Evergreen alpine shrubs have high freezing resistance in spring,irrespective of snowmelt timing and exposure to frost: an investigation from the Snowy Mountains,Australia

Over winter, alpine plants are protected from low-temperature extremes by a blanket of snow. Climate change predictions indicate an overall reduction in snowpack and an earlier thaw; a situation which could expose the tips of shrubs which extend above the snowpack to freezing events in early spring, and cause foliar frost damage during the onset of physiological activity. We assessed the photosynthetic responses of freezing-damaged shrub leaves from an assay of freezing temperatures in the Snowy Mountains in south-eastern Australia, using chlorophyll fluorometery ex situ. We sampled leaves that were exposed early during the spring thaw and leaves that were buried in snow for up to two extra weeks, from four evergreen shrub species at monthly intervals following the period of snowmelt. Freezing resistance (estimated from LT₅₀) was poorest at the earliest spring sampling time, in both exposed above-snow and protected below-snow foliage in all species. Protected foliage in early spring had lower freezing resistance than exposed foliage, but not significantly so. By the third sampling time, freezing resistance was significantly better in the lower protected foliage (LT₅₀ of ? 14) compared with the upper exposed foliage (LT₅₀ of ? 10) in one species. Over the course of spring, freezing resistance improved significantly in all species, with LT₅₀ values of between ? 10 and ? 15 °C by the third sampling time, which is lower than the minimum air temperatures recorded at that time (> ? 5 °C). The results indicate that the dominant evergreen shrub species in this area may only be susceptible to freezing events very early in spring, before a period of frost-hardening occurs after snowmelt. Later in spring, these alpine shrubs appear frost hardy, thus further perpetuating the positive feedbacks surrounding shrub expansion in alpine areas.     

Drought increases the freezing resistance of high-elevation plants of the Central Chilean Andes

Freezing temperatures and summer droughts shape plant life in Mediterranean high-elevation habitats. Thus, the impacts of climate change on plant survival for these species could be quite different to those from mesic mountains. We exposed 12 alpine species to experimental irrigation and warming in the Central Chilean Andes to assess whether irrigation decreases freezing resistance, irrigation influences freezing resistance when plants are exposed to warming, and to assess the relative importance of irrigation and temperature in controlling plant freezing resistance. Freezing resistance was determined as the freezing temperature that produced 50 % photoinactivation [lethal temperature (LT₅₀)] and the freezing point (FP). In seven out of 12 high-Andean species, LT₅₀ of drought-exposed plants was on average 3.5 K lower than that of irrigated plants. In contrast, most species did not show differences in FP. Warming changed the effect of irrigation on LT₅₀. Depending on species, warming was found to have (1) no effect, (2) to increase, or (3) to decrease the irrigation effect on LT₅₀. However, the effect size of irrigation on LT₅₀ was greater than that of warming for almost all species. The effect of irrigation on FP was slightly changed by warming and was sometimes in disagreement with LT₅₀ responses. Our data show that drought increases the freezing resistance of high-Andean plant species as a general plant response. Although freezing resistance increases depended on species-specific traits, our results show that warmer and moister growing seasons due to climate change will seriously threaten plant survival and persistence of these and other alpine species in dry mountains.     

Vascular plant diversity and climate change in the alpine zone of the Snowy Mountains,Australia

This study examines vascular plant species richness along an altitudinal gradient in alpine Australia. Vascular plant composition and soil temperature records were obtained for five summits (from 1729 m to 2114 m a.s.l.) using sampling protocols from the Global Observation Research Initiative in Alpine Environments program. Species richness was examined against altitude, aspect and climatic variables at different spatial scales (10 × 10 cm quadrats, 1 m² quadrats, clusters of 4 * 1 m² quadrats, for the summit area above a line 5 m altitudinally below the summit (the −5 m isoline), for the extended summit down to the −10 m isoline). About 75 taxa (70 species, 5 graminoid genera) were recorded, 9 of which are endemic to the small alpine area of ∼100 km². There were significant linear relationships between species richness and altitude and climatic variables for the top to −5 isolines on the summits. However, there was no consistent pattern for species richness at other spatial scales, altitude, aspect or climatic variables. The proportion of species for the whole summits with localised distributions (local endemics) increased with altitude. Predicted increasing temperatures and reduced snowcover is likely to result in an increase in species richness as shrubs, herbs and introduced weeds become more common at higher altitude. Because Australian alpine areas occur in narrow altitudinal bands with no nival zone, there are no higher altitudinal refuges available for alpine species. Therefore many of these species are likely to be at risk of extinction from climate change.     

