth of transplanted Pinus uncinata seedlings during three growing seasons in a multi-factorial design (forest vs. treeline, +/– shrub removal, +/– temperature increase and +/– nutrient addition).

Results: There was better seedling growth at the treeline compared to the forest, and the presence of shrubs prevented seedling winter damage and herbivory. Both temperature and nutrient increase enhanced seedling performance.

Conclusions: Although some particular details were exclusive to the Pyrenees or to the Scandes, the similarities indicated general patterns and help to interpret the underlying mechanisms of treeline dynamics in both regions. This convergence of responses points to the potential of the development of a robust mechanistically founded predictive framework for scaling up shrub impacts on treeline dynamics to other regions.     

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.     

Mosses modify effects of warmer and wetter conditions on tree seedlings at the alpine treeline

Climate warming enables tree seedling establishment beyond the current alpine treeline, but to achieve this, seedlings have to establish within existing tundra vegetation. In tundra, mosses are a prominent feature, known to regulate soil temperature and moisture through their physical structure and associated water retention capacity. Moss presence and species identity might therefore modify the impact of increases in temperature and precipitation on tree seedling establishment at the arctic‐alpine treeline. We followed Betula pubescens and Pinus sylvestris seedling survival and growth during three growing seasons in the field. Tree seedlings were transplanted along a natural precipitation gradient at the subarctic‐alpine treeline in northern Sweden, into plots dominated by each of three common moss species and exposed to combinations of moss removal and experimental warming by open‐top chambers (OTCs). Independent of climate, the presence of feather moss, but not Sphagnum, strongly supressed survival of both tree species. Positive effects of warming and precipitation on survival and growth of B. pubescens seedlings occurred in the absence of mosses and as expected, this was partly dependent on moss species. P. sylvestris survival was greatest at high precipitation, and this effect was more pronounced in Sphagnum than in feather moss plots irrespective of whether the mosses had been removed or not. Moss presence did not reduce the effects of OTCs on soil temperature. Mosses therefore modified seedling response to climate through other mechanisms, such as altered competition or nutrient availability. We conclude that both moss presence and species identity pose a strong control on seedling establishment at the alpine treeline, and that in some cases mosses weaken climate‐change effects on seedling establishment. Changes in moss abundance and species composition therefore have the potential to hamper treeline expansion induced by 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.     

Feathermoss seedbeds facilitate black spruce seedling recruitment in the forest–tundra ecotone (Labrador,Canada)

As climate warms, conifers are expected to expand their ranges into alpine tundra where ecological factors such as seedbed availability, and post‐dispersal seed and seedling predation may control local recruitment. Seedbed composition may influence microhabitat, nutrients, physical structure, and predation level and, therefore, affect the success of conifer recruitment, thereby providing the template for future expansion. In the boreal forest, seedbed–seedling competition dominates such that seedbed removal increases black spruce recruitment. In the harsher climate of the Mealy Mountains boreal forest–tundra ecotone (Labrador, Canada) the Stress gradient hypothesis (SGH) predicts that facilitation may dominate seedbed–seedling interactions. This study investigated potential mechanisms of seedbed facilitation (temperature, water, nutrients, physical protection) in three conifer seedbeds (Pleurozium schreberi, Cladonia spp., bare soil) and examined whether seed predation and/or seedling herbivory varied among seedbeds over three years. Seed emergence was low overall (< 10% on all treatments), but highest on Pleurozium (6.3%), followed by bare ground (4.6%) and Cladonia (0.3%). Facilitation was observed between Pleurozium and black spruce as seedling height increase (31%) and survival (55%) were highest; herbivory, seed predation and overwinter mortality were lowest compared to both Cladonia and bare ground seedbeds. Unlike in the closed canopy boreal forest, seedlings recruited poorly on bare soil as seedling height increase and survival were 20.5% and 26%. Temperature and water availability were similar across seedbeds, while nutrient availability was higher on Pleurozium. The physical structure of Pleurozium likely protects first to third‐year seedlings from temperature extremes and predators. As climate warms and seed availability increases, Pleurozium may facilitate black spruce recruitment and treeline expansion.     

