Plant species composition shifts in the Tatra Mts as a response to environmental change: a resurvey study after 90 years

Mountain vegetation is often considered highly sensitive to climate and land-use changes due to steep environmental gradients determining local plant species composition. In this study we present plant species compositional shifts in the Tatra Mts over the past 90 years and discuss the potential drivers of the changes observed. Using historical vegetation studies of the region from 1927, we resurveyed 76 vegetation plots, recording the vascular flora of each plot using the same methodology as in the original survey. We used an indirect method to quantify plant species compositional shifts and to indicate which environmental gradients could be responsible for these shifts: by calculating shifts in estimated species optima as reflected in shifts in the ecological indicator values of co-occurring species. To find shifts in species composition, focusing on each vegetation type separately, we used ordination (DCA). The species optimum changed significantly for at least one of the tested environmental gradients for 26 of the 95 plant species tested; most of these species changed in terms of the moisture indicator value. We found that the strongest shifts in species composition were in mylonite grassland, snowbed and hygrophilous tall herb communities. Changes in precipitation and increase in temperature were found to most likely drive compositional shifts in vegetation resurveyed. It is likely that the combined effect of climate change and cessation of sheep grazing has driven a species composition shift in granite grasslands communities.     

Snowbeds are more affected than other subalpine–alpine plant communities by climate change in the Swiss Alps

While the upward shift of plant species has been observed on many alpine and nival summits, the reaction of the subalpine and lower alpine plant communities to the current warming and lower snow precipitation has been little investigated so far. To this aim, 63 old, exhaustive plant inventories, distributed along a subalpine–alpine elevation gradient of the Swiss Alps and covering different plant community types (acidic and calcareous grasslands; windy ridges; snowbeds), were revisited after 25–50 years. Old and recent inventories were compared in terms of species diversity with Simpson diversity and Bray–Curtis dissimilarity indices, and in terms of community composition with principal component analysis. Changes in ecological conditions were inferred from the ecological indicator values. The alpha‐diversity increased in every plant community, likely because of the arrival of new species. As observed on mountain summits, the new species led to a homogenization of community compositions. The grasslands were quite stable in terms of species composition, whatever the bedrock type. Indeed, the newly arrived species were part of the typical species pool of the colonized community. In contrast, snowbed communities showed pronounced vegetation changes and a clear shift toward dryer conditions and shorter snow cover, evidenced by their colonization by species from surrounding grasslands. Longer growing seasons allow alpine grassland species, which are taller and hence more competitive, to colonize the snowbeds. This study showed that subalpine–alpine plant communities reacted differently to the ongoing climate changes. Lower snow/rain ratio and longer growing seasons seem to have a higher impact than warming, at least on plant communities dependent on long snow cover. Consequently, they are the most vulnerable to climate change and their persistence in the near future is seriously threatened. Subalpine and alpine grasslands are more stable, and, until now, they do not seem to be affected by a warmer climate.     

Alpine flora dynamics – a critical review of responses to climate change in the Swedish Scandes since the early 1950s

Reports about changes of alpine plant species richness over the past 60 years in the Swedish Scandes are reviewed, synthesized and updated with data from recent reinventories. Methodologically, this endeavour is based on resurveys of the floristic composition on the uppermost 20 m of four high‐mountain summits. The key finding is that the species pool has increased by 60–170% since the 1950s and later. Some of the invading species are new to the alpine tundra, with more silvine and thermophilic properties than the extant alpine flora. Not a single species of the original flora has disappeared from any of the summits. This circumstance is discussed in perspective of widespread expectations of pending temperature‐driven extinction of alpine species in an alleged future warmer climate. These progressive changes coincided with distinct warming (summer and winter) since the late 1980s. During a short cooler period (1974–1994), the species numbers decreased and the upper elevational limits of some ground cover species descended. Thus, discernible responses, concurrent with both warming and cooling intervals, sustain a strong causal link between climate variability and alpine plant species richness. Methodologically, plot‐less revisitation studies of the present kind are beset with substantial uncertainties, which may overstate floristic changes over time. However, it is argued here that carefully executed and critically interpreted, no other method can equally effectively sense the earliest phases of plant invasions into alpine vegetation.     