No increase in alpine snowbed productivity in response to experimental lengthening of the growing season

Climate change effects on snow cover and thermic regime in alpine tundra might lead to a longer growing season, but could also increase risks to plants from spring frost events. Alpine snowbeds, i.e. alpine tundra from late snowmelt sites, might be particularly susceptible to such climatic changes. Snowbed communities were grown in large monoliths for two consecutive years, under different manipulated snow cover treatments, to test for effects of early (E) and late (L) snowmelt on dominant species growth, plant functional traits, leaf area index (LAI) and aboveground productivity. Spring snow cover was reduced to assess the sensitivity of snowbed alpine species to severe early frost events, and dominant species freezing temperatures were measured. Aboveground biomass, productivity, LAI and dominant species growth did not increase significantly in E compared to L treatments, indicating inability to respond to an extended growing season. Edapho‐climatic conditions could not account for these results, suggesting that developmental constraints are important in controlling snowbed plant growth. Impaired productivity was only detected when harsher and more frequent frost events were experimentally induced by early snowmelt. These conditions exposed plants to spring frosts, reaching temperatures consistent with the estimated freezing points of the dominant species (～?10 °C). We conclude that weak plasticity in phenological response and potential detrimental effects of early frosts explain why alpine tundra from snowbeds is not expected to benefit from increased growing season length.     

Cold in the common garden: comparative low-temperature tolerance of boreal and temperate conifer foliage

Because they maintain green foliage throughout the winter season, evergreen conifers may face special physiological challenges in a warming world. We assessed the midwinter low-temperature (LT) tolerance of foliage from eight temperate and boreal species in each of the genera Abies, Picea, and Pinus growing in an arboretum in Trondheim, Norway, using relative electrolyte leakage (REL) as an index of cell injury. Relatively LT sensitive species came from temperate coastal and Mediterranean environments and displayed a well-defined sigmoidal response to LT stress, with LT₅₀ ranging from −27 to −38°C. Species originating from boreal regions were not lethally stressed by slow freezing to temperatures as low as −80°C, while species from temperate mountains and continental interiors displayed intermediate responses, with LT50s ranging from −33 to −44°C. Further evaluation of one sensitive and one insensitive species in each genus showed that boreal species can survive quenching in liquid nitrogen at −196°C provided they are first slowly cooled to −30°C or lower. Quantitative image analysis of color changes resulting from LT stress followed by exposure to light showed that foliage from nonlethally stressed boreal species developed mild to moderate chlorosis while more sensitive species developed a mixture of chlorosis and necrosis, with significant necrosis occurring mainly at temperatures resulting in REL of 50% or more. Sensitive and insensitive trees differed significantly in total raffinose, sucrose, and total sugar concentrations, and raffinose and sucrose correlated significantly with LT₅₀ within the sensitive group. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Short-term signals of climate change along an altitudinal gradient in the South Alps

Short-term changes in plant species number, frequency and composition were studied along an altitudinal gradient crossing four summits from the treeline ecotone to the subnival zone in the South Alps (Dolomites, Italy). Large-scale (summit areas) and small-scale patterns (16 plots of 1 m2/summit) were monitored. After 5 years, a re-visitation of the summit areas revealed a considerable increase of species richness at the upper alpine and subnival zone (10% and 9%, respectively) and relatively modest increases at the lower alpine zone and the treeline ecotone (3% and 1%, respectively). At the small scale, the results were partly different, with species richness decreasing at the lower summits and increasing at the higher summits. The changes can most likely be attributed to climate warming effects and to competitive interactions. The main newcomers at the lower three summits were species from the treeline and the lower altitudinal zones. Only at the highest summit, the newcomers came from the alpine species pool. At the treeline ecotone, the abundance of Pinus cembra, of dwarf shrubs and clonal graminoid species increased. Here, displacements of alpine species may be predicted for the near future. At the higher summits, expansions of the established alpine species and further invasions of species from lower altitudes are forecasted.     

