A climate-induced tree species bottleneck for forest management in Europe.

Large pulses of tree mortality have ushered in a major reorganization of Europe's forest ecosystems. To initiate a robust next generation of trees, the species that are planted today need to be climatically suitable throughout the entire twenty-first century. Here we developed species distribution models for 69 European tree species based on occurrence data from 238,080 plot locations to investigate the option space for current forest management in Europe. We show that the average pool of tree species continuously suitable throughout the century is smaller than that under current and end-of-century climate conditions, creating a tree species bottleneck for current management. If the need for continuous climate suitability throughout the lifespan of a tree planted today is considered, climate change shrinks the tree species pool available to management by between 33% and 49% of its current values (40% and 54% of potential end-of-century values), under moderate (Representative Concentration Pathway 2.6) and severe (Representative Concentration Pathway 8.5) climate change, respectively. This bottleneck could have strong negative impacts on timber production, carbon storage and biodiversity conservation, as only 3.18, 3.53 and 2.56 species of high potential for providing these functions remain suitable throughout the century on average per square kilometre in Europe. Our results indicate that the option space for silviculture is narrowing substantially because of climate change and that an important adaptation strategy in forestry-creating mixed forests-might be curtailed by widespread losses of climatically suitable tree species.

Post-glacial range formation of temperate forest understorey herbs - Insights from a spatio-temporally explicit modelling approach.

Aim: Our knowledge of Pleistocene refugia and post-glacial recolonization routes of forest understorey plants is still very limited. The geographical ranges of these species are often rather narrow and show highly idiosyncratic, often fragmented patterns indicating either narrow and species-specific ecological tolerances or strong dispersal limitations. However, the relative roles of these factors are inherently difficult to disentangle. Location: Central and south-eastern Europe. Time period: 17,100 BP - present. Major taxa studied: Five understorey herbs of European beech forests: Aposeris foetida, Cardamine trifolia, Euphorbia carniolica, Hacquetia epipactis and Helleborus niger. Methods: We used spatio-temporally explicit modelling to reconstruct the post-glacial range dynamics of the five forest understorey herbs. We varied niche requirements, demographic rates and dispersal abilities across plausible ranges and simulated the spread of species from potential Pleistocene refugia identified by phylogeographical analyses. Then we identified the parameter settings allowing for the most accurate reconstruction of their current geographical ranges. Results: We found a largely homogenous pattern of optimal parameter settings among species. Broad ecological niches had to be combined with very low but non-zero rates of long-distance dispersal via chance events and low rates of seed dispersal over moderate distances by standard dispersal vectors. However, long-distance dispersal events, although rare, led to high variation among replicated simulation runs. Main conclusions: Small and fragmented ranges of many forest understorey species are best explained by a combination of broad ecological niches and rare medium- and long-distance dispersal events. Stochasticity is thus an important determinant of current species ranges, explaining the idiosyncratic distribution patterns of the study species despite strong similarities in refugia, ecological tolerances and dispersal abilities.

Human appropriation of net primary production as driver of change in landscape-scale vertebrate richness.

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 R 2-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.

Niche breadth explains the range size of European-centred butterflies, but dispersal ability does not.

Aim: The breadth of ecological niches and dispersal abilities have long been discussed as important determinants of species' range sizes. However, studies directly comparing the relative effects of both factors are rare, taxonomically biased and revealed inconsistent results. Location: Europe. Time Period: Cenozoic. Major Taxa: Butterflies, Lepidoptera. Methods: We relate climate, diet and habitat niche breadth and two indicators of dispersal ability, wingspan and a dispersal tendency index, to the global range size of 369 European-centred butterfly species. The relative effects of these five predictors and their variation across the butterfly phylogeny were assessed by means of phylogenetic generalized least squares models and phylogenetically weighted regressions respectively. Results: Climate niche breadth was the most important single predictor, followed by habitat and diet niche breadth, while dispersal tendency and wingspan showed no relation to species' range size. All predictors together explained 59% of the variation in butterfly range size. However, the effects of each predictor varied considerably across families and genera. Main Conclusions: Range sizes of European-centred butterflies are strongly correlated with ecological niche breadth but apparently independent of dispersal ability. The magnitude of range size-niche breadth relationships is not stationary across the phylogeny and is often negatively correlated across the different dimensions of the ecological niche. This variation limits the generalizability of range size-trait relationships across broad taxonomic groups.

Evidence for Glacial Refugia of the Forest Understorey Species Helleborus niger (Ranunculaceae) in the Southern as Well as in the Northern Limestone Alps.