Recent cooling and dynamic responses of alpine summit floras in the southern Swedish Scandes

Changes in plant species richness on alpine summits in the southern Swedish Scandes were analyzed between 2004/2006 and 2012. This period experienced consistent summer and winter cooling and finalized with a cold and snow rich summer 2012. Re‐surveys of these summits had previously documented substantial increases in species numbers in concordance with climate warming since the mid‐20th century. Over the present study period, species richness decreased by 25–46%. The majority of lost species were those that had advanced upslope during the previous warm episode. Cooling since the mid 2000s and particularly the unusually short and snow‐rich growth period in 2012 caused a floristic retrogression. Taken together with extensive upshifts of many species during previous relative warm decades, recent downshifts highlight the large capability of certain alpine species to track their ecological niches as climate changes. The pivotal importance of unusually late‐lying snow in 2012, suggests that snow cover phenology exerts a more direct effect on the composition of the alpine flora than ambient temperatures. Dynamic modeling of future ecological landscape evolution needs to consider episodes of the kind reported here.     

Short-term signals of climate change in Italian summit vegetation: observations at two GLORIA sites

Short-term changes occurring in high mountain vegetation were analysed using the data from two Italian sites already part of the GLobal Observation Research Initiative in Alpine environments (GLORIA – central Apennines and southwestern Alps). The study focused on a set of floristic (endemics), structural (life forms) and ecological (thermic vegetation indicator) variables. Vegetation data were collected according to the GLORIA multi-summit standardized method during the last decade. The re-visitation revealed a moderate decrease in regional endemic flora and significant variations in structural and ecological parameters. The increase in caespitose hemicryptophytes in both sites, in suffruticose chamaephytes in the central Apennines and in rosette-forming hemicryptophytes in the southwestern Alps emerged, highlighting the rapid responses of the alpine vegetation to climate warming. The increase in perennial life forms is related with the expansion of graminoids and small woody plants. These life forms seem to be most suitable to face climate warming in Italian summits. The increase in the thermic vegetation indicator exceeds the mean European summits increment, and this is due to the expansion of thermophilic species. Short-term analyses with fine spatial and temporal resolutions are still necessary to improve our understanding concerning species behaviour in high-elevation ecosystems.     

Low impact of climate change on subalpine grasslands in the Swiss Northern Alps

While phenological shifts and migration of isolated species under climate change have already been observed on alpine summits, very few studies have focused on community composition changes in subalpine grasslands. Here we use permanent plots monitored since 1954 and precisely located phytosociological censuses from 1970 to study compositional changes of subalpine grasslands in two distinct regions of the Swiss Northern Alps. In both areas, warming trends during the monitoring period were associated with changes in land management (abandonment of goat and sheep pasturing or grazing replaced by mowing). Old and recent inventories were compared with correspondence analyses (CA). Ecological indicator values, community‐affinities and biological traits of the species were used to infer the factors responsible for triggering the observed changes. In both regions, subalpine grasslands were stable with smaller changes than have previously been observed in alpine environments. Only a few species appeared or disappeared and changes were generally limited to increasing or decreasing frequency and cover of certain taxa. At one site, grazing abandonment favored fallow species. Some of these species were located at their upper altitudinal distribution limits and may have spread because of rising temperatures. In both areas, declining species were predominantly alpine and low‐growing species; their decline was probably due to increased competition (e.g., shadow) with more vigorous subalpine taxa no longer limited by grazing. We conclude that vegetation communities can respond rapidly to warming as long as colonization is facilitated by available space or structural change. In the subalpine grasslands studies, changes were mainly driven by land management. These communities have a dense vegetation cover and newly arriving herbaceous species preferring warmer conditions may take some time to establish themselves. However, climate disturbances, such as exceptional drought, may accelerate community changes by opening gaps for new species.     

Impact of climate change on alpine vegetation of mountain summits in Norway

Climate change is affecting the composition and functioning of ecosystems across the globe. Mountain ecosystems are particularly sensitive to climate warming since their biota is generally limited by low temperatures. Cryptogams such as lichens and bryophytes are important for the biodiversity and functioning of these ecosystems, but have not often been incorporated in vegetation resurvey studies. Hence, we lack a good understanding of how vascular plants, lichens and bryophytes respond interactively to climate warming in alpine communities. Here we quantified long-term changes in species richness, cover, composition and thermophilization (i.e. the increasing dominance of warm-adapted species) of vascular plants, lichens and bryophytes on four summits at Dovrefjell, Norway. These summits are situated along an elevational gradient from the low alpine to high alpine zone and were surveyed for all species in 2001, 2008 and 2015. During the 15-year period, a decline in lichen richness and increase in bryophyte richness was detected, whereas no change in vascular plant richness was found. Dwarf-shrub abundance progressively increased at the expense of lichens, and thermophilization was most pronounced for vascular plants, but occurred only on the lowest summits and northern aspects. Lichens showed less thermophilization and, for the bryophytes, no significant thermophilization was found. Although recent climate change may have primarily caused the observed changes in vegetation, combined effects with non-climatic factors (e.g. grazing and trampling) are likely important as well. At a larger scale, alpine vegetation shifts could have a profound impact on biosphere functioning with feedbacks to the global climate.     

