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International audience; Aim: Land use is the most pervasive driver of biodiversity loss. Predicting its impact on species richness (SR) is often based on indicators of habitat loss. However, the degradation of habitats, especially through land-use intensification, also affects species. Here, we evaluate whether an integrative metric of land-use intensity, the human appropriation of net primary production, is correlated with the decline of SR in used landscapes across the globe. Location Global. Time period Present. Major taxa studied Birds, mammals and amphibians. Methods: Based on species range maps (spatial resolution: 20 km × 20 km) and an area-of-habitat approach, we calibrated a “species–energy model” by correlating the SR of three groups of vertebrates with net primary production and biogeographical covariables in “wilderness” areas (i.e., those where available energy is assumed to be still at pristine levels). We used this model to project the difference between pristine SR and the SR corresponding to the energy remaining in used landscapes (i.e., SR loss expected owing to human energy extraction outside wilderness areas). We validated the projected species loss by comparison with the realized and impending loss reconstructed from habitat conversion and documented by national Red Lists. Results Species–energy models largely explained landscape-scale variation of mapped SR in wilderness areas (adjusted R2-values: 0.79–0.93). Model-based projections of SR loss were lower, on average, than reconstructed and documented ones, but the spatial patterns were correlated significantly, with stronger correlation in mammals (Pearson's r = 0.68) than in amphibians (r = 0.60) and birds (r = 0.57). Main conclusions: Our results suggest that the human appropriation of net primary production is a useful indicator of heterotrophic species loss in used landscapes, hence we recommend its inclusion in models based on species–area relationships to improve predictions of land-use-driven biodiversity loss.

Aim: Species' ecological traits influence their spatial genetic patterns. Bedrock preference strongly shapes the phylogeography of alpine plants, but its interactions with other ecological traits have rarely been disentangled. Here, we explore whether dispersal ability and degree of habitat specialization account for divergent postglacial expansion patterns of high-elevation plants in spite of similar bedrock preference. Location: The Pyrenees, southwestern Europe. Taxon: Cirsium glabrum (Asteraceae), Salix pyrenaica (Salicaceae) and Silene borderei (Caryophyllaceae). Methods: Phylogenetic, genetic structure and demographic modelling analyses based on restriction-site- associated DNA sequencing (RADseq) data from a range-wide populational sampling were conducted. Occurrence data and environmental variables were used to construct species distribution models, which were projected under current and Last Glacial Maximum conditions, and were combined with RADseq data to reconstruct the postglacial history of the study species. The degree of habitat specialization of each species was estimated based on the plant communities within which they occur, and their climatic niche breadth. Results: Salix pyrenaica, which occupies a broad range of habitats, shows a high level of range filling, a blurred genetic structure and an admixture cline between the two main genetic groups, congruent with rapid postglacial expansion. The microsite specialists C. glabrum and S. borderei exhibit a strong genetic structure and low levels of range filling, indicative of slow postglacial expansion. The good disperser C. glabrum shows higher levels of admixture between genetic groups and weaker population differentiation than the poor disperser S. borderei. Main Conclusions: Factors other than bedrock preference have a strong impact on the postglacial range dynamics of high-elevation species. Habitat specialization plays an important role, allowing species occupying a broad range of habitats to more rapidly expand their ranges after environmental change. The effect of dispersal ability is lower than expected for the study species.

Deciduous forests form the dominant natural vegetation of Europe today, but were restricted to small refugia during Pleistocene cold stages, implying an evolutionary past shaped by recurrent range contractions and expansions. Cold-stage forest refugia were probably widespread in southern and central Europe, with the northwestern Balkan Peninsula being of particular importance. However, the actual number and location of deciduous forest refugia, as well as the connections between them, remain disputed. Here, we address the evolutionary dynamics of the deciduous forest understorey species Euphorbia carniolica as a proxy for past forest dynamics. To do so, we obtained genomic and morphometric data from populations representing the species’ entire range, investigated phylogenetic position and intraspecific genetic variation, tested explicit demographic scenarios and applied species distribution models. Our data support two disjoint groups linked to separate refugia on the northwestern and central Balkan Peninsula. We find that genetic differentiation between groups started in the early Pleistocene via vicariance, suggesting a larger distribution in the past. Both refugia acted as sources for founder events to the southeastern Alps and the Carpathians, whereas the latter were likely colonized before the last cold stage. In line with traditional views on the pre-Pleistocene origin of many southeastern European deciduous forest species, the origin of E. carniolica was dated to the late Pliocene. The fact that E. carniolica evolved at a time when a period of continuous forestation was ending in much of Eurasia provides an interesting biogeographical perspective on the past links between Eurasian deciduous forests and their biota.