The potential for mycobiont sharing between shrubs and seedlings to facilitate tree establishment after wildfire at Alaska arctic treeline

Root‐associated fungi, particularly ectomycorrhizal fungi (EMF), are critical symbionts of all boreal tree species. Although climatically driven increases in wildfire frequency and extent have been hypothesized to increase vegetation transitions from tundra to boreal forest, fire reduces mycorrhizal inoculum. Therefore, changes in mycobiont inoculum may potentially limit tree‐seedling establishment beyond current treeline. We investigated whether ectomycorrhizal shrubs that resprout after fire support similar fungal taxa to those that associate with tree seedlings that establish naturally after fire. We then assessed whether mycobiont identity correlates with the biomass or nutrient status of these tree seedlings. The majority of fungal taxa observed on shrub and seedling root systems were EMF, with some dark septate endophytes and ericoid mycorrhizal taxa. Seedlings and adjacent shrubs associated with similar arrays of fungal taxa, and there were strong correlations between the structure of seedling and shrub fungal communities. These results show that resprouting postfire shrubs support fungal taxa compatible with tree seedlings that establish after wildfire. Shrub taxon, distance to the nearest shrub and fire severity influenced the similarity between seedling and shrub fungal communities. Fungal composition was correlated with both foliar C:N ratio and seedling biomass and was one of the strongest explanatory variables predicting seedling biomass. While correlative, these results suggest that mycobionts are important to nutrient acquisition and biomass accrual of naturally establishing tree seedlings at treeline and that mycobiont taxa shared by resprouting postfire vegetation may be a significant source of inoculum for tree‐seedling establishment beyond current treeline.     

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.     

Microtopographic Control of Treeline Advance in Noatak National Preserve, Northwest Alaska

Arctic treeline positions are of fundamental importance to the function of high latitude landscapes, as regulators of surface energy exchange and carbon cycling. Most studies aimed at explaining current and predicting future treeline positions have examined growth trends in mature treeline trees, but treeline advance requires seed production, germination, seedling establishment, and recruitment of new trees beyond the treeline and these processes may not be well correlated with growth of mature trees. Tussock tundra is a widespread, microtopographically complex vegetation type, covering vast areas of northern Alaska and Siberia. We examined a site where the white spruce (Picea glauca) treeline has recently advanced into tussock tundra and asked if white spruce seedlings (~36-cm tall) occurred in tussocks, inter-tussocks, and frost boils disproportionate to the abundance of these habitats in the treeline ecotone. We found that seedlings disproportionately occurred in tussocks and frost boils. Seedlings found in tussocks and frost boils also showed greater branch growth than those in inter-tussocks. Tussocks and frost boils had higher soil temperatures than inter-tussocks. Tussocks had higher N, P, and K availability, whereas frost boils had greater secondary and micronutrient availability. The disproportionate occurrence of seedlings in tussocks and frost boils can likely be explained by the combination of warmer soils and greater nutrient availability. It is also possible that low competition for primary nutrients contributes to the success of seedlings in frost boils. Results of the study highlight the importance of biotic and abiotic facilitation as mediators of treeline advance in a changing Arctic.     

The importance of biotic filtering on boreal conifer recruitment at alpine treeline