Littoral benthic macroinvertebrates of mountain lakes in the Tatra Mountains (Slovakia,Poland)

Littoral benthic macroinvertebrates of 45 mountain lakes in the Tatra Mountains were sampled using a semi-quantitative method in September 2000. A total of 32,852 specimens were identified to 93 taxa belonging to 14 higher taxonomic groups. Multivariate statistics (CCA, RDA) and nine biotic metrics (AQEM/STAR) were used to explain relationships between macroinvertebrate assemblages and environmental variables. Up to 57% of the ecological position of littoral macroinvertebrate assemblages were explained by variance of environmental variables divided into chemical, trophic, physical, catchment and location. Five types of Tatra lakes were recognized using CCA: A — strongly acidified lakes (small catchment, low pH, high concentration of TP, DOC, highest amount of POM in littoral); B — alpine acidified lakes (low amount of POM, low values of biotic metrics); C — alpine non-acidified lakes (high value of diversity index, predominance of Diptera); D — subalpine acidified lakes (high values of biotic metrics: number of families, proportion of crenal and rhithral taxa/total taxa); E — subalpine non-acidified lakes (high values of biotic metrics: number of families, number of genera, BMWP score, number of taxa and abundance of EPT taxa). RDA was used to design five levels of macroinvertebrate taxa acidification tolerance. The Tatra Acidification Index (TAI) was established to assess the acidification status of the lakes in the Tatra Mts.     

Studies of the alpine flora along an east-west gradient in central Western Norway

Central Western Norway was divided up into seven mountain areas, representing an east-west gradient from Hardangervidda (area A) westwards to the islands of Tysnes and Stord (area G). The alpine flora of totally 138 species thins out gradually where A has the greatest number (137 species) and G has the fewest (55 species). A drop of 83 species from east to west must be due to ecological barriers which prevent growth on the coastal mountains. Several factors have been taken into account and discussed. Rich Cambro-Ordovician rocks are present in all the explored areas, but the zone narrows towards the west due to the overlying base-poor thrusted rocks in the Caledonian nappe system. West of Hardangervidda, the summits of the mountains are almost entirely composed of acid rocks, and habitats for basiphilous species are therefore not present at the highest altitudes. Climate during the season of winter dormancy differs much more than during the growing season between east and west. Alpine plants require climatic conditions conducive to the termination of growth in autumn, a state of quiescence due to cold winters, and stable snow conditions which give a rapid transition from winter to summer. As the oceanic climate does not comply with these requirements, the number of alpine species declines. The alpine flora of Westem Norway is closely related to that of Great Britain. This is probably explained historically by a common origin during the Late Weichselian period.     

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.     

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.     

Intensified grazing affects endemic plant and gastropod diversity in alpine grasslands of the Southern Carpathian mountains (Romania)

Alpine grasslands in the Southern Carpathian Mts, Romania, harbour an extraordinarily high diversity of plants and invertebrates, including Carpathic endemics. In the past decades, intensive sheep grazing has caused a dramatic decrease in biodiversity and even led to eroded soils at many places in the Carpathians. Because of limited food resources, sheep are increasingly forced to graze on steep slopes, which were formerly not grazed by livestock and are considered as local biodiversity hotspots. We examined species richness, abundance and number of endemic vascular plants and terrestrial gastropods on steep slopes that were either grazed by sheep or ungrazed by livestock in two areas of the Southern Carpathians. On calcareous soils in the Bucegi Mts, a total of 177 vascular plant and 19 gastropod species were recorded. Twelve plant species (6.8%) and three gastropod species (15.8%) were endemic to the Carpathians. Grazed sites had lower plant and gastropod species richness than ungrazed sites. Furthermore, grazed sites harboured fewer gastropod species endemic to the Carpathians than ungrazed sites. On acid soils in the Fagaras Mts, a total of 96 vascular plant and nine gastropod species were found. In this mountain area, however, grazed and ungrazed sites did not differ in species richness, abundance and number of endemic plant and gastropod species. Our findings confirm the high biodiversity of grasslands on steep slopes in the Southern Carpathian Mts and caution against increasing grazing pressure in these refuges for relic plants and gastropods as well as for other invertebrates.     