Mountain plant species are changing their ranges in response to global warming. However, these shifts vary tremendously in rate, extent and direction. The reasons for this variation are yet poorly understood. A process potentially important for mountain plant re-distribution is a competition between colonizing species and the resident vegetation. Here, we focus on the impact of this process using the recent elevational shift of the sedge Carex humilis in the northern Italian Alps as a model system. We repeated and extended historical sampling (conducted in 1976) of the species in the study region. We used the historical distribution data and historical climatic maps to parameterize a species distribution model (SDM) and projected the potential distribution of the species under current conditions. We compared the historical and the current re-survey for the species in terms of the cover of important potential competitor species as well as in terms of the productivity of the resident vegetation indicated by the Normalized Difference Vegetation Index (NDVI). We found that Carex humilis has shifted its leading range margin upward rapidly (51.2 m per decade) but left many sites that have become climatically suitable since 1976 according to the SDM uncolonized. These suitable but uncolonized sites show significantly higher coverage of all dwarf shrub species and higher NDVI than the sites occupied by the sedge. These results suggest that resistance of the resident vegetation against colonization of migrating species can indeed play an important role in controlling the re-distribution of mountain plants under climate change.

The abstract is available here: https://uscholar.univie.ac.at/o:1633096

Climate change and nitrogen (N) deposition are among the most important drivers of biodiversity change at a global scale. Although commonly considered separately in applied research, conservation, and policy development, both empirical evidence and modelling studies indicate that these two stressors may act synergistically and trigger a more pronounced biodiversity loss than each of them alone. Interactive effects thus need to be considered in biodiversity conservation. In this study, we propose a new method for jointly assessing risk from both stressors to habitats by combining climate envelope modelling and the evaluation of exceedance of empirical critical loads for eutrophication through N deposition. Based on this approach we can calculate a combined risk metric, which is the scaled exceedance of the N critical load and climatic thresholds in the three-dimensional space of N deposition, temperature, and water limitation. We use a habitat map to project this risk metric for 34 EUNIS habitat types across Austria. Resulting risk maps indicated that about 16 % of the area of these natural and semi-natural habitat types is currently at risk from either N exceedance, drought or warming. Using combinations of future climatic (RCP2.5 and RCP8.5) and N deposition (business as usual, current legislation and maximum feasible reduction) scenarios, we predict that the area at risk might increase to up to 19% until 2050, with risk from N-deposition decreasing and risk from climate change increasing. Combined impacts of both stressors affect only up to 2% of the entire evaluated area, but are much more frequent in particular habitat types such as oligotrophic bogs and subalpine to alpine grasslands. We conclude that this method provides a useful screening procedure to identify and compare areas and habitats under combined risk from both stressors and to thus support prioritization and decision making in biodiversity conservation.

The Balkan Peninsula as one of Europe’s biodiversity hotspots came into the focus of biogeographic research during the last decade, but the phylogeography of its mountain flora still remains widely...

Climate driven species’ range shifts may interfere with existing protected area (PA) networks, resulting in a mismatch between places where species are currently protected and places where they can thrive in the future. Here, we assess the climate-smartness of the Austrian PA network by focusing on endemic species’ climatic niches and their future representation within PAs. We calculated endemic species’ climatic niches and climate space available in PAs within their dispersal reach under current and future climates, with the latter represented by three climate change scenarios and three time-steps (2030, 2050, and 2080). Niches were derived from the area of occupancy of species and the extent of PAs, respectively, and calculated as bivariate density kernels on gradients of mean annual temperature and annual precipitation. We then computed climatic representation of species’ niches in PAs as the proportion of the species’ kernel covered by the PA kernel. We found that under both a medium (RCP 4.5) and severe (RCP 8.5) climate change scenario, representation of endemic species’ climatic niches by PAs will decrease to a sixth for animals and to a third for plants, on average, toward the end of the century. Twenty to thirty percent of Austrian endemic species will then have no representation of their climatic niches in PAs anymore. Species with larger geographical and wider elevational ranges will lose less climatic niche representation. The declining representation of climatic niches in PAs implies that, even if PAs may secure the persistence of a part of these endemics, only a small portion of intraspecific diversity of many species may be represented in PAs in the future. We discuss our findings in the context of the varied elevational gradients found in Austria and suggest that the most promising strategies for safeguarding endemic species’ evolutionary potential are to limit the magnitude of climate change and to reduce other pressures that additionally threaten their survival.

