Treeline Ecotone Research Articles

Soil properties as constraints to seedling regeneration beyond alpine treelines in the Canadian Rocky Mountains

ABSTRACT Plants growing at the edges of their range limits are expected to be particularly sensitive to changes in precipitation and temperature regimes associated with climatic change. However, non-climatic factors are increasingly recognized as important constraints to species’ range expansions. Therefore, we assessed the effects of soil provenance with respect to the alpine treeline on the germination, growth, and survival of Engelmann spruce (Picea engelmannii) seedlings. Seedlings were grown under controlled conditions in a growth chamber and greenhouse for ninety days in soils collected from four treeline ecotones in the Canadian Rocky Mountains. By controlling seed source and climatic conditions, and eliminating competition and predation, we attribute differences in seedling viability to soil properties that differed across elevation zones and individual treeline sites. Overall, alpine soils originating from beyond the species’ current elevational range were least amenable to growth, and there was some indication of reduced germination and survival in high-elevation soils. Forest soils, which were coarser and more nutrient rich, hosted seedlings with greater above- and below-ground biomass. Thus, the physical and chemical characteristics of alpine soils in our study region may constrain future treeline expansion, underscoring the importance of incorporating soil properties when considering species’ distributions under climate change.

Peculiarities and Determinants of Regeneration of Siberian Larch on the Upper Limit of Its Growth in the Urals

Quantitative and qualitative characteristics of cones and seeds and the emergence and survival of seedlings in the Siberian larch (Larix sibirica Ledeb.) have been studied in different parts of the ecotone at the upper boundary of arboreal vegetation (treeline ecotone) on Serebryanskii Kamen’ Mountain (Northern Urals) and on the hills (with a landmark 312 m above sea level) surrounding Chernaya Mountain (Polar Urals) from 2005 to 2011. We have found a decrease in the parameters of cones, number of seeds in the cones, their viability in laboratory with an increase in altitude, and differences in the number of seedlings between parts of the treeline ecotone. It is shown that the formation of Siberian larch generations on the Northern Urals occurs only after moist years. It is noted that, in the Polar Urals, Siberian larch produces a big seed crop every 2–3 years. It is proven that the seed production and mortality of seedlings in Siberian larch are influenced not only by air temperature and soil moisture, but also by wind load, snow depth, and soil temperature rapidly changing along the slope.

Subalpine Forest and Treeline Ecotone under the Influence of Disturbances: A Review

This article considers disturbances caused by abiotic and biotic factors and human impact in the ecological region extending from subalpine forest to the upper tree limit. Both abiotic and biotic factors may cause reversible or irreversible disturbances. Disturbances by mass movement and avalanches give the subalpine forest and the treeline ecotone a distinct spatial pattern characterized by forest on safe topography and sites that preclude forest. Removal of the upper subalpine forests by humans has enlarged the snow-catchment area of avalanches and elongated the avalanche pathways. Consequently, avalanche destructive potential has increased. Hazards will probably increase due to climate change. External factors, like cyclonic storms, may cause fundamental disturbances. Fires have played a major role in the removal of high-elevation forests. Forest destruction by fire is often followed by soil erosion. Wild fires are likely to increase as a result of warming climate and would possibly prevent climatically-driven treeline advance. Cyclic or episodic mass outbreaks of defoliating insects and bark beetles, and pathogens also cause severe disturbances. Oversized populations of wild ungulates impede tree regeneration and can cause local soil erosion. Inadequate game management is the primary cause of intolerable ungulate numbers. Due to man-caused habitat fragmentation, the animals’ impact on the remained habitats has increased. Subalpine forest may recover from disturbance or become replaced by a substitute formation (e.g. krummholz). A subsequent absence of natural disturbances may also be considered a disturbance initiating a new development. Both natural and anthropogenic disturbances may counteract positive influences of climatic warming on subalpine forests and treeline. Effective measures to reduce or prevent abiotic and biotic disturbances of high-elevation forest may contribute to greater safety for people living in the endangered areas of the mountain valleys and also improve other ecosystem services of the subalpine forest.

