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High mountain ecosystems and their biota are governed by low-temperature conditions and thus can be used as indicators for climate warming impacts on natural ecosystems, provided that long-term data exist. We used data from the largest alpine to nival permanent plot site in the Alps, established in the frame of the Global Observation Research Initiative in Alpine Environments (GLORIA) on Schrankogel in the Tyrolean Alps, Austria, in 1994, and resurveyed in 2004 and 2014. Vascular plant species richness per plot increased over the entire period, albeit to a lesser extent in the second decade, because disappearance events increased markedly in the latter period. Although presence/absence data could only marginally explain range shift dynamics, changes in species cover and plant community composition indicate an accelerating transformation towards a more warmth-demanding and more drought-adapted vegetation, which is strongest at the lowest, least rugged subsite. Divergent responses of vertical distribution groups of species suggest that direct warming effects, rather than competitive displacement, are the primary causes of the observed patterns. The continued decrease in cryophilic species could imply that trailing edge dynamics proceed more rapidly than successful colonisation, which would favour a period of accelerated species declines.

Complex topography regulates near‐surface temperature above the treeline. It may thus sustain microrefugia for alpine plants and relax the need of shifting upward when the climate warms. The effectiveness of these microrefugia rests on the premise that plant distributions in alpine landscapes are mainly controlled by fine‐scale topographic variation. We tested this assumption by relating the distribution of 79 plant species and 10 community attributes across 900 1 m² plots in a landscape spanning 1677 m of elevation to 17 topographical descriptors at resolutions between 1 and 301 m. We found that the presence of most species and most community attributes were better explained by topographic variation at coarser scales (> 20 m). Fine‐scale topography is more clearly reflected in moisture than in temperature requirements of species. The elevational gradient rather than topographic variation at any scale, is the single most important driver of both species distributions and the variation in community attributes in the area studied. We hypothesise that our results reveal a hitherto underestimated influence of spatial mass effects on alpine plant distributions. These effects can override environmental filtering at fine scales and will thus impede the survival of cold‐adapted plants in small and fragmented refugia under climate warming.

1. High alpine plants endure a cold climate with short growing seasons entailing severe consequences of an improper timing of development. Hence, their flowering phenology is expected to be rigorously controlled by climatic factors. 2. We studied ten alpine plant species from habitats with early and late melting snow cover for 2 years and compared the synchronizing effect of temperature sums (TS), time of snowmelt (SM) and photoperiod (PH) on their flowering phenology. Intraseasonal and habitat-specific variation in the impact of these factors was analysed by comparing predictions of time-to-event models using linear mixed-effects models. 3. Temperature was the overwhelming trigger of flowering phenology for all species. Its synchronizing effect was strongest at or shortly after flowering indicating the particular importance of phenological control of pollination. To some extent, this pattern masks the common trend of decreasing phenological responses to climatic changes from the beginning to the end of the growing season for lowland species. No carry-over effects were detected. 4. As expected, the impact of photoperiod was weaker for snowbed species than for species inhabiting sites with early melting snow cover, while for temperature the reverse pattern was observed. 5. Our findings provide strong evidence that alpine plants will respond quickly and directly to increasing temperature without considerable compensation due to photoperiodic control of phenology.

Background Anthropogenic climate change leads to increasing temperatures and altered precipitation and snowmelt patterns, especially in alpine ecosystems. To understand species’ responses to climate change, assessment of genetic structure and diversity is crucial as the basis for the evaluation of migration patterns, genetic adaptation potential as well as the identification of adaptive alleles. Results We studied genetic structure, diversity and genome-environment associations of two snowbed species endemic to the Eastern Alps with a large elevational range, Achillea clusiana Tausch and Campanula pulla L. Genotyping-by-sequencing was employed to assemble loci de novo, call variants and perform population genetic analyses. Populations of either species were distinguishable by mountain, and to some extent by elevation. We found evidence for gene flow between elevations. Results of genome-environment associations suggested similar selective pressures acting on both species, emanating mainly from precipitation and exposition rather than temperature. Conclusions Given their genetic structure and amount of gene flow among populations the two study species are suitable to serve as a model for genetic monitoring of climate change adaptation along an elevation gradient. Consequences of climate change will predominantly manifest via changes in precipitation and, thus, duration of snow cover in the snowbeds and indirectly via shrub encroachment accompanied by increasing shading of snowbeds at lower range margins. Assembling genomes of the study species and studying larger sample sizes and time series will be necessary to functionally characterize and validate the herein identified genomic loci putatively involved in adaptive processes.