Global warming has been strongly accelerating in the last decades. Climate models tell us that this trend will continue in the future, accompanied by a marked decline in precipitation in Southern Europe, whereas the Alps will likely receive more winter and less summer precipitation. Climate factors and additionally nitrogen deposition and land-use changes have been identified as global change factors posing threats on high-mountain biodiversity, ecosystem stability and services. On the other hand, the characteristic micro-topographic variability of high mountain ecosystems may buffer them against global change impacts. Monitoring data from European mountain peaks show that changes in biodiversity patterns are closely related to rising temperatures. However, the effects of climate change on plant biodiversity differ significantly between temperate and Mediterranean biomes with species richness increases synchronously with warming in the former and richness decreases in the latter. The MediAlps project aimed at disentangling anthropogenic and natural factors underlying differential changes in plant species composition and richness observed on mountain summits in the European Alps and the Mediterranean biome at the local and regional spatial scale. Changes in plant species richness and composition and present land-use impact based on systematic field observations were recorded on long-term monitoring plots on 23 summits. Soil temperature, water potential and local dry nitrogen deposition were measured in situ. Topographic parameters were recorded with photogrammetric methods. At the regional level, climate data and regional nitrogen deposition data from online resources (CHELSA, EMEP) were used and past land-use impact was assessed via guideline-aided semi-structured interviews. (Generalized) linear mixed-effects models and structural equation models (SEM) were employed to assess the impact of these drivers on biodiversity changes. Furthermore, spatio-temporal analyses based on satellite images were conducted. Climate change is and will probably continue to be the main driver of plant biodiversity, species composition and their changes on mountain summits in both biomes. However, there are biome-specific differences with precipitation playing an important role in the Mediterranean biome in addition to temperature, which clearly is the most important single factor in the temperate biome. These changes will likely lead to a further thermophilisation in both biomes. The upwards movement of species from lower elevations will likely also result in a biotic homogenization of the vegetation, exacerbated by the decline of high-elevation endemic species. Species richness will likely continue to increase in the temperate biome until the “pay-off” of extinction debts or threshold effects of population size on extinction risks set in. With decreasing precipitation species richness in the Mediterranean biome will probably decline in the long run, too. Nevertheless, other anthropogenic drivers have to be considered as well, although their influence is arguably much smaller than that of climate variables, namely nitrogen deposition with a negative influence on species richness change in the temperate biome and present land-use with a positive one in the Mediterranean biome. In addition to MediAlps’ main focus on comparing multiple anthropogenic ecological drivers in the Alps with the Mediterranean mountains, the project substantially contributed to a spatially larger scaled long-term observation effort in the frame of the GLORIA (Global Observation Research Initiative in Alpine Environments) program.

The alpine life zone is expected to undergo major changes with ongoing climate change. While an increase of plant species richness on mountain summits has generally been found, competitive displacement may result in the long term. Here, we explore how species richness and surface cover types (vascular plants, litter, bare ground, scree and rock) changed over time on different bedrocks on summits of the European Alps. We focus on how species richness and turnover (new and lost species) depended on the density of existing vegetation, namely vascular plant cover. We analyzed permanent plots (1 m × 1 m) in each cardinal direction on 24 summits (24 × 4 × 4), with always four summits distributed along elevation gradients in each of six regions (three siliceous, three calcareous) across the European Alps. Mean summer temperatures derived from downscaled climate data increased synchronously over the past 30 years in all six regions. During the investigated 14 years, vascular plant cover decreased on siliceous bedrock, coupled with an increase in litter, and it marginally increased on higher calcareous summits. Species richness showed a unimodal relationship with vascular plant cover. Richness increased over time on siliceous bedrock but slightly decreased on calcareous bedrock due to losses in plots with high plant cover. Our analyses suggest contrasting and complex processes on siliceous versus calcareous summits in the European Alps. The unimodal richness-cover relationship and species losses at high plant cover suggest competition as a driver for vegetation change on alpine summits.