Spatio-temporal dynamics of two alpine treeline ecotones and ecological characteristics of their dominate species at the eastern margin of Qinghai-Xizang Plateau

Spatio-temporal dynamics of two alpine treeline ecotones and ecological characteristics of their dominate species at the eastern margin of Qinghai-Xizang Plateau

Palynology and the Ecology of the New Zealand Conifers

The New Zealand conifers (20 species of trees and shrubs in the Araucariaceae, Podocarpaceae and Cupressaceae) are often regarded as ancient Gondwanan elements, but mostly originated much later. Often thought of as tall trees of humid, warm forests, they are present throughout in alpine shrublands, tree lines, bogs, swamps, and in dry, frost-prone regions. The tall conifers rarely form purely coniferous forest and mostly occur as an emergent stratum above evergreen angiosperm trees. During Maori settlement in the 13th century, these fire-sensitive trees succumbed rapidly, most of the drier forests being lost. As these were also the more conifer-rich forests, ecological research has been skewed towards conifer dynamics of forests wetter and cooler than the pre-human norm. Conifers are well represented in the pollen record and we here we review their late Quaternary history in the light of what is known about their current ecology with the intention of countering this bias. During glacial episodes, all trees were scarce south of c. 40°S, and extensive conifer-dominant forest was confined to the northern third of the North Island. Drought- and cold-resistant Halocarpus bidwillii and Phyllocladus alpinus formed widespread scrub in the south. During the deglacial, beginning 18,000 years ago, tall conifers underwent explosive spread to dominate the forest biomass throughout. Conifer dominance lessened in favour of angiosperms in the wetter western lowland forests over the Holocene but the dryland eastern forests persisted largely unchanged until settlement. Mid to late Holocene climate change favoured the more rapidly growing Nothofagaceae which replaced the previous conifer-angiosperm low forest or shrubland in tree line ecotones and montane areas. The key to this dynamic conifer history appears to be their bimodal ability to withstand stress, and dominate on poor soils and in cool, dry regions but, in wetter, warmer locations, to slowly grow thorough competing broadleaves to occupy an exposed, emergent stratum where their inherent stress resistance ensures little effective angiosperm competition.

Seedling recruitment and facilitation dependence on safe site characteristics in a Himalayan treeline ecotone

Seedling establishment is a crucial factor for spatial dynamics of treelines. However, safe sites for successful tree regeneration which is a basic prerequisite for a supposed treeline advance in response to climate warming have not been fully characterised, especially in the Himalayas. In this study, we aim at analysing safe sites of the tree species Abies spectabilis, Betula utilis and Rhododendron campanulatum, dominating the near-natural treeline ecotone in Rolwaling Himal, Nepal. We investigated microhabitat ground cover of established seedlings in rectangular plots along the treeline ecotone from subalpine forest stands via the krummholz belt to alpine dwarf shrub heaths (3800–4200 m). We examined protective elements that potentially ameliorate seedlings’ growth conditions and quantified the distance of seedlings to the nearest protective element. Our results showed species-specific microhabitat preferences for seedling establishment. While A. spectabilis seedlings establish on ground that is covered in average by 70% with litter, B. utilis and R. campanulatum seedlings primarily emerge on bryophyte mats (mean site cover 80%). Microhabitat structure did not vary significantly in relation to different vegetation types encompassing the treeline ecotone. The majority of seedlings of all species grew in maximum 2 m distance to at least one potentially protective element. Geomorphic elements and deadwood were the most important shelters for seedling establishment. Our results emphasise the importance of safe sites for successful tree regeneration in harsh environments. As growth conditions in high mountain ecosystems may substantially alter under contemporary climate change, the necessity of safe sites may increase to buffer potential extremes.

Typhoon diverged forest succession from natural trajectory in the treeline ecotone of the Changbai Mountains, Northeast China

Abstract Windthrow is a typical gap phrase disturbance that changes forest composition but rarely alter succession trajectories of a forest stand. Catastrophic windthrow such as typhoons is a large but infrequent disturbance that can have long-lasting effects on forest ecosystems. Whether catastrophic windthrow resets successional pathways of forest ecosystems remains inconclusive. We compared forest structure and regeneration patterns between undisturbed and disturbed stands 30 years after a typhoon event in the treeline ecotone of the Changbai Mountains, Northeastern China. Measurements were made using WorldView-1 satellite imaging and fieldwork in two types of large plots, one located in areas affected by the typhoon event and the other in adjacent undisturbed areas. Results showed that community divergence at landscape scale occurred following the typhoon. In non-wind–disturbed stands, forest development followed the successional pathway of both broadleaved and coniferous trees. By contrast, the late-successional coniferous rather than the early successional broadleaved trees occupied the wind–disturbed stands. The alteration mechanism on the forest succession pathway was that herbs formed tall, dense, and persistent mono-dominant thickets after the typhoon, which inhibited the regeneration of broadleaved trees. In contrast to the common knowledge that forest would return to initial successional stage after catastrophic disturbance, we concluded that catastrophic windthrow, different from gap phase windthrow, altered forest successional trajectory.