Increased spring freezing vulnerability for alpine shrubs under early snowmelt

Alpine dwarf shrub communities are phenologically linked with snowmelt timing, so early spring exposure may increase risk of freezing damage during early development, and consequently reduce seasonal growth. We examined whether environmental factors (duration of snow cover, elevation) influenced size and the vulnerability of shrubs to spring freezing along elevational gradients and snow microhabitats by modelling the past frequency of spring freezing events. We sampled biomass and measured the size of Salix herbacea, Vaccinium myrtillus, Vaccinium uliginosum and Loiseleuria procumbens in late spring. Leaves were exposed to freezing temperatures to determine the temperature at which 50 % of specimens are killed for each species and sampling site. By linking site snowmelt and temperatures to long-term climate measurements, we extrapolated the frequency of spring freezing events at each elevation, snow microhabitat and per species over 37 years. Snowmelt timing was significantly driven by microhabitat effects, but was independent of elevation. Shrub growth was neither enhanced nor reduced by earlier snowmelt, but decreased with elevation. Freezing resistance was strongly species dependent, and did not differ along the elevation or snowmelt gradient. Microclimate extrapolation suggested that potentially lethal freezing events (in May and June) occurred for three of the four species examined. Freezing events never occurred on late snow beds, and increased in frequency with earlier snowmelt and higher elevation. Extrapolated freezing events showed a slight, non-significant increase over the 37-year record. We suggest that earlier snowmelt does not enhance growth in four dominant alpine shrubs, but increases the risk of lethal spring freezing exposure for less freezing-resistant species.     

Effect of seasonal freeze–thaw cycle on net nitrogen mineralization of soil organic layer in the subalpine/alpine forests of western Sichuan,China

Nitrogen mineralization, a main way that soil organic nitrogen converts to mineral nitrogen, is one of the key processes in soil nitrogen cycle. The mineral nitrogen has an important role in plant growth in the growing season. It has been widely accepted that soil freezing in winter can kill a number of microorganisms, weakening soil nitrogen mineralization. However, more and more recent studies have documented that soil microorganisms still have high activity during the deep freezing period, and obvious nitrogen mineralization in winter. Seasonal freeze–thaw cycle is a common phenomenon in the subalpine/alpine forest region, which may have a strong effect on soil ecological processes. Furthermore, the changing pattern of seasonal freeze–thaw cycles might have a significant influence on soil nitrogen mineralization in this region in the scenarios of global warming. As yet, little attention has been given to nitrogen mineralization of soil organic layer as affected by changed seasonal freeze–thaw pattern, although the increasing studies have demonstrated that winter warming might give strong effects on the litter decomposition and microbial activity in the subalpine/alpine forest regions. Therefore, a method of intact soil core incubation in combination with natural environmental gradient was employed by transferring forest soils from 3582 m (A₁) of altitude to 3298 m (A₂) of altitude and 3023 m (A₃) of altitude in the subalpine/alpine forests of western Sichuan, respectively. The amounts and rates of net nitrogen mineralization in soil organic layer were measured. The incubation period included the growing season and the freeze–thaw season from May 24, 2010 to April 19, 2011. The results suggested that significant net nitrogen mineralization was only observed in soil organic layer at low altitude (A₃) during the whole incubation period. Forest soils at higher altitudes (A₁ and A₂) showed obvious soil nitrogen immobilization. In comparison with the growing season which showed remarkable nitrogen immobilization characteristic, the freeze–thaw season showed obvious nitrogen mineralization at lower altitudes (A₂ and A₃). In contrast, the nitrogen immobilization amounts at high altitude (A₁) in freeze–thaw period were less than those in the growing season. Besides, the maximum of net nitrogen mineralization amounts and rates at high altitude (A₁) in soil organic layer mainly occurred in the late stage of growing season and the onset of freezing, soil nitrogen mineralization at the middle altitude (A₂) mainly occurred in the onset of freezing and the deep freezing period, while the highest amount and rate of net nitrogen mineralization at low altitude (A₃) occurred in the early stage of thawing and the late stage of growing season. Furthermore, the amount and rate of soil net nitrogen mineralization during the freeze–thaw season were increasing with the decrease of altitude, which correlated with soil freeze–thaw cycle and freezing process at different altitudes. These results indicated that increasing soil temperature in the future could not only significantly enhance soil nitrogen mineralization in the freeze–thaw season, but also improve soil nitrogen mineralization by increasing freeze–thaw cycle times and shortening freeze–thaw period. However, the processes were significantly influenced by soil micro-environment of subalpine/alpine forest regions.     