Climate and land‐use change jointly affect the future of biodiversity. Yet, biodiversity scenarios have so far concentrated on climatic effects because forecasts of land use are rarely available at appropriate spatial and thematic scales. Agent‐based models (ABMs) represent a potentially powerful but little explored tool for establishing thematically and spatially fine‐grained land‐use scenarios. Here, we use an ABM parameterized for 1,329 agents, mostly farmers, in a Central European model region, and simulate the changes to land‐use patterns resulting from their response to three scenarios of changing socio‐economic conditions and three scenarios of climate change until the mid of the century. Subsequently, we use species distribution models to, first, analyse relationships between the realized niches of 832 plant species and climatic gradients or land‐use types, respectively, and, second, to project consequent changes in potential regional ranges of these species as triggered by changes in both the altered land‐use patterns and the changing climate. We find that both drivers determine the realized niches of the studied plants, with land use having a stronger effect than any single climatic variable in the model. Nevertheless, the plants' future distributions appear much more responsive to climate than to land‐use changes because alternative future socio‐economic backgrounds have only modest impact on land‐use decisions in the model region. However, relative effects of climate and land‐use changes on biodiversity may differ drastically in other regions, especially where landscapes are still dominated by natural or semi‐natural habitat. We conclude that agent‐based modelling of land use is able to provide scenarios at scales relevant to individual species distribution and suggest that coupling ABMs with models of species' range change should be intensified to provide more realistic biodiversity forecasts., Land‐use and climate change will have important effects on the future of biodiversity. We predict changes to the potential distribution of vascular plants in response to these two drivers in a region of the Austrian Alps. We therefore combine agent‐based modelling of land owners' decisions with species distribution modelling. We find that both drivers determine the realized niches of the studied plants, with land use having a stronger effect than any single climatic variable in the model. Nevertheless, the plants' future distributions appear more responsive to climate than to land‐use changes.

Distribution modelling of vascular plants, butterflies and grasshoppers in central Europe suggests that habitat-based conservation strategies will be insufficient to save species from regional extinction under twenty-first-century climate change. Anthropogenic habitat fragmentation represents a major obstacle to species shifting their range in response to climate change1. Conservation measures to increase the (meta-)population capacity2 and permeability of landscapes3 may help but the effectiveness of such measures in a warming climate has rarely been evaluated. Here, we simulate range dynamics of 51 species from three taxonomic groups (vascular plants, butterflies and grasshoppers) in Central Europe as driven by twenty-first-century climate scenarios and analyse how three habitat-based conservation strategies (establishing corridors, improving the landscape matrix, and protected area management) modify species’ projected range size changes. These simulations suggest that the conservation strategies considered are unable to save species from regional extinction. For those persisting, they reduce the magnitude of range loss in lowland but not in alpine species. Protected area management and corridor establishment are more effective than matrix improvement. However, none of the conservation strategies evaluated could fully compensate the negative impact of climate change for vascular plants, butterflies or grasshoppers in central Europe.

Mountain plant species shift their elevational ranges in response to climate change. However, to what degree these shifts lag behind current climate change, and to what extent delayed extinctions and colonizations contribute to these shifts, are under debate. Here, we calculate extinction debt and colonization credit of 135 species from the European Alps by comparing species distribution models with 1576 re-surveyed plots. We find extinction debt in 60% and colonization credit in 38% of the species, and at least one of the two in 93%. This suggests that the realized niche of very few of the 135 species fully tracks climate change. As expected, extinction debts occur below and colonization credits occur above the optimum elevation of species. Colonization credits are more frequent in warmth-demanding species from lower elevations with lower dispersal capability, and extinction debts are more frequent in cold-adapted species from the highest elevations. Local extinctions hence appear to be already pending for those species which have the least opportunity to escape climate warming., Nature Communications, 10, ISSN:2041-1723