AIM: Emerging polyploids may depend on environmental niche shifts for successful establishment. Using the alpine plant Ranunculus kuepferi as a model system, we explore the niche shift hypothesis at different spatial resolutions and in contrasting parts of the species range. LOCATION: European Alps. METHODS: We sampled 12 individuals from each of 102 populations of R. kuepferi across the Alps, determined their ploidy levels, derived coarse‐grain (100 × 100 m) environmental descriptors for all sampling sites by downscaling WorldClim maps, and calculated fine‐scale environmental descriptors (2 × 2 m) from indicator values of the vegetation accompanying the sampled individuals. Both coarse and fine‐scale variables were further computed for 8239 vegetation plots from across the Alps. Subsequently, we compared niche optima and breadths of diploid and tetraploid cytotypes by combining principal components analysis and kernel smoothing procedures. Comparisons were done separately for coarse and fine‐grain data sets and for sympatric, allopatric and the total set of populations. RESULTS: All comparisons indicate that the niches of the two cytotypes differ in optima and/or breadths, but results vary in important details. The whole‐range analysis suggests differentiation along the temperature gradient to be most important. However, sympatric comparisons indicate that this climatic shift was not a direct response to competition with diploid ancestors. Moreover, fine‐grained analyses demonstrate niche contraction of tetraploids, especially in the sympatric range, that goes undetected with coarse‐grained data. MAIN CONCLUSIONS: Although the niche optima of the two cytotypes differ, separation along ecological gradients was probably less decisive for polyploid establishment than a shift towards facultative apomixis, a particularly effective strategy to avoid minority cytotype exclusion. In addition, our results suggest that coarse‐grained analyses overestimate niche breadths of widely distributed taxa. Niche comparison analyses should hence be conducted at environmental data resolutions appropriate for the organism and question under study.

Range shifts (especially during the Pleistocene), polyploidisation and hybridization are major factors affecting high-mountain biodiversity. A good system to study their role in the European high mountains is the Doronicum clusii aggregate (Asteraceae), whose four taxa (D. clusii s.s., D. stiriacum, D. glaciale subsp. glaciale and D. glaciale subsp. calcareum) are differentiated geographically, ecologically (basiphilous versus silicicolous) and/or via their ploidy levels (diploid versus tetraploid). Here, we use DNA sequences (three plastid and one nuclear spacer) and AFLP fingerprinting data generated for 58 populations to infer phylogenetic relationships, origin of polyploids-whose ploidy level was confirmed by chromosomally calibrated DNA ploidy level estimates-and phylogeographic history. Taxonomic conclusions were informed, among others, by a Gaussian clustering method for species delimitation using dominant multilocus data. Based on molecular data we identified three lineages: (i) silicicolous diploid D. clusii s.s. in the Alps, (ii) silicicolous tetraploid D. stiriacum in the eastern Alps (outside the range of D. clusii s.s.) and the Carpathians and (iii) the basiphilous diploids D. glaciale subsp. glaciale (eastern Alps) and D. glaciale subsp. calcareum (northeastern Alps); each taxon was identified as distinct by the Gaussian clustering, but the separation of D. glaciale subsp. calcareum and D. glaciale subsp. glaciale was not stable, supporting their taxonomic treatment as subspecies. Carpathian and Alpine populations of D. stiriacum were genetically differentiated suggesting phases of vicariance, probably during the Pleistocene. The origin (autopolyploid versus allopolyploid) of D. stiriacum remained unclear. Doronicum glaciale subsp. calcareum was genetically and morphologically weakly separated from D. glaciale subsp. glaciale but exhibited significantly higher genetic diversity and rarity. This suggests that the more widespread D. glaciale subsp. glaciale originated from D. glaciale subsp. calcareum, which is restricted to a prominent Pleistocene refugium previously identified in other alpine plant species.

Reproductive interactions among cytotypes in their contact zones determine whether these cytotypes can co-exist and form stable contact zones or not. In autopolyploids, heteroploid cross-compatibilities might depend on parental ploidy, but tests of this hypothesis in autopolyploid systems with more than two ploidies are lacking. Here, we study Jacobaea carniolica, which comprises diploid, tetraploid, and hexaploid individuals regularly forming contact zones. Seeds obtained from in situ cross-pollinations within and among cytotypes were subjected to DNA flow cytometry and greenhouse germination experiments. Hybrid fitness and parental effects on hybrid fitness were tested with regression models comparing fitness parameters of early life stages. Irrespective of the direction of crosses, seed viability and seedling survival in diploid-polyploid crosses were substantially lower than in tetraploid-hexaploid crosses. In contrast, seedling growth traits indicated neither transgressive character expression nor any selection against hybrid offspring. Congruent with a model of genome dosage effects, these traits differed between reciprocal crosses, especially of diploids and tetraploids, where trait values resembled those of the maternal parent. The strong effect of parental ploidy on offspring fitness in heteroploid crosses may cause contact zones involving exclusively polyploid cytotypes to be less stable over longer terms than those involving diploids and polyploids.