Causes for treeline stability under climate warming: Evidence from seed and seedling transplant experiments in southeast Tibet

Abstract Few data have examined the warming-related controls on seedling recruitment above the treeline, which is critical to understand causes for treeline stability under climate warming. We tested the hypothesis that compared to controls on seed dispersal and germination, the warming-induced decrease of seedling survival above the treeline is more likely to cause treeline stability. Long-term microclimate observations and 4-year seed and seedling transplant experiments were conducted in two contrasting treeline ecotones on the opposite slopes of a U-shaped valley at the peak of the Sergyemla Mountains. Seeds and young seedlings of Smith-fir (Abies georgei var. smithii) were transplanted to the treeline (4320 m) and Rhododendron shrub (4390 m) on the north-facing slope and to the juniper treeline (4425 m) on the south-facing slope. Transplanted seeds were collected from the Smith-fir treeline and its distribution center (3800 m). We also investigated the distribution of naturally-established seedlings and its correlation with shrub cover along elevation transects above the treeline. Smith-fir seedlings growing under shrubs were observed up to 40 m in elevation higher above the treeline, and there was either no correlation or a low correlation between seedling density and shrub cover. Harsh environments above the treeline did not limit the germination of seeds from different seed sources. However, transplanted seedlings cannot survival well above the treeline or at the juniper treeline due to high frequency of early-season freezing events and strong light intensity, while the absence of early-season freezing events in the Smith-fir treeline forest was more beneficial to seedling survival. During 2006–2015, the frequency of early-season freezing events increased with increasing annual mean air temperature above the treeline and at the juniper treeline but not in the Smith-fir treeline forest. The warming-induced increase of early-season freezing events, combined with high light intensity, mainly limits the survival of young seedlings above the treeline, which is more likely to cause treeline stability. Early-season freezing events should be an important index for evaluating the vulnerability of treeline forests to global warming.

Deconstructing human-shaped treelines: Microsite topography and distance to seed source control Pinus nigra colonization of treeless areas in the Italian Apennines

Human-shaped treelines are a common feature in mountain landscapes across Europe, and particularly in secularly managed Mediterranean high-elevation areas. The abandonment of traditional land use, and especially the reduction in grazing pressure at high elevations, triggered secondary succession in treeless grassland areas, and favored the upward shift of anthropogenic treelines in some cases. We investigated this process in four anthropogenic treeline ecotone sites in the Central Apennines, Italy, populated by European black pine (Pinus nigra Arn.). The upward treeline shift was controlled by microsite topography and the proximity to plantations acting as seed source. We found a 50% probability of producing cones in trees with basal diameters of 15–25 cm, heights of 2–5 m and ages of 20–25 years. The role played by climate on growth and the recruitment processes seems to be secondary, or could be masked by human-shaped processes. The presence of reproductive age trees at the treeline, mainly growing on debris-rich and steep slope sites, could indicate that the recruitment process will increase in future, leading to patchy tree patterns at different elevations. The high growth and encroachment rates observed at these human-shaped treelines would indicate that general growth dynamics are speeding up, including the tree colonization of treeless areas. These succession processes could cause a significant long-term decline in plant diversity in species-rich grasslands. Nonetheless, tree encroachment could increase forest protection against landslides and avalanches in the context of global change.