Short-term variation in species richness across an altitudinal gradient of alpine summits

In response to climate warming, high altitude alpine vegetation may be replaced by typically lower altitude species, as species re-assemble and migrate to new areas. However, empirical evidence showing vegetation change in response to climate warming is largely unavailable for Australian alpine areas. Here, we examine changes in species richness with respect to climate and altitude over a 7?year period at a range of spatial scales in a re-survey of five alpine summits that are part of the Global Observation Research Initiative in Alpine Environments monitoring network. Eighty species were recorded in 2011 across all summits, an increase of 6 species since 2004. Mean species richness increased at the whole-of-summit scale from 45 to 50 species (about 12?%). At this scale, the rate of species richness increase was almost one new species per year, with 15 new species recorded at one summit. Here, shrub and graminoid species showed the largest increases. At the smaller spatial scales, changes in species richness were less pronounced. Turnover at the species and community level was typically moderate at all spatial scales and on all summits. The strength and direction of species richness change (the difference in species richness between the two sample periods, +/?) was not related to altitude nor variation in climate. Future re-surveys of the summits will confirm whether these short-term variations in species richness, particularly increases in shrubs, are indeed signals of longer-term trends and interactions with a changing climate.     

Assessment of climate change effects on alpine summit vegetation in the transition of tropical to subtropical humid climate

Mountain ecosystems are globally among the most vulnerable to the impacts of global warming. This study is carried out on high-mountain summits dominated by dwarf bamboo, Yushania niitakayamensis, in the alpine zone of subtropical Taiwan. A total of six summits were selected using a multi-summit approach, following the common protocol of the Global Observation Research Initiative in Alpine Environments (GLORIA). To assess the climate change effects, we used the soil temperature and statistical downscaling precipitation data to analyze the climatic changing patterns, also applied enhanced vegetation index (EVI) and vegetation diversity index to assess the vegetation responses among summits along the timeline. The results showed that the temperature has a significant increase in all summits, especially in winter. The annual chilling days have reduced around 50% from 2010 to 2019, which would imply the extension of plant growing season. The EVI values showed a tendency to increase over time. Furthermore, the long-term trend of EVI had an abnormal drop in 2018. This coincided with low precipitation and extremely high temperature in the early growing season, which may have caused water shortage and a decline in plant growth. Although the temperature was increasing significantly, the species diversity was slightly increasing in the first survey, but significantly decreased in the third survey. We suggest that the increase in temperature and the extension of the growing season might have led to the expansion of the bamboo species Y. niitakayamensis. Its expansion, combined with an extreme climate event could have caused the decline in species diversity.

     

The relationship between soil water storage capacity and plant species diversity in high alpine vegetation

Background: In those alpine regions where growing season precipitation is decreasing due to climate change, the capacity of soils to retain water may become an important factor for the persistence of plant species. However, the importance of soil water storage capacity (WSC) for plant species diversity has not been studied so far.

Aims: We aim to evaluate the relevance of WSC for species diversity of alpine plant communities in relation to temperature and length of growing season.

Methods: Species diversity was determined in 150 plots from a broad range of alpine vegetation types in the calcareous western part of the central Swiss Alps. WSC of soil cores sampled in every plot was determined, as well as rooting zone temperature and snowmelt date. Linear mixed models were used to assess the relationship between environmental data and species diversity.

Results: Species diversity was most strongly and positively related to WSC, followed by mean daily minimum temperature (T_min) of the growing season. Species diversity was significantly related to date of snowmelt only in sites with high WSC and/or T_min.

Conclusions: WSC represents an integrative measure for habitat quality and accounts for differences in species diversity within the study region. In order to understand and predict responses of plant species to climate change in high mountain regions, it may be crucial to also take changes in plant water supply into account.     