High mountain areas above the alpine zone are, despite the low-temperature conditions, inhabited by evolutionary and functionally differing organism groups. We compared the abundance and species richness of vascular plants, oribatid mites, springtails, spiders, and beetles, as well as bacterial and methanogenic archaeal prokaryotes (only abundance), at 100 m vertical intervals from 2,700-3,400 m in the Central Alps. We hypothesized that the less mobile microarthropods and microorganisms are more determined by and respond in similar ways to soil properties as do vascular plants. In contrast, we expected the more mobile surface-dwelling groups to forage also in places devoid of vegetation and thus to show patterns that deviate from that of vascular plants. Surprisingly, the observed patterns were diametrically opposed to our expectations: soil-living oribatid mites and springtails showed high individual numbers at high elevations, even where vascular plants barely occurred. Springtails also showed a rather constant species richness throughout the entire gradient. In contrast, patterns of surface-dwelling organisms and of archaeal prokaryotes did not differ significantly from vascular plants, because of either comparable climate sensitivity or their dependency on vegetated habitats. This study may serve as a baseline to estimate the risks of biodiversity losses in response to climate change across different biotic ecosystem components and to explore the potential and limitations of vascular plants as proxy for other organism groups that are far more challenging to monitor.

Aim We test for the congruence between allele-based range boundaries (break zones) in silicicolous alpine plants and species-based break zones in the silicicolous flora of the European Alps. We also ask whether such break zones coincide with areas of large elevational variation. Location The European Alps. Methods On a regular grid laid across the entire Alps, we determined areas of allele- and species-based break zones using respective clustering algorithms, identifying discontinuities in cluster distributions (breaks), and quantifying integrated break densities (break zones). Discontinuities were identified based on the intra-specific genetic variation of 12 species and on the floristic distribution data from 239 species, respectively. Coincidence between the two types of break zones was tested using Spearman's correlation. Break zone densities were also regressed on topographical complexity to test for the effect of elevational variation. Results We found that two main break zones in the distribution of alleles and species were significantly correlated. Furthermore, we show that these break zones are in topographically complex regions, characterized by massive elevational ranges owing to high mountains and deep glacial valleys. We detected a third break zone in the distribution of species in the eastern Alps, which is not correlated with topographic complexity, and which is also not evident from allelic distribution patterns. Species with the potential for long-distance dispersal tended to show larger distribution ranges than short-distance dispersers. Main conclusions We suggest that the history of Pleistocene glaciations is the main driver of the congruence between allele-based and species-based distribution patterns, because occurrences of both species and alleles were subject to the same processes (such as extinction, migration and drift) that shaped the distributions of species and genetic lineages. Large elevational ranges have had a profound effect as a dispersal barrier for alleles during post-glacial immigration. Because plant species, unlike alleles, cannot spread via pollen but only via seed, and thus disperse less effectively, we conclude that species break zones are maintained over longer time spans and reflect more ancient patterns than allele break zones.

BACKGROUND AND AIMS: Seeds of epiphytes must land on branches with suitable substrates and microclimates to germinate and for the resulting seedlings to survive. It is important to understand the fate of seeds and seedlings in order to model populations, but this is often neglected when only established plants are included in analyses. METHODS: The seeds of five bromeliad species were exposed to different canopy positions in a Mexican montane forest, and germination and early seedling survival were recorded. Additionally, the survival of naturally dispersed seedlings was monitored in a census over 2·5 years. Survival analysis, a procedure rarely used in plant ecology, was used to study the influence of branch characteristics and light on germination and seedling survival in natural and experimental populations. KEY RESULTS: Experimental germination percentages ranged from 7·2 % in Tillandsia deppeana to 33·7 % in T. juncea, but the seeds of T. multicaulis largely failed to germinate. Twenty months after exposure between 3·5 and 9·4 % of the seedlings were still alive. There was no evidence that canopy position affected the probability of germination, but time to germination was shorter in less exposed canopy positions indicating that higher humidity accelerates germination. More experimental seedlings survived when canopy openness was high, whereas survival in census-seedlings was influenced by moss cover. While mortality decreased steadily with age in juveniles of the atmospheric Tillandsia, in the more mesomorphic Catopsis sessiliflora mortality increased dramatically in the dry season. CONCLUSIONS: Seedling mortality, rather than the failure to germinate, accounts for the differential distribution of epiphytes within the canopy studied. With few safe sites to germinate and high seedling mortality, changes of local climate may affect epiphyte populations primarily through their seedling stage.