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Predicting climate change impact on ecosystem structure and services is one of the most important challenges in ecology. Until now, plant species response to climate change has been described at the level of fixed plant functional types, an approach limited by its inflexibility as there is much interspecific functional variation within plant functional types. Considering a plant species as a set of functional traits greatly increases our possibilities for analysis of ecosystem functioning and carbon and nutrient fluxes associated therewith. Moreover, recently assembled large-scale databases hold comprehensive per-species data on plant functional traits, allowing a detailed functional description of many plant communities on Earth. Here, we show that plant functional traits can be used as predictors of vegetation response to climate warming, accounting in our test ecosystem (the species-rich alpine belt of Caucasus mountains, Russia) for 59% of variability in the per-species abundance relation to temperature. In this mountain belt, traits that promote conservative leaf water economy (higher leaf mass per area, thicker leaves) and large investments in belowground reserves to support next year’s shoot buds (root carbon content) were the best predictors of the species increase in abundance along with temperature increase. This finding demonstrates that plant functional traits constitute a highly useful concept for forecasting changes in plant communities, and their associated ecosystem services, in response to climate change.

1. Although the consequences of facilitation at individual and population levels are well known, the community-level consequences of these processes have received much less attention. In particular, the importance of facilitation in determining richness at the entire community level has seldom been evaluated. 2. In this study, we sampled 11 alpine plant communities along the southern Andes in South America, spanning from tropical (25°S) to sub-antarctic latitudes (55°S). Plant communities were dominated by cushion plants, a particular growth form that acts as a nurse plant for other plant species. Through rarefaction curves, we assessed the effectiveness of community sampling and estimated the number of species present within and outside cushions. Non-metric Multidimensional Scaling ordinations (NMDS) were used to assess differences between the species assemblages growing within and outside cushions. Finally, samples from cushions and open areas were combined in a single matrix accounting for the difference in cover between both microhabitats, and through rarefaction curves we assessed how many more species are added to the community due to the presence of cushions. 3. Samples taken within cushions always contained more species than equivalent samples from open areas. However, the magnitude of this difference varied among communities. NMDS ordination indicated that cushions generate species assemblages structurally different from those found in open areas. Inclusion of samples from cushion and open areas in synthetic analyses - where differences in cover were accounted for - indicated that the presence of cushions consistently increased species richness at the entire community level. The magnitude of these increases in species richness varied with habitat severity, with lower values at both extremes of the environmental severity gradient. 4.Synthesis. Facilitative interactions with cushion nurse plants along the high Andes of southern South America changed plant assemblage structure and increased species richness at the entire community level, indicating that facilitative interactions are pivotal in maintaining the diversity of these harsh environments.

Aim: Global warming is predicted to shift distributions of mountain species upwards, driven by a release from climatic restrictions at their upper distribution limit and increased biotic pressure at their lower distribution limit. In alpine ecosystems, which are characterized by large microclimatic diversity and sparse vegetation cover, the relative importance of abiotic and biotic drivers for species distribution is poorly understood. To disentangle abiotic and biotic mechanisms affecting distributions of alpine species, we investigated how alpine plant species with differing elevational ranges and frequency trends over the past century differ in their microhabitat distribution, and how they respond to neighbouring vegetation. Location: A total of 11 summits (2635-3410 m a.s.l.) in SE-Switzerland. Methods: We quantified the microscale abundance of 12 species in relation to biogeographic (frequency trend, i.e., change in occurrences over the past century, and elevational range on summits) and local microhabitat characteristics (temperature, substrate type). We assessed species size traits in relation to neighbouring vegetation characteristics to investigate possible neighbour interactions. Results: Species with increasing frequency on summits over the past century were most abundant on scree and warmer slopes. Species with negative or stable frequency trends on summits were more abundant on organic soil and colder slopes. The preferred microhabitats of the latter were rarest overall, decreased with increasing elevation, and had the most competitive neighbours. Size of one high-alpine specialist, Ranunculus glacialis was negatively related to cover of neighbouring vegetation, whereas other species showed no response to neighbours. Main conclusions: Long-term frequency trends of species correlate with their microhabitat association. Species with most negative frequency trends show preferences for the rarest microhabitat conditions, where they likely experience higher competitive pressure in a warming climate. This finding emphasizes the importance of characterizing microhabitat associations and microclimatic diversity to assess present and future distributions of alpine plant species.