Using plant functional traits to explain community composition across a strong environmental filter in Australian alpine snowpatches

Environmental filters act to limit the local community assemblage from the regional species pool by restricting the viable trait states that can occur there. In alpine snowpatches, the timing of snowmelt is a strong environmental filter. In coming decades, the strength of this filter is likely to relax with global climate change. We used three continuous plant functional traits (leaf area, plant height, seed mass) and their divergence (using the FD_var index) to document current patterns of community assembly and predict plant community responses to future environmental filters in alpine snowpatch vegetation. The community trait-weighted mean for leaf area and height, but not seed mass, was significantly higher in early snowmelt zones relative to mid and late melting zones across all snowpatches. Mean FD_var for height (but not leaf area or seed mass), by contrast, was substantially lower in early snowmelt zones, indicating that species growing in early melt zones are consistently taller than those growing in other zones. These results suggest that if climate change leads to earlier snowmelt and hence, a longer growing season, taller (more competitive) species with larger leaf areas (more productive) may replace short species in snowpatches as these plant communities re-assemble in response to changing environmental filters.     

Vascular plant diversity and climate change in the alpine zone of the Lefka Ori,Crete

The aim of this study is to analyse the vascular flora and the local climate along an altitudinal gradient in the Lefka Ori massif Crete and to evaluate the potential effects of climate change on the plant diversity of the sub-alpine and alpine zones. It provides a quantitative/qualitative analysis of vegetation-environment relationships for four summits along an altitude gradient on the Lefka Ori massif Crete (1664–2339 m). The GLORIA multi-summit approach was used to provide vegetation and floristic data together with temperature records for every summit. Species richness and species turnover was calculated together with floristic similarity between the summits. 70 species were recorded, 20 of which were endemic, belonging to 23 different families. Cretan endemics dominate at these high altitudes. Species richness and turnover decreased with altitude. The two highest summits showed greater floristic similarity. Only 20% of the total flora recorded reaches the highest summit while 10% is common among summits. Overall there was a 4.96°C decrease in temperature along the 675 m gradient. Given a scenario of temperature increase the ecotone between the sub-alpine and alpine zone would be likely to have the greatest species turnover. Southern exposures are likely to be invaded first by thermophilous species while northern exposures are likely to be more resistant to changes. Species distribution shifts will also depend on habitat availability. Many, already threatened, local endemic species will be affected first.     

Effects of temperature and date of snowmelt on growth, reproduction, and flowering phenology in the arctic/alpine herb, Ranunculus glacialis

Changes in growing season temperature and duration may have profound effects on the population dynamics of arctic and alpine plant species in snow-bed and fell-field habitats. We examined how a typical herbaceous pioneer species, Ranunculus glacialis, responded to experimental climate change in open-top chambers for three seasons at an alpine site in southern Norway. Warming had no significant effect on any reproductive, growth or phenological variables, except for seed weight, which increased significantly during the first 2 ears. Despite large differences in average date of snowmelt among years, average reproductive output and ramet size differed little among years. Within-year variation in date of snowmelt had no impact on seed number or weight in either control or warmed plots. Leaf width and ramet leaf number decreased significantly with later snowmelt within a year. Experimental warming reduced the negative effect on ramet size of late snowmelt within a year to some extent. In general, R. glacialis reacts contrary to most other arctic/alpine species to experimental warming. Species with such low responsiveness to environmental conditions may be particularly vulnerable to climatic change, especially if their habitat is invaded by other species with higher phenotypic plasticity and a better competitive ability.     

Nutrient dynamics of reindeer forage species along snowmelt gradients at different ecological scales

Growing season nutrient dynamics of four reindeer forage species (Betula nana, Eriophorum angustifolium, Rumex acetosa and Vaccinium myrtillus) were studied in a mountainous sub-arctic landscape in N Sweden. Changes in nitrogen (N), phosphorus (P) and fibre (ADF) concentrations in leaves and shoots were analysed (from a reindeer foraging perspective) along four ecological gradients significant to regional and local snowmelt regimes: season, oceanicity, altitude, and snow-patch retreat. N and P concentrations showed marked seasonal variations with peaks occurring from the middle of June to the end of July depending on species and snowmelt progression. The seasonal pattern for ADF concentrations, as well as differences between snowmelt regimes, were less consistent and showed large differences between species. N concentrations increased along snowmelt gradients at small (within snow-patches) and medium (along altitude gradients) spatial scales. Furthermore, variations in N concentrations were temporally and spatially scale-dependent, which underlines the importance of scale for understanding plant nutrient dynamics in sub-arctic and alpine systems, as well for plant–animal interactions. The greatest temporal and spatial differences in nutrient quality were observed early in the season, i.e., at the time of highest nutrient requirements for reindeer. Climate-induced changes of the start and the progress of the growing season may result in significant consequences to the development of calves and subsequently to the population demography of reindeer, mediated through spatial and temporal changes in the distribution of high quality food. These results provide basic knowledge for reindeer management in the light of the ongoing global warming.     