Species distribution models (SDMs) are used to project how suitable ranges of species shift under a warming climate. Conservation management, however, commonly targets habitat types rather than individual species. Such habitat types are often defined by the co-occurrence of a set of characteristic species. Here, we develop a co-occurrence-based index (CRI); which measures how the representation of habitat types in a particular area may change in a future climate. The index is based on stacking projections of distribution models of characteristic species and accounts for changes both in potential range size of each species individually and in spatial range overlap among characteristic species, i.e. co-occurrence patterns. We illustrate the approach by modelling the changing representation of 68 habitat types in Austria under two different climate scenarios. We base index calculations on SDM projections under either the assumption of unrestricted mobility (‘full-dispersal’) or of complete immobility (‘no-dispersal’) of individual species. Moreover, we compare results to those achieved with a simpler occurrence-based index (OI); which only accounts for change in species’ range sizes. All three alternative index calculations suggest that most modelled habitat types will lose area (in particular mires, wetlands and siliceous alpine grasslands) and only a minority will profit from a warming climate (in particular forests of dry and warm sites). ‘Full-dispersal’ CRI and OI are closely, but not perfectly correlated. Importantly, for more than half of the habitat types, accounting for changing co-occurrence patterns amplifies projected losses. The ‘no-dispersal’ CRI (CRInd) delivers the most alarming projections, indicating considerable spatial turn-over of sites suitable to the habitat types. Taken together, our results suggest that modelling re-distribution of habitat types which are defined by species combinations needs to account for modifications of co-occurrence patterns. Moreover, conservation should acknowledge that novel combinations of species will likely emerge under a warming climate.

Many studies report that mountain plant species are shifting upward in elevation. However, the majority of these reports focus on shifts of upper limits. Here, we expand the focus and simultaneously analyze changes of both range limits, optima, and abundances of 183 mountain plant species. We therefore resurveyed 1,576 vegetation plots first recorded before 1970 in the European Alps. We found that both range limits and optima shifted upward in elevation, but the most pronounced trend was a mean increase in species abundance. Despite huge species-specific variation, range dynamics showed a consistent trend along the elevational gradient: Both range limits and optima shifted upslope faster the lower they were situated historically, and species’ abundance increased more for species from lower elevations. Traits affecting the species’ dispersal and persistence capacity were not related to their range dynamics. Using indicator values to stratify species by their thermal and nutrient demands revealed that elevational ranges of thermophilic species tended to expand, while those of cold-adapted species tended to contract. Abundance increases were strongest for nutriphilous species. These results suggest that recent climate warming interacted with airborne nitrogen deposition in driving the observed dynamics. So far, the majority of species appear as “winners” of recent changes, yet “losers” are overrepresented among high-elevation, cold-adapted species with low nutrient demands. In the decades to come, high-alpine species may hence face the double pressure of climatic changes and novel, superior competitors that move up faster than they themselves can escape to even higher elevations.

Aim Interspecific hybridization can promote invasiveness of alien species. In many regions of the world, public and domestic gardens contain a huge pool of non‐native plants. Climate change may relax constraints on their naturalization and hence facilitate hybridization with related species in the resident flora. Here, we evaluate this possible increase in hybridization risk by predicting changes in the overlap of climatically suitable ranges between a set of garden plants and their congeners in the resident flora. Location Europe. Methods From the pool of alien garden plants, we selected those which (1) are not naturalized in Europe, but established outside their native range elsewhere in the world; (2) belong to a genus where interspecific hybridization has been previously reported; and (3) have congeners in the native and naturalized flora of Europe. For the resulting set of 34 alien ornamentals as well as for 173 of their European congeners, we fitted species distribution models and projected suitable ranges under the current climate and three future climate scenarios. Changes in range overlap between garden plants and congeners were then assessed by means of the true skill statistic. Results Projections suggest that under a warming climate, suitable ranges of garden plants will increase, on average, while those of their congeners will remain constant or shrink, at least under the more severe climate scenarios. The mean overlap in ranges among congeners of the two groups will decrease. Variation among genera is pronounced; however, and for some congeners, range overlap is predicted to increase significantly. Main conclusions Averaged across all modelled species, our results do not indicate that hybrids between potential future invaders and resident species will emerge more frequently in Europe when climate warms. These average trends do not preclude, however, that hybridization risk may considerably increase in particular genera.