The study of phylogenetic conservatism in alpine plant phenology is critical for predicting climate change impacts; currently we have a poor understanding of how phylogeny and climate factors interactively influence plant phenology. Therefore, we explored the influence of phylogeny and climate factors on flowering phenology in alpine meadows. For two different types of alpine plant communities, we recorded phenological data, including flowering peak, first flower budding, first flowering, first fruiting and the flowering end for 62 species over the course of 5 years (2008–2012). From sequences in two plastid regions, we constructed phylogenetic trees. We used Blomberg’s K and Pagel’s lambda to assess the phylogenetic signal in phenological traits and species’ phenological responses to climate factors. We found a significant phylogenetic signal in the date of all reproductive phenological events and in species’ phenological responses to weekly day length and temperature. The number of species in flower was strongly associated with the weekly day lengths and followed by the weekly temperature prior to phenological activity. Based on phylogenetic eigenvector regression (PVR) analysis, we found a highly shared influence of phylogeny and climate factors on alpine species flowering phenology. Our results suggest the phylogenetic conservatism in both flowering and fruiting phenology may depend on the similarity of responses to external environmental cues among close relatives.

Geranium erianthum is an alpine plant growing in dry habitats, which is distributed from East Asia to northern coastal regions of the northern Pacific. The ice-free area around the current Bering Strait (i.e., Beringia) had played an important role in range expansion into neighboring regions such as East Asia and North America for some alpine plants. However, recent studies suggest that some alpine plants in snowbed environment spread from East Asia to northern coastal regions of the northern Pacific. In this study, we investigated phylogenetic relationships and genetic differentiations among populations of G. erianthum and the related species using the chloroplast genome and single-nucleotide polymorphisms, to evaluate the alternative biogeographic hypotheses in which region of Beringia, British Columbia or East Asia is probable for its distributional origin. Range reconstruction based on phylogenetic tree of chloroplast genome indicated G. erianthum and related species originated in East Asia, from where G. erianthum migrated eastward into Beringia and British Columbia. In addition, nuclear genome-wide SNPs indicated that no significant genetic differentiation was detected between Japanese and Beringian populations. The lack of genetic differentiation suggests that the current range of G. erianthum resulted from rapid range expansion during the latter period of the last glacial era. Overall, the East Asian refugium hypothesis was applicable to the alpine plant G. erianthum in dry habitat, indicating that range expansion pattern from East Asia into the northern Pacific may be more common rather than limited for snowbed species.

Questions: How do phylogenetic and functional trait dispersions respond to multiple abiotic gradients? Are functional trait and phylogenetic dispersions coupled across different spatial scales? Does phylogenetic signal on functional trait data help to elucidate the degree to which phylogenetic information is providing novel information? Location: Three massifs in mediterranean-type climate zone of the high Andes, central Chile. Methods: We sampled plant species composition in 20 alpine sites above the tree line at three different spatial scales: plot (20 m × 20 m), subplot (2.4 m × 2.4 m) and cell (30 cm × 30 cm). Functional and phylogenetic mean pair-wise distances (MPD) calculated using data on six functional traits (maximum plant height, plant size, leaf area, specific leaf area, leaf dry matter content and leaf thickness) and a molecular phylogeny (rbcL and matK) were compared to the patterns expected under a null model to characterize the functional and phylogenetic dispersion along interacting elevation and potential solar radiation gradients. Results: Our results show that functional and phylogenetic dispersion were related and influenced by potential solar radiation, but the effect of this factor varied with elevation. Overdispersion was found in the most stressful sites, while clustering was observed where the conditions were milder, suggesting a relevant role of facilitation and competitive interactions, respectively. While Blomberg's K statistic indicated no phylogenetic signal for the studied plant traits, Pagel's λ indicated phylogenetic signal, but not of strong intensity (< 1), suggesting that the correlation between the functional and phylogenetic diversities was low and that additional unmeasured traits with phylogenetic signal were likely to be important in determining the structure of the studied communities. Conclusions: Our results support the hypothesis that biotic interactions modulated by environmental conditions are important for alpine plant community assembly. Moreover, they reinforce the notion that multiple processes shape community structure, and this can be elucidated by examining interacting environmental gradients, such as elevation and potential solar radiation, and taking into account multiple spatial scales. Our results reinforce the use of both functional and phylogenetic diversities simultaneously and discourage the use of phylogenetic diversity as a surrogate of functional structure.