Population Spatial Dynamics of Larix potaninii in Alpine Treeline Ecotone in the Eastern Margin of the Tibetan Plateau, China

The high-altitude treeline is known to be sensitive to climate variability, and is thus considered as a bio-monitoring indicator of climate change. However, our understanding of the population dynamics and the cumulative climate-change effects on the alpine treeline ecotone in recent decades is limited. Here, we investigated the population dynamics of Larix potainii on the south- and north-facing slopes in the alpine treeline ecotone in the eastern margin of the Tibetan Plateau, China, including treeline position, population density, and tree recruitment. Results showed that on both south- and north-facing slopes, the treeline did not show a significant advancement in the past four decades. The population was dominated by young individuals, which tend to be established in the lower areas. Larix, here, tends to be clustered, especially in the upper areas. However, population density increased dramatically only on north-facing slopes. Larix here suffer from the stressful environment, but the warmer winter due to climate warming could facilitate the vertical growth of seedlings and saplings. Aggregated spatial patterns also provide a positive feedback in ameliorating the harsh environment. The slope-climate-moisture interactions have a pronounced impact on tree recruitment, including snow-limited tree establishment on the north-facing slopes and moisture-limited tree establishment on the south-facing slopes.

Assessing Fractional Cover in the Alpine Treeline Ecotone Using the WSL Monoplotting Tool and Airborne Lidar

ABSTRACTAs forest cover in mountain areas impacts headwater properties like habitat extent and downstream water resources, it is important to assess and understand the changes that occur across forest transition zones like the alpine treeline ecotone (ATE). Such changes occur slowly and manifest at decadal to century time scales; however, the present benchmark spatial resolution for land cover analysis from oblique repeat photographs, 100 m, is insufficient to analyze anticipated changes in ATE over a century-scale. In this research note, fractional cover classification of the ATE is achieved through oblique photography analysis with the WSL Monoplotting Tool. Seven oblique photographs of the West Castle Watershed (Alberta, Canada), collected by the Mountain Legacy Project, were gridded to a 20-m resolution and assigned canopy cover classes by manual interpretation. Four canopy cover classes (i.e., no cover, low vegetation, partial canopy, full canopy) were compared to lidar-derived fractional cover. The extraction of canopy cover information from oblique photography at a resolution of 20 m introduces the ability to assess and quantify changes in the ATE using century-scale oblique photographic records.

Among-tree variability and feedback effects result in different growth responses to climate change at the upper treeline in the Swiss Alps.

Upper treeline ecotones are important life form boundaries and particularly sensitive to a warming climate. Changes in growth conditions at these ecotones have wide‐ranging implications for the provision of ecosystem services in densely populated mountain regions like the European Alps. We quantify climate effects on short‐ and long‐term tree growth responses, focusing on among‐tree variability and potential feedback effects. Although among‐tree variability is thought to be substantial, it has not been considered systematically yet in studies on growth–climate relationships. We compiled tree‐ring data including almost 600 trees of major treeline species (Larix decidua, Picea abies, Pinus cembra, and Pinus mugo) from three climate regions of the Swiss Alps. We further acquired tree size distribution data using unmanned aerial vehicles. To account for among‐tree variability, we employed information‐theoretic model selections based on linear mixed‐effects models (LMMs) with flexible choice of monthly temperature effects on growth. We isolated long‐term trends in ring‐width indices (RWI) in interaction with elevation. The LMMs revealed substantial amounts of previously unquantified among‐tree variability, indicating different strategies of single trees regarding when and to what extent to invest assimilates into growth. Furthermore, the LMMs indicated strongly positive temperature effects on growth during short summer periods across all species, and significant contributions of fall (L. decidua) and current year's spring (L. decidua, P. abies). In the longer term, all species showed consistently positive RWI trends at highest elevations, but different patterns with decreasing elevation. L. decidua exhibited even negative RWI trends compared to the highest treeline sites, whereas P. abies, P. cembra, and P. mugo showed steeper or flatter trends with decreasing elevation. This does not only reflect effects of ameliorated climate conditions on tree growth over time, but also reveals first signs of long‐suspected negative and positive feedback of climate change on stand dynamics at treeline.

Dissimilar responses of larch stands in northern Siberia to increasing temperatures-a field and simulation based study.