Sward structure and diet selection after sheep introduction on abandoned grassland in the Giant Mts, Czech Republic

The effect of rotational grazing on the sward structure of long-term abandoned grassland and the sheep diet selection were investigated in the Giant Mts (Krkono?e/Karkonosze). The aim of the study was to answer the following questions: (1) Does sheep grazing affect the sward structure of previously abandoned mountain grassland? (2) How does sheep diet selection develop within and among grazing seasons? (3) Which section of a pasture do sheep prefer to graze? Data was collected three times during each grazing season in spring, summer and in autumn in the years 2001, 2002 and 2003 Fifty 1 × 1 m plots were used where the plant species occurrence and damage of plants by grazing for all species were repeatedly recorded. 1) Nonsignificant changes in plant species occurrences were recorded in spite of obvious visual changes in sward structure due to grazing e.g. the retreat of tall dominant species characteristic of long-term unmanaged grasslands. 2) Seasonal as well as inter-annual changes in sheep diet selection were detected. In spring 2001, sheep grazed over a wide variety of plant species in low quantities. In subsequent springs, they preferred species favoured during previous autumns (Veratrum album subsp. lobelianum; Ranunculus platanifolius; Senecio ovatus) and changed the diet only as a result of the elimination of favoured plants. 3) Sheep preferred to graze at the highest elevated part of the pasture probably as a consequence of anti-predator behaviour not due to higher occurrence of favoured plant species or the presence of a drinking place or salt licks. Changes in the sward structure were mostly of a quantitative not qualitative character, thus the presence-absence data collection is not a suitable method for monitoring the effects of management restoration. The diet selection changed probably due to the animals’ experience; sheep had no experience with montane species in spring 2001. The sheep were able to recognize favoured plant species after 6 months of wintering in lowland.     

Grazing as a factor structuring grasslands in the Pyrenees

Questions: What are the relative roles of abiotic and grazing management factors on plant community distribution in landscapes? How are livestock type and stocking rate related to changes in vegetation structure and composition? Location: Sub‐alpine grasslands in the central and eastern Pyrenees. Methods: Multivariate analysis and variance partitioning methods were used to evaluate the relative roles of environmental factors in structuring vegetation composition and diversity patterns in three surveys on differently managed grasslands. Results: Vegetation composition within a region was affected by environmental factors hierarchically, changing first according to abiotic factors and then to grazing management. At landscape scales, abiotic factors explained two‐fold more variation in vegetation composition than grazing factors. Within landscape units, cattle grazing increased vegetation heterogeneity at landscape and patch scales, while sheep grazing favoured the presence of a specific set of species with high conservation value. Species composition was highly responsive to management variables compared to diversity components. Conclusions: The combination of sheep and cattle grazing at various stocking rates is an effective tool to preserve the diversity of plant species and communities within a region with a long tradition of livestock management, through the scaling up of effects by local processes occurring in patches at smaller scales.     

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.     

Changes in the Altitudinal Distribution of Alpine Plants in Katunskiy Biosphere Reserve (Central Altai) Revealed on the Basis of Multiyear Monitoring Data

In order to reveal climate-related changes in the plant diversity of alpine ecosystems in recent decades, a target region of the GLORIA (Global Observation Research Initiative in Alpine Environments) worldwide network consisting of four mountain summits representing an elevation gradient from the subalpine to the upper part of alpine ecotone (2181, 2231, 2358, and 2475 m above sea level) has been established in Katunskiy Biosphere Reserve (Russia, Central Altai). In the course of the observation period (2005–2015), species in the target region have shifted towards higher altitudes by 5.3 m on average. The plant species richness has increased on the three higher summits and decreased on the lowest summit.     