Oceanic islands are vulnerable ecosystems and their flora has been under pressure since the arrival of the first humans. Human activities and both deliberately and inadvertently introduced biota have had and continue to have a severe impact on island endemic plants. The number of alien plants has increased nearly linearly on many islands, perhaps resulting in extinction‐based saturation of island floras. Here, we provide evidence for such a scenario in Alejandro Selkirk, Robinson Crusoe Islands (Archipelago Juan Fernández, Chile). We compared species richness and species composition of historical vegetation samples from 1917 with recent ones from 2011. Changes in species’ relative occurrence frequency were related to their taxonomic affiliation, dispersal mode, distribution status, and humidity and temperature preferences. While total species richness of vascular plants remained relatively similar, species composition changed significantly. Plants endemic to the Robinson Crusoe Islands declined, exotic species increased substantially within the period of ca. 100 years. Further, the relative occurrence frequency of plants with preferences for very warm and humid climate decreased, while the opposite was found for plants preferring drier and colder environments. Potential drivers responsible for this dramatic shift in the vegetation within only one century might have been the large goat population affecting especially small populations of endemic plants and climatic changes. Taking into account a substantial extinction debt, we expect further shifts in the vegetation of this small oceanic island toward alien plants. This would have significant negative consequences on global biodiversity, considering that island floras contribute substantially to global plant species richness due to their high proportion of endemics.

Withstanding extinction while facing rapid climate change depends on a species' ability to track its ecological niche or to evolve a new one. Current methods that predict climate-driven species' range shifts use ecological modelling without eco-evolutionary dynamics. Here we present an eco-evolutionary forecasting framework that combines niche modelling with individual-based demographic and genetic simulations. Applying our approach to four endemic perennial plant species of the Austrian Alps, we show that accounting for eco-evolutionary dynamics when predicting species' responses to climate change is crucial. Perennial species persist in unsuitable habitats longer than predicted by niche modelling, causing delayed range losses; however, their evolutionary responses are constrained because long-lived adults produce increasingly maladapted offspring. Decreasing population size due to maladaptation occurs faster than the contraction of the species range, especially for the most abundant species. Monitoring of species' local abundance rather than their range may likely better inform on species' extinction risks under climate change., Environmental niche models are often used to predict species responses to climate change but they neglect the potential for evolutionary responses. Here, Cotto et al. develop a model incorporating demographic processes and evolutionary dynamics and show that perennial alpine plants persist in unsuitable habitats but produce maladapted offspring.

Aim:Plant invasions often follow initial introduction with a considerable delay. The current non-native flora of a region may hence contain species that are not yet naturalized but may become so in the future, especially if climate change lifts limitations on species spread. In Europe, non-native garden plants represent a huge pool of potential future invaders. Here, we evaluate the naturalization risk from this species pool and how it may change under a warmer climate.Location:Europe.Methods:We selected all species naturalized anywhere in the world but not yet in Europe from the set of non-native European garden plants. For this subset of 783 species, we used species distribution models to assess their potential European ranges under different scenarios of climate change. Moreover, we defined geographical hotspots of naturalization risk from those species by combining projections of climatic suitability with maps of the area available for ornamental plant cultivation.Results:Under current climate, 165 species would already find suitable conditions in > 5% of Europe. Although climate change substantially increases the potential range of many species, there are also some that are predicted to lose climatically suitable area under a changing climate, particularly species native to boreal and Mediterranean biomes. Overall, hotspots of naturalization risk defined by climatic suitability alone, or by a combination of climatic suitability and appropriate land cover, are projected to increase by up to 102% or 64%, respectively.Main conclusions:Our results suggest that the risk of naturalization of European garden plants will increase with warming climate, and thus it is very likely that the risk of negative impacts from invasion by these plants will also grow. It is therefore crucial to increase awareness of the possibility of biological invasions among horticulturalists, particularly in the face of a warming climate. published

The authors incorporate intraspecific variation into a dynamic range model to predict the consequences of twenty-first century warming on six European alpine plants. As well as overall range loss, their model predicts a decrease in the frequency of warm-adapted haplotypes in five out of six species. Modelling of climate-driven range shifts commonly treats species as ecologically homogeneous units. However, many species show intraspecific variation of climatic niches and theory predicts that such variation may lead to counterintuitive eco-evolutionary dynamics. Here, we incorporate assumed intraspecific niche variation into a dynamic range model and explore possible consequences for six high-mountain plant species of the European Alps under scenarios of twenty-first century climate warming. At the species level, the results indicate massive range loss independent of intraspecific variation. At the intraspecific level, the model predicts a decrease in the frequency of warm-adapted haplotypes in five species. The latter effect is probably driven by a combination of leading-edge colonization and priority effects within the species' elevational range and was weakest when leading-edge expansion was constrained by mountain topography The resulting maladaptation may additionally increase the risk that alpine plants face from shrinkage of their ranges in a warming climate.