Arctic and alpine treelines worldwide differ in their reactions to climate change. A northward advance of or densification within the treeline ecotone will likely influence climate-vegetation feedback mechanisms. In our study, which was conducted in the Taimyr Depression in the North Siberian Lowlands, w present a combined field- and model-based approach helping us to better understand the population processes involved in the responses of the whole treeline ecotone, spanning from closed forest to single-tree tundra, to climate warming. Using information on stand structure, tree age, and seed quality and quantity from seven sites, we investigate effects of intra-specific competition and seed availability on the specific impact of recent climate warming on larch stands. Field data show that tree density is highest in the forest-tundra, and average tree size decreases from closed forest to single-tree tundra. Age-structure analyses indicate that the trees in the closed forest and forest-tundra have been present for at least ~240yr. At all sites except the most southerly ones, past establishment is positively correlated with regional temperature increase. In the single-tree tundra, however, a change in growth form from krummholz to erect trees, beginning ~130yr ago, rather than establishment date has been recorded. Seed mass decreases from south to north, while seed quantity increases. Simulations with LAVESI (Larix Vegetation Simulator) further suggest that relative density changes strongly in response to a warming signal in the forest-tundra while intra-specific competition limits densification in the closed forest and seed limitation hinders densification in the single-tree tundra. We find striking differences in strength and timing of responses to recent climate warming. While forest-tundra stands recently densified, recruitment is almost non-existent at the southern and northern end of the ecotone due to autecological processes. Palaeo-treelines may therefore be inappropriate to infer past temperature changes at a fine scale. Moreover, a lagged treeline response to past warming will, via feedback mechanisms, influence climate change in the future.

Sensitivity to environmental change of the treeline ecotone and its associated biodiversity in European mountains

Sensitivity to environmental change of the treeline ecotone and its associated biodiversity in European mountains

Phytosociology and ecology of treeline ecotone vegetation in Rolwaling Himal, Nepal

Questions: Species composition and ecology of treeline vegetation types in the Rolwaling Valley is largely unknown to date. The aim of this study is to differentiate plant communities along the elevational gradient in the treeline ecotone, and to identify site factors which cause the differentiation of the treeline ecotone vegetation. Study area: Rolwaling Valley (27 degrees 52' N;86 degrees 25' E) in the central Himalaya, Nepal. Methods: We included 91 phytosociological releves along three elevational transects in this analysis, ranging from 3745 to 4300 m a.s.l. Plant communities were differentiated and vegetation-environment relationships analysed using cluster analysis and ordination. For each community and higher ranked unit, we detected diagnostic species according to their fidelity index (phi coefficient). Ecological preferences of each community were examined using one-way analysis of variance (ANOVA). Results and conclusions: We identified five communities, belonging to two different classes: the Synotis alata-Abies spectabilis and Ribes glaciale-Abies spectabilis communities occupy the upper part of the subalpine forest zone, representing the uppermost forest stands below treeline. These communities are primarily comprised of Abies spectabilis and Betula utilis in the tree layer, with varying cover of several tree species and distinct differences within the herb layer. Above treeline, the mixed forest stands are replaced by the species-poor Boschniakia himalaica-Rhododendron campanulatum community, forming a dense krummholz belt. We assigned the forest and krummholz communities to the class Betula utilis-Abies spectabilis forests. In the alpine zone two communities were classified and assigned to the class Dasiphora arbuscula-Rhododendron anthopogon dwarf shrub heaths.The species-poor Pedicularis cf. micro calyx-Rho dodendron anthopogon community has a less developed herbaceous layer, whereas the Anaphalis royleana-Rhododendron anthopogon community is characterized by higher cover and higher species richness of herbs and grasses. Species composition of identified communities is mainly differentiated by soil temperature, nitrogen supply and availability, and soil moisture content.

Phytosociology and ecology of treeline ecotone vegetation in Rolwaling Himal, Nepal

Phytosociology and ecology of treeline ecotone vegetation in Rolwaling Himal, Nepal

Changes in plant taxonomic and functional diversity patterns following treeline advances in the South Urals

Background: Treeline ecotones represent environmental boundaries that fluctuate in space and time and thus induce changes in plant taxonomic and functional diversity.Aims: To study changes through time in taxonomic and functional plant diversity patterns along the treeline ecotone.Methods: In 2002, vegetation was sampled along a gradient from upper montane forest to the treeline–alpine transition in the South Ural Mountains, Russia. In 2014, vegetation was resampled and plant functional traits were collected. We studied spatial and temporal changes in plant species composition, functional composition and functional diversity.Results: Species composition and diversity changed along the elevational gradient. The functional composition in height, leaf area, specific leaf area and leaf nitrogen content decreased with elevation, whereas functional composition of leaf carbon content increased. We found a temporal shift towards shorter plants with smaller leaves in treeline sites. Functional richness varied in several traits along the elevational gradient, while functional dispersion showed a trend towards increased functional dispersion in height, specific leaf area and leaf nitrogen in the treeline–tundra transition.Conclusions: Tree encroachment across the treeline ecotone has resulted in a shift in plant species relative abundances and functional diversity, possibly affecting plant community assembly patterns.