Upward shift of alpine plants increases floristic similarity of mountain summits

Question: Does the upward shift of species and accompanied increase in species richness, induced by climate change, lead to homogenization of Alpine summit vegetation? Location: Bernina region of the Swiss Alps. Methods: Based on a data set from previous literature we expand the analysis from species richness to beta‐diversity and spatial heterogeneity. Species compositions of mountain summits are compared using a two‐component heterogeneity concept including the mean and the variance of Sørensen similarities calculated between the summits. Non‐metric multidimensional scaling is applied to explore developments of single summits in detail. Results: Both heterogeneity components (mean dissimilarity and variance) decrease over time, indicating a trend towards more homogeneous vegetation among Alpine summits. However, the development on single summits is not strictly unidirectional. Conclusions: The upward shift of plant species leads to homogenization of alpine summit regions. Thus, increasing alpha‐diversity is accompanied by decreasing beta‐diversity. Beta‐diversity demands higher recognition by scientists as well as nature conservationists as it detects changes which cannot be described using species richness alone.     

高寒草地植物群落关键种对不同放牧家畜组合放牧的响应

如何通过合理的利用方式提高高寒草地管理水平,实现其可持续利用一直是草地生态学领域的研究热点。为明确不同放牧家畜组合下高寒草地植物群落关键种的演替规律及其驱动因素,基于中等放牧强度设置了不同放牧家畜组合放牧样地（牦牛单牧、藏羊单牧、牦牛藏羊1 ： 2混牧、牦牛藏羊1 ： 4混牧、牦牛藏羊1 ： 6混牧）以及围封样地,并在连续放牧处理7年后系统分析了不同放牧家畜组合下植物群落特征与土壤理化性质变化,以期确定最优放牧组合。结果表明：（1）不同放牧家畜组合均会显著降低高寒草地植物盖度,但对其物种丰富度与多样性指数的影响并不显著。（2）牦牛藏羊1 ： 2混牧下植物群落特征与禁牧处理下植物群落特征较为相似,且牦牛藏羊1 ： 2混牧下的植物群落稳定性最强、组织水平最高。（3）牦牛单牧与1 ： 2混牧下关键种为矮生嵩草;藏羊单牧下关键种为天山针茅;1 ： 4混牧下关键种为星毛委陵菜;1 ： 6混牧与围封下关键种为赖草。（4）围封能够显著降低土壤容重,藏羊单牧则会显著增加土壤容重与土壤速效氮、磷含量。（5）土壤速效氮、容重与含水量是驱动不同放牧家畜组合下植物群落关键种演替的重要理化因子。综上所述,中等放牧强度下,牦牛藏羊1 ： 2混牧是青藏高原高寒草地较为理想的利用方式。此外,相较于单纯增加草地物种丰富度或多样性,建立植物群落物种之间的有效关联是提高青藏高原高寒草地管理水平的另一关键途径。     

Potential warm-stage microrefugia for alpine plants: Feedback between geomorphological and biological processes

During interglacial stages, microrefugia are sites that support locally favorable climates within larger areas with unfavorable warmer climates. Despite recent theoretical representations of microrefugia, an appropriate ecological characterization is still lacking, mostly for warm periods. Across mountain/alpine areas, cold-adapted plant species could adopt different strategies to manage the effects of climate warming: (A) migration toward higher elevations and summits; (B) in situ resilience of communities and species populations within microrefugia; and C) adaptation and evolution by genetic differentiation. This review aims to distinguish and characterize from an ecological perspective glacial, nival, periglacial and composite landforms and deposits that may function as potential microrefugia during interglacial warm periods.We conducted a literature screening related to the geomorphological processes and landforms associated with vegetation and plant communities in alpine/mountain environments of Europe. They include glacial deposits rock glaciers, debris-covered glaciers, composite cones and channels. In Alpine regions, geomorphologic niches that constantly maintain cold-air pooling and temperature inversions are the main candidates for microrefugia. Within such microrefugia, microhabitat diversity modulates the responses of plants to disturbances caused by geomorphologic processes and supports their aptitude for surviving under extreme conditions on unstable surfaces in isolated patches. Currently, European marginal mountain chains may be considered as examples of macrorefugia where relict boreo-alpine species persist within peculiar geomorphological niches that act as microrefugia.This review contributes to identifying potential warm-stage microrefugia areas across alpine and mountain regions and determining certain landforms that play or may play such role under global-change scenarios. The occurrence of warm-stage microrefugia within these locations may be of great importance for the modeling of future distributions of species and assessing the risk of extinction for alpine species. Microrefugia may have important implications in micro-evolutionary processes that occur across alternating climatic phases.