Reduced early growing season freezing resistance in alpine treeline plants under elevated atmospheric CO2

The frequency of freezing events during the early growing season and the vulnerability to freezing of plants in European high‐altitude environments could increase under future atmospheric and climate change. We tested early growing season freezing sensitivity in 10 species, from four plant functional types (PFTs) spanning three plant growth forms (PGFs), from a long‐term in situ CO₂ enrichment (566 vs. 370 ppm) and 2‐year soil warming (+4 K) experiment at treeline in the Swiss Alps (Stillberg, Davos). By additionally tracking plant phenology, we distinguished indirect phenology‐driven CO₂ and warming effects from direct physiology‐related effects on freezing sensitivity. The freezing damage threshold (lethal temperature 50) under ambient conditions of the 10 treeline species spanned from ?6.7±0.3 °C (Larix decidua) to ?9.9±0.6 °C (Vaccinium gaultherioides). PFT, but not PGF, explained a significant amount of this interspecific variation. Long‐term exposure to elevated CO₂ led to greater freezing sensitivity in multiple species but did not influence phenology, implying that physiological changes caused by CO₂ enrichment were responsible for the effect. The elevated CO₂ effect on freezing resistance was significant in leaves of Larix, Vaccinium myrtillus, and Gentiana punctata and marginally significant in leaves of Homogyne alpina and Avenella flexuosa. No significant CO₂ effect was found in new shoots of Empetrum hermaphroditum or in leaves of Pinus uncinata, Leontodon helveticus, Melampyrum pratense, and V. gaultherioides. Soil warming led to advanced leaf expansion and reduced freezing resistance in V. myrtillus only, whereas Avenella showed greater freezing resistance when exposed to warming. No effect of soil warming was found in any of the other species. Effects of elevated CO₂ and soil warming on freezing sensitivity were not consistent within PFTs or PGFs, suggesting that any future shifts in plant community composition due to increased damage from freezing events will likely occur at the individual species level.     

高山林线交错带高山杜鹃的凋落物分解

凋落物分解是维持生态系统生产力、养分循环、土壤有机质形成的关键生态过程。高山林线交错带是陆地生态系统中对气候变化响应的敏感区域。季节变化和海拔梯度上的植被类型差异可能会影响该区域凋落物的分解,进而对高山生态系统的碳氮循环产生重要影响。采用凋落物分解袋的方法,研究了川西高山林线交错带优势种高山杜鹃(Rhododendron lapponicum)凋落叶在雪被期和生长季的分解特征。结果显示:(1)季节变化和植被类型对高山杜鹃凋落物的分解均具有显著影响(P0.05),凋落叶的质量损失主要发生在生长季且在高山林线最大,暗针叶林中雪被期的质量损失略高于生长季,但差异不显著;(2)林线交错带上高山杜鹃凋落叶分解缓慢,一年干物质失重率为9.62%,拟合分解系数k为0.145;(3)高山杜鹃凋落叶的质量变化主要体现在纤维素降解显著且集中在雪被期,木质素无明显降解,在高山林线上C/N、C/P、木质素/N变化幅度较小且C、N、P的释放表现得稳定而持续。结果表明,季节性雪被对林线交错带内高山杜鹃分解的影响不仅局限在雪被期内,雪被融化期间频繁的冻融作用和雪融水淋洗作用可能会促进高山杜鹃凋落物在生长季初期的分解。总的来看,在气候变暖的情景下,雪被的缩减、生长季的延长和高山杜鹃群落的扩张可能加速高山林线交错带高山杜鹃凋落物的分解。     

Vascular plant diversity and climate change in the alpine belt of the central Apennines (Italy)