Why input matters: Selection of climate data sets for modelling the potential distribution of a treeline species in the Himalayan region

Betula utilis is a major constituent of alpine treeline ecotones in the western and central Himalayan region. The objective of this study is to analyse for the first time the performance of different climatic predictors in modelling the potential distribution of B. utilis in the subalpine and alpine belts of the Himalayan region. Using Generalized Linear Models (GLM) we aim at examining climatic factors controlling the species distribution under current climate conditions. We evaluate the predictive ability of climate data derived from different statistical methods GLMs were created using least correlated bioclimatic variables derived from two different climate data sets: 1) interpolated climate data (i.e. WORLDCLIM; Hijmans et al., 2005), and 2) quasi-mechanistical statistical downscaling (i.e. CHELSA; Karger et al., 2016). Model accuracy was evaluated using threshold-independent (Area Under the Curve) and threshold-dependent (True Skill Statistics) measures. Although there were no significant differences between the models in AUC, we found highly significant differences (p≤0.01) in TSS. We conclude that models based on variables of CHELSA climate data had higher predictive power, whereas models using WORLDCLIM climate data consistently overpredicted the potential suitable habitat for B. utilis.Although climatic variables of WORLDCLIM are widely used in modelling species distribution, our results suggest to treat them with caution when topographically complex regions like the Himalaya are in focus. Unmindful usage of climatic variables for environmental niche models potentially causes misleading projections.

Growth response of Scots pines in polar-alpine tree-line to a warming climate

Coniferous trees at the alpine and polar tree lines of the Northern Hemisphere represent the outermost limit of their ecological range. Under such conditions, even small temperature variations may cause growth responses, which therefore can be used as indicators for changing environmental conditions. In this study we analysed the radial growth of Scots pine (Pinus sylvestris L.) along a latitudinal and altitudinal gradient at six locations in the polar and alpine tree-line ecotone in Finnish Lapland. We evaluated the effect of climatic trends on pine growth in relation to tree age and region, specifying a northern and a southern region in the study area. We found a response of Scots pine to climatic variations until the 1980s, but not to the current warming period. Increasing growth trends could be detected since 2000 in the radial growth of southern located trees, predominantly of juvenile ages, while the northern trees did not respond significantly to the current warming. In the north recent warmer and wetter conditions during winter time, inducing snow loads, wind damages, diseases and frost damages possibly masked the benefits of warmer conditions. The missing link between warming and radial growth would affect the use of tree-rings as proxy for past climate and for predictions for forest extension in polar-alpine tree-line sites.

Decreasing nutrient concentrations in soils and trees with increasing elevation across a treeline ecotone in Rolwaling Himal, Nepal

At a global scale, tree growth in alpine treeline ecotones is limited by low temperatures. At a local scale, however, tree growth at its upper limit depends on multiple interactions of influencing factors and mechanisms. The aim of our research was to understand local scale effects of soil properties and nutrient cycling on tree growth limitation, and their interactions with other abiotic and biotic factors in a near-natural Himalayan treeline ecotone. Soil samples of different soil horizons, litter, decomposition layers, and foliage samples of standing biomass were collected in four altitudinal zones along three slopes, and were analysed for exchangeable cations and nutrient concentrations, respectively. Additionally, soil and air temperature, soil moisture, precipitation, and tree physiognomy patterns were evaluated. Both soil nutrients and foliar macronutrient concentrations of nitrogen (N), magnesium (Mg), potassium (K), and foliar phosphorus (P) decrease significantly with elevation. Foliar manganese (Mn) concentrations, by contrast, are extraordinarily high at high elevation sites. Potential constraining factors on tree growth were identified using multivariate statistical approaches. We propose that tree growth, treeline position and vegetation composition are affected by nutrient limitation, which in turn, is governed by low soil temperatures and influenced by soil moisture conditions.