Treeline, the ecotone where forest transitions to alpine or tundra ecosystems, is considered the thermal limit to tree growth and survival. Despite temperature increases across mountainous areas and high latitudes globally, there has been no ubiquitous change in treeline position. The process of range expansion must initially depend on increased recruitment at, or beyond current range limits and recruitment limitations have been hypothesized as a mechanism for the variable response of treeline position to climate warming. We conducted a unique series of observational and experimental studies to quantify early-life stage constraints, from seed production to seedling establishment, on black spruce Picea mariana and tamarack Larix laricina recruitment at a model alpine treeline in Newfoundland, Canada. We found recruitment at treeline to be simultaneously seed and establishment limited. The treeline population produced fewer seeds than the forest population and black spruce seeds produced at treeline were less viable. Tamarack was more seed limited than black spruce where seed viability was low regardless of altitudinal position. Post-dispersal seed predation greatly constrained recruitment across the altitudinal gradient; however, black spruce seeds experienced the lowest levels of invertebrate seed predation on the lichen mat at treeline. If seeds were not consumed, individuals at treeline were establishment limited where germination and seedling establishment was both less abundant and delayed on lichen substrate. Our study highlights the need for multiple factors to align temporally for significant recruitment at treeline to occur.     

Performance of germinating tree seedlings below and above treeline in the Swiss Alps

The germination and early survival of tree seedlings is a critical process for the understanding of treeline dynamics with ongoing climate change. Here we analyzed the performance of 0–4 year-old seedlings of seven tree species at three sites above and below the current altitudinal treeline in the Swiss Central Alps near Davos. Starting from sown seeds, we monitored the seedling performance as proportions of living seedlings, seedling shoot height growth, and biomass allocation over 4 years to examine changes along an elevational gradient. We evaluated the relative importance of the environmental factors soil temperature, light conditions, water use efficiency, and nitrogen availability on seedling performance. During the 4 years, the proportions of living seedlings differed only slightly along the elevational gradient even in species currently occurring at lower elevations. Microsite-specific soil temperature and light availability had only little effect on the proportion of living seedlings and seedling biomass across the elevational gradient. Conversely, seedling biomass and biomass allocation correlated well with the foliar stable nitrogen isotope abundance (δ ¹⁵N) that was used as an indicator for nitrogen availability. Collectively, our results suggested that the early establishment of seedlings of a variety of tree species in the treeline ecotone was not limited by current climatic conditions even beyond the species’ actual upper distribution limit. Nitrogen dynamics appeared to be an important environmental co-driver for biomass production and allocation in very young tree seedlings.     

植物群落类型和围栏封育对高山林线岷江冷杉幼苗成活的影响

林线树种幼苗的生长紧密关系着高山林线的形成。通过室内发芽实验测定了岷江冷杉种子萌发能力。2015年春,在川西巴郎山岷江冷杉林线之上的高山林线交错带,通过播种处理(播种和不播种),研究了不同植物群落类型(灌木和草地群落)和围栏处理(围栏和不围栏)对岷江冷杉幼苗成活率的影响,以期为进一步探讨高山林线形成机制提供科学参考。结果表明:(1)岷江冷杉室内发芽率31.4%,成活率随时间下降。(2)灌木群落的全年日平均空气温湿度显著高于草本群落,而两群落的全年日平均土壤温度无显著差异。(3)未播种样地没有发现岷江冷杉幼苗,说明林线交错带缺乏岷江冷杉种子。(4)与不围栏样地相比,围栏封育分别显著提高了岷江冷杉幼苗成活率2.0%(2015年)和2.2%(2016年);与灌木群落相比,草本群落中岷江冷杉幼苗成活率显著提高0.8%(2015年)和1.2%(2016年),说明灌木对林线交错带幼苗更新具有更强的竞争作用。围栏处理下,草本群落中岷江冷杉幼苗成活率显著高于灌木群落;不围栏处理下,草本群落中岷江冷杉幼苗成活率显著低于灌木群落,2017年在不围栏样地没有幼苗存活,说明动物干扰对林线交错带森林更新有阻碍作用。研究表明,高山林线交错带森林更新,受种子、植物竞争和动物干扰(例如,放牧)等多方面因素的共同影响,可以通过播种、植物剔除(去除竞争)和围栏等促进幼苗更新和成活。     

Seedling occurrence in alpine treeline conifers: A case study from the central Rocky Mountains,USA