The aim of this study is to analyse the vascular flora and the local climate along the altitude gradient in the largest alpine belt of the central Apennines (Majella National Park), and to contribute to the evaluation of the possible effects of global climate changes on the biodiversity of the alpine ecosystem. For this purpose floristic-quantitative analyses and temperature records on three different summits have been carried out by using the methodological protocol of the UE-GLORIA project (2001 2003); the project aims toward a standardised monitoring of flora and temperature in the alpine environment of the main European chains. From the analysis of the changes in species richness along the altitude gradient (2405 m versus 2730 m a.s.l.), it emerged that 70% of species do not reach the highest summit and only 11% of the overall flora is shared by all of the summits examined; a drop in mean temperature has been observed at soil level, along the same gradient from 3.11 to 0.03 °C. Floristic-quantitative and climatic analyses have been carried out even along the horizontal gradient (principal exposures), highlighting a great species richness and vegetation cover in eastward aspects. We singled out some endangered rare species and we proved that the slopes facing east will be the first to be affected by the coming of subalpine species from below, whereas northward exposures will be the most conservative, showing greater inertia toward the invasive process caused by the climate warming.     

Carbon sink limitation and frost tolerance control performance of the tree <Emphasis Type="Italic">Kageneckia angustifolia</Emphasis> D. Don (Rosaceae) at the treeline in central Chile

The decrease in temperature with increasing elevation may determine the altitudinal tree distribution in different ways: affecting survival through freezing temperatures, by a negative carbon balance produced by lower photosynthetic rates, or by limiting growth activity. Here we assessed the relative importance of these direct and indirect effects of altitudinal decrease in temperature in determining the treeline in central Chile (33°S) dominated by Kageneckia angustifolia. We selected two altitudes (2000 and 2200 m a.s.l.) along the treeline ecotone. At each elevation, leaf non-structural carbohydrates (NSC) and gas exchange parameters were measured on five individuals during the growing season. We also determined the cold resistance of K.␣angustifolia, by measuring temperatures that cause 50% seedling mortality (LT₅₀) and ice nucleation (IN). No differences in net photosynthesis were found between altitudes. Although no differences were detected on NSC concentration on a dry matter basis between 2000 and 2200 m, when NSC concentration was expressed on a leaf area basis, higher contents were found at the higher elevation. Thus, carbon sink limitations may occur at the K. angustifolia’s upper altitudinal limit. For seedlings derived from seeds collected at the 2200 m, LT₅₀ of cold-acclimated and non-acclimated plants were −9.5 and −7 °C, respectively. However, temperatures as low as −10 °C can frequently occur at this altitude during the end of winter. Therefore, low temperature injury of seedlings seems also be involved in the treeline formation in this species. Hence, a confluence of global (carbon sink limitation) and regional (freezing tolerance) mechanisms explains the treeline formation in the Mediterranean-type climate zone of central Chile.     

Freezing resistance of high-elevation plant species is not related to their height or growth-form in the Central Chilean Andes

The level and mechanism of resistance to damage from freezing temperatures is related to the ambient temperatures experienced by plants. High-elevation habitats exhibit a thermal gradient in the air–soil profile, exposing ground-level species to more rigorous thermal microhabitats than well aboveground species. Such gradients could explain an inverse relationship between freezing temperature damage (LT₅₀) and plant height, such that leaves of smaller plants tolerate freezing conditions, while leaves of taller plants avoid them. However, microclimatic data have not been included in previous studies evaluating those patterns. On the other hand, a proposed trade-off between plant functional traits and freezing resistance suggest that leaves of growth-forms with higher intensity of growth are less freezing resistant than those with lower growth intensity. In this work, we determined whether the ability to resist freezing temperatures is related to plant height or growth-form in 37 high-elevation species from the Central Chilean Andes. We proposed that (1) plants with contrasting heights are exposed to different thermal conditions during freezing events, (2) the level and the mechanism of freezing resistance are inversely related to plant height, and that (3) the level of freezing resistance varied between plants with low- and high-growth intensities. We found that high-Andean species of different heights are exposed to different thermal conditions depending on the distance from the ground. However, neither level nor mechanisms of freezing resistance were related to plant height. Leaves of both short and tall plants showed similar LT₅₀ and their main freezing resistance mechanism was tolerance. Moreover, leaves of growth-forms with high- and low-growth intensity resisted similar freezing temperatures. Our results are discussed in relation to environmental conditions that characterize the Central Chilean Andes.