Cryptic colouration is a common predation‐avoidance strategy in animals that is postulated to occur in plants, but few experimental studies have rigorously tested this hypothesis. We investigated the colouration of Corydalis benecincta, an alpine plant with remarkably dimorphic leaf colours (grey and green), based on a cost–benefit analysis. First we tested the premise that herbivores (Parnassius butterflies) cannot distinguish grey leaves from a scree background by spectrographic measurements and by estimating discriminability between leaves and scree using a butterfly colour vision model. Then we estimated the potential costs of inconspicuousness by comparing the photosynthetic performance and visual attractiveness to flower visitors of the two colour morphs. Finally, we examined the potential benefits of inconspicuousness by comparing damage, survivorship and female reproductive success. It is difficult for herbivores to distinguish grey‐coloured morphs against the background. This grey colour originates in a combination of anthocyanins and chlorophylls. The two colour morphs had similar photosynthetic performance, visual attractiveness and female reproductive success. However, grey morphs had significantly lower herbivore damage and higher survivorship. Grey leaves benefit C. benecincta by reducing herbivory with low investment in anthocyanin synthesis, and little cost on photosynthesis and mating opportunity. This cryptic colouration may have evolved through selection pressure imposed by visually foraging herbivores.

Aim In recent decades species ranges have shifted upwards in elevation and northwards in latitude. These shifts are commonly interpreted as a response to recent climate warming. However, several alternative hypotheses have been proposed to explain the elevational shifts, including increased deposition of atmospheric nitrogen, changes in precipitation and dispersal limitation. We evaluate these hypotheses and attempt to identify the dominant drivers for the observed shifts in the upper range limits of alpine plant species. Location European mountains from Svalbard to the southern Alps. Methods We assembled data on observed shifts in the upper range limit of alpine plants over 40 to 100 years on 114 mountains. We related the observed shifts to recent changes in temperature and precipitation and to recent deposition of atmospheric nitrogen. Changes in traits and habitat preferences of species in the summit assemblages were used to evaluate the potential role of different drivers. Results Seventy per cent of the species that showed a detectable change in their upper range limits between surveys shifted their range limits upwards. The same species tend to move up on different mountains. There are, however, large differences between mountains in the proportion of species shifting upwards. This proportion is not found to be statistically related to local changes in temperature. Correspondingly, warmth-demanding species did not move upward more frequently than expected by chance. Snow-bed species have become more common on summits. Main conclusions Our data do not support the idea that climate warming is the dominant factor causing the observed range shifts of alpine plant species on European mountains: first, the amount of change in species assemblages on the summits studied is not related statistically to the amount of climate warming; second, those species that have moved upwards are not particularly warmth demanding.

Aim Species distribution models (SDMs) are commonly used to forecast climate change impacts. These models, however, are subject to important assumptions and limitations. By integrating two independent but complementary methods, ensemble SDMs and statistical phylogeography, we addressed key assumptions and created robust assessments of climate change impacts on species distributions while improving the conservation value of these projections. Location North American cordillera. Methods This approach was demonstrated using the arctic-alpine plant Rhodiola integrifolia (Crassulaceae). SDMs were fitted to current and past climates using eight models, two thresholds and one to three climate data sets. These projections were combined to create a map of stable climate (refugia) since the Last Interglacial (124,000 kya). Five biogeographic hypotheses were developed based on the configuration of refugia and tested using statistical phylogeography. Projection of SDMs into the future was contingent on agreement across approaches; future projections (to 2085) used five climate data sets and two greenhouse gas scenarios. Results A multiple-refugia hypothesis was supported by both methods, confirming the assumption of niche conservatism in R. integrifolia and justifying the projection of SDMs onto future climates. Future projections showed substantial loss of climatically suitable habitat. Southern populations had the greatest losses, although the biogeographic scale of modelling may overpredict extinction risks in areas of topographic complexity. Past and future SDMs were assessed for novel values of climate variables; areas of novel climate were flagged as having higher uncertainty. Main conclusions Integrating molecular approaches with spatial analyses of species distributions under global change has great potential to improve conservation decision-making. Molecular tools can support and improve current methods for understanding the vulnerability of species to climate change and provide additional data upon which to base conservation decisions, such as prioritizing the conservation of areas of high genetic diversity to build evolutionary resiliency within populations.