Information concerning the occurrence of very young (1- to 10-year-old) tree seedlings in the alpine treeline ecotone is rare. Seedling occurrence of the dominant conifers Picea engelmannii and Abies lasiocarpa was measured in the treeline ecotone of the Medicine Bow Mountains, Wyoming (central Rocky Mountains, USA), an area composed of elongated tree islands separated by open meadows (ribbon forest) that grade into the closed forest. No seedlings were found on the windward sides of tree islands, while a mean of 0.6 seedlings.m^–2 occurred on the leeward (downward) sides. These values compared to the 4.2 seedlings.m^–2 in the closed forest. In addition, a strong correspondence was found between snowpack depth and seedling abundance, with depths that were either too shallow (< 0.5 m) or too deep (> 1.5 m) associated with fewer or no seedlings. A. lasiocarpa seedlings made up much less of the overall seedling population in the ribbon forest (6 %) than in the closed forest (22 %). Seedling establishment in this portion of the alpine treeline ecotone appears to be occurring at a low rate that differs between the two dominant species and may be strongly influenced by wind-driven snow accumulation patterns.     

Limited prospects for future alpine treeline advance in the Canadian Rocky Mountains

Treeline advance has occurred throughout the twentieth century in mountainous regions around the world; however, local variation and temporal lags in responses to climate warming indicate that the upper limits of some treelines are not necessarily in climatic equilibrium. These observations suggest that factors other than climate are constraining tree establishment beyond existing treelines. Using a seed addition experiment, we tested the effects of seed availability, predation and microsite limitation on the establishment of two subalpine tree species (Picea engelmannii and Abies lasiocarpa) across four treelines in the Canadian Rocky Mountains. The effect of vegetation removal on seedling growth was also determined, and microclimate conditions were monitored. Establishment limitations observed in the field were placed in context with the effects of soil properties observed in a parallel experiment. The seed addition experiment revealed reduced establishment with increasing elevation, suggesting that although establishment within the treeline ecotone is at least partially seed limited, other constraints are more important beyond the current treeline. The effects of herbivory and microsite availability significantly reduced seedling establishment but were less influential beyond the treeline. Microclimate monitoring revealed that establishment was negatively related to growing season temperatures and positively related to the duration of winter snow cover, counter to the conventional expectation that establishment is limited by low temperatures. Overall, it appears that seedling establishment beyond treeline is predominantly constrained by a combination of high soil surface temperatures during the growing season, reduced winter snowpack and unfavourable soil properties. Our study supports the assertion that seedling establishment in alpine treeline ecotones is simultaneously limited by various climatic and nonclimatic drivers. Together, these factors may limit future treeline advance in the Canadian Rocky Mountains and should be considered when assessing the potential for treeline advance in alpine systems elsewhere     

高山林线生态交错区木本植物幼苗分布特征、更新机制及其对气候变化的响应

木本植物幼苗是高山林线生态交错区的重要组成部分,其更新对气候变化背景下树线的移动至关重要.本研究通过对近几十年来全球范围内林线生态交错区的木本植物幼苗分布特征、更新机制及其对气候变化响应的研究总结得出:林线生态交错区木本植物幼苗的空间分布类型主要为渐变型和聚集型,且不同分布类型对树线动态的指示意义各异.在全球尺度上,其分布的海拔高限通常与生长季长度、均温和物种特性等有关,而在区域尺度上则多受降水影响.在幼苗更新初期,种源在很大程度上决定了种子的萌发及分布位置,之后微环境的促进作用为幼苗的定植提供庇护,提高其存活率,而在更新后期多种生物和非生物因素及其相互作用则非常关键.气候变暖促使林线生态交错区气温升高、降水充沛,有利于幼苗生长,使其向高海拔区域扩张而成为树线上移的先兆,但部分物种受遗传特性或适应策略影响,仅表现为密度增加,使树线保持相对稳定.未来应借助树轮、14C等精确定年技术,通过长期的野外定位观测和室内模拟,加强多时空尺度下林线幼苗的空间分布特征和更新机制研究,分析不同类型林线内木本植物幼苗的适应策略,预测气候变化背景下的树线动态,为山地生态系统恢复及保护提供科学依据.     

Promoting survival prospects of rare plants

Many hypotheses have been put forward to explain the structure and position of alpine treelines. The spatial complexity of the ecotone, ranging from sharp boundaries to networks of tree patches within a heath matrix, may explain why no consensus has been reached. In this paper, we discuss factors from abiotic disturbances to herbivory that may help understand the spatial structure of the alpine treeline ecotone in Fennoscandia. The ecotone is dominated by mountain birch (Betula pubescens ssp. tortuosa), and may show a wide range of spatial structures. We discuss the influence of topography, seed limitations, seedling establishment, growth limitations, abiotic disturbances and herbivory as structuring factors. All of these factors may operate, but their relative importance in space and time is unknown. There is a basic difference between factors that prevent the establishment of trees, and thus act on early life history stages, and factors that thin out a previously dense forest, and thus act on adult trees. Mortality caused directly or indirectly by geometrid moths may belong to the latter category. We suggest that seedling and sapling mortality is more important than seed limitation for the establishment of new individuals in the treeline ecotone. Important mortality factors may be abiotic disturbances, competition (or allelopathy) from field layer plants and herbivory. The relative role of these factors needs to be examined further.     

Latitudinal response of subarctic tree lines to recent climate change in eastern Canada

Aim The predictions from biogeographical models of poleward expansion of biomes under a warmer 2 × CO2 scenario might not be warranted, given the non‐climatic influences on vegetation dynamics. Milder climatic conditions have occurred in northern Québec, Canada, in the 20th century. The purpose of this study was to document the early signs of a northward expansion of the boreal forest into the subarctic forest‐tundra, a vast heterogeneous ecotone. Colonization of upland tundra sites by black spruce (Picea mariana (Mill.) BSP.) forming local subarctic tree lines was quantified at the biome scale. Because it was previously shown that the regenerative potential of spruce is reduced with increasing latitude, we predicted that tree line advances and recent establishment of seedlings above tree lines will also decrease northwards. Location Black spruce regeneration patterns were surveyed across a > 300‐km latitudinal transect spanning the forest‐tundra of northern Québec, Canada (55°29′–58°27′ N). Methods Elevational transects were positioned at forest–tundra interfaces in two regions from the southern forest‐tundra and two regions from the northern forest‐tundra, including the arctic tree line. The surroundings of stunted black spruce, forming the species limit in the shrub tundra, were also examined. Position, total height and origin (seed or layer) of all black spruce stems established in the elevational transects were determined. Dendrochronological and topographical data allowed recent subarctic tree line advances to be estimated. Age structures of spruce recently established from seed (< 2.5 m high) were constructed and compared between forest‐tundra regions. Five to 20‐year heat sum (growing degree‐days, > 5 °C) and precipitation fluctuations were computed from regional climatic data, and compared with seedling recruitment patterns. Results During the 20th century, all tree lines from the southern forest‐tundra rose slightly through establishment of seed‐origin spruce, while some tree lines in the northern forest‐tundra rose through height growth of stunted spruce already established on the tundra hilltops. However, the rate of rise in tree lines did not slow down with latitude. The density of < 2.5‐m spruce established by seed declined exponentially with latitude. While the majority of < 2.5‐m spruce has established since the late 1970s on the southernmost tundra hilltops, the regeneration pool was mainly composed of old, suppressed individuals in the northern forest‐tundra. Spruce age generally decreased with increasing elevation in the southern forest‐tundra stands, therefore indicating current colonization of tundra hilltops. Although spruce reproductive success has improved over the twentieth century in the southern forest‐tundra, there was hardly any evidence that recruitment of seed‐origin spruce was controlled by 5‐ to 20‐year regional climatic fluctuations, except for winter precipitation. Main conclusions Besides the milder 20th century climate, local topographic factors appear to have influenced the rise in tree lines and recent establishment by seed. The effect of black spruce's semi‐serotinous cones in trapping seeds and the difficulty of establishment on exposed, drought‐prone tundra vegetation are some factors likely to explain the scarcity of significant correlations between tree establishment and climatic variables in the short term. The age data suggest impending reforestation of the southernmost tundra sites, although the development of spruce seedlings into forest might be slowed down by the harsh wind‐exposure conditions.     

Does shrub invasion indirectly limit grass establishment via seedling herbivory? A test at grassland‐shrubland ecotones

Question: Does shrub invasion at ecotones indirectly limit grass establishment by increasing mammalian seedling herbivory? Location: Chihuahuan Desert, New Mexico, USA. Methods: We tested the hypothesis that herbivore‐related mortality of seedlings of the dominant perennial grass Bouteloua eriopoda would be highest in shrub‐dominated portions of grassland‐shrubland ecotones. We tested the hypothesis in two Chihuahuan Desert sites featuring similar shrub encroachment patterns but different shrub species, grass cover, and different abundances of small mammals. Within each site we transplanted B. eriopoda seedlings to grass‐dominated, middle, and shrub‐dominated positions of replicate ecotones during the time of year (mid‐summer) when they would naturally appear and monitored seedling fates. We estimated population size/activity of putative small mammal herbivores. Results: Seedlings were killed by mammals in greater numbers in shrubland than in grassland or middle ecotone positions at the site with large herbivore numbers. At the site with low herbivore numbers, most seedlings were killed in middle ecotone positions. The abundance patterns of herbivores did not parallel patterns of seedling herbivory across the ecotones or between sites. Conclusions: Seedling herbivory is an important process and is related to vegetation composition, but the mechanisms underlying the relationship are not clear. We speculate that variation in small mammal foraging behavior may contribute to seedling herbivory patterns. Restoration strategies in the Chihuahuan Desert need to account for the abundance and/or behavior of native herbivores.     

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.     

Seedling establishment in relation to microhabitat variation in a windthrow gap in a boreal Pinus sylvestris forest

Abstract. The characteristics of microhabitats of established Pinus sylvestris and Betula seedlings were studied in a small windthrow gap in a mature P. sylvestris-dominated forest in the Petkeljärvi National Park in eastern Finland. Seedlings were strongly clustered in disturbed microhabitats, particularly uprooting pits and mounds, formed by tree falls. They covered 3% of the 0.3.ha study area consisting of the gap and some of the forest edge. Although Betula occurred only as scattered individuals in the dominant canopy layer of the forest, it accounted for 30% of the seedlings found in the study area. Betula regeneration was almost completely restricted to pits and mounds, where 91% of the seedlings were found. Uprooting spots were also the most important regeneration microhabitats for Pinus, where 60% of the seedlings grew, even though the seedlings were found in other substrates as well, particularly on sufficiently decomposed coarse wood. Undisturbed field- and bottom-layer vegetation had effectively hindered tree seedling establishment, which emphasises the role of soil disturbance for regeneration. While the establishment of seedlings was found to be clearly determined by the availability of favourable regeneration microhabitats, the early growth of seedlings was affected by a complex interaction of environmental variables, including the type of microhabitat, radiation environment and interferences caused by competing seedlings and adjacent trees. In the most important regeneration microhabitats, i.e. in uprooting pits and on mounds, the distributions of the local elevations of Pinus and Betula seedlings were different. Pinus seedlings occurred closer to ground level, i.e. on the fringes of pits and lower on mounds, while Betula seedlings grew deeper in pits and higher on mounds. The position of the Betula seedlings indicate that they may have a competitive advantage over Pinus seedlings in the dense seedling groups occurring in uprooting spots. We suggest that this initial difference in Pinus and Betula establishment may affect the subsequent within-gap tree species succession and can, in part, explain the general occurrence of Betula in conifer-dominated boreal forests.