Glacial refugia of alpine and subnival biota have been intensively studied in the European Alps but the fate of forests and their understory species in that area remains largely unclear. In order to fill this gap, we aimed at disentangling the spatiotemporal diversification of disjunctly distributed black hellebore Helleborus niger (Ranunculaceae). We applied a set of phylogeographic analyses based on restriction-site associated DNA sequencing (RADseq) data and plastid DNA sequences to a range-wide sampling of populations. These analyses were supplemented with species distribution models generated for the present and the Last Glacial Maximum (LGM). We used exploratory analyses to delimit genomically coherent groups and then employed demographic modeling to reconstruct the history of these groups. We uncovered a deep split between two major genetic groups with western and eastern distribution within the Southern Limestone Alps, likely reflecting divergent evolution since the mid-Pleistocene in two glacial refugia situated along the unglaciated southern margin of the Alps. Long-term presence in the Southern Limestone Alps is also supported by high numbers of private alleles, elevated levels of nucleotide diversity and the species' modeled distribution at the LGM. The deep genetic divergence, however, is not reflected in leaf shape variation, suggesting that the morphological discrimination of genetically divergent entities within H. niger is questionable. At a shallower level, populations from the Northern Limestone Alps are differentiated from those in the Southern Limestone Alps in both RADseq and plastid DNA data sets, reflecting the North-South disjunction within the Eastern Alps. The underlying split was dated to ca. 0.1 mya, which is well before the LGM. In the same line, explicit tests of demographic models consistently rejected the hypothesis that the partial distribution area in the Northern Limestone Alps is the result of postglacial colonization. Taken together, our results strongly support that forest understory species such as H. niger have survived the LGM in refugia situated along the southern, but also along the northern or northeastern periphery of the Alps. Being a slow migrator, the species has likely survived repeated glacial-interglacial circles in distributional stasis while the composition of the tree canopy changed in the meanwhile.

High-Resolution Maps of Material Stocks in Buildings and Infrastructures in Austria and Germany.

The dynamics of societal material stocks such as buildings and infrastructures and their spatial patterns drive surging resource use and emissions. Two main types of data are currently used to map stocks, night-time lights (NTL) from Earth-observing (EO) satellites and cadastral information. We present an alternative approach for broad-scale material stock mapping based on freely available high-resolution EO imagery and OpenStreetMap data. Maps of built-up surface area, building height, and building types were derived from optical Sentinel-2 and radar Sentinel-1 satellite data to map patterns of material stocks for Austria and Germany. Using material intensity factors, we calculated the mass of different types of buildings and infrastructures, distinguishing eight types of materials, at 10 m spatial resolution. The total mass of buildings and infrastructures in 2018 amounted to ∼5 Gt in Austria and ∼38 Gt in Germany (AT: ∼540 t/cap, DE: ∼450 t/cap). Cross-checks with independent data sources at various scales suggested that the method may yield more complete results than other data sources but could not rule out possible overestimations. The method yields thematic differentiations not possible with NTL, avoids the use of costly cadastral data, and is suitable for mapping larger areas and tracing trends over time.

A socio-ecological model for predicting impacts of land-use and climate change on regional plant diversity in the Austrian Alps.

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.

Extinction debts and colonization credits of non-forest plants in the European Alps.

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.

Reconstructing geographical parthenogenesis: effects of niche differentiation and reproductive mode on Holocene range expansion of an alpine plant.

Asexual taxa often have larger ranges than their sexual progenitors, particularly in areas affected by Pleistocene glaciations. The reasons given for this 'geographical parthenogenesis' are contentious, with expansion of the ecological niche or colonisation advantages of uniparental reproduction assumed most important in case of plants. Here, we parameterized a spread model for the alpine buttercup Ranunculus kuepferi and reconstructed the joint Holocene range expansion of its sexual and apomictic cytotype across the European Alps under different simulation settings. We found that, rather than niche broadening or a higher migration rate, a shift of the apomict's niche towards colder conditions per se was crucial as it facilitated overcoming of topographical barriers, a factor likely relevant for many alpine apomicts. More generally, our simulations suggest potentially strong interacting effects of niche differentiation and reproductive modes on range formation of related sexual and asexual taxa arising from their differential sensitivity to minority cytotype disadvantage.

Will climate change increase hybridization risk between potential plant invaders and their congeners in Europe?

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.

Climate change will increase the naturalization risk from garden plants in Europe.

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.

A novel method to infer the origin of polyploids from Amplified Fragment Length Polymorphism data reveals that the alpine polyploid complex of Senecio carniolicus (Asteraceae) evolved mainly via autopolyploidy.

Despite its evolutionary and ecological relevance, the mode of polyploid origin has been notoriously difficult to be reconstructed from molecular data. Here, we present a method to identify the putative parents of polyploids and thus to infer the mode of their origin (auto- vs. allopolyploidy) from Amplified Fragment Length Polymorphism (AFLP) data. To this end, we use Cohen's d of distances between in silico polyploids, generated within a priori defined scenarios of origin from a priori delimited putative parental entities (e.g. taxa, genetic lineages), and natural polyploids. Simulations show that the discriminatory power of the proposed method increases mainly with increasing divergence between the lower-ploid putative ancestors and less so with increasing delay of polyploidization relative to the time of divergence. We apply the new method to the Senecio carniolicus aggregate, distributed in the European Alps and comprising two diploid, one tetraploid and one hexaploid species. In the eastern part of its distribution, the S. carniolicus aggregate was inferred to comprise an autopolyploid series, whereas for western populations of the tetraploid species, an allopolyploid origin involving the two diploid species was the most likely scenario. Although this suggests that the tetraploid species has two independent origins, other evidence (ribotype distribution, morphology) is consistent with the hypothesis of an autopolyploid origin with subsequent introgression by the second diploid species. Altogether, identifying the best among alternative scenarios using Cohen's d can be straightforward, but particular scenarios, such as allopolyploid origin vs. autopolyploid origin with subsequent introgression, remain difficult to be distinguished.

Uncertainty in predicting range dynamics of endemic alpine plants under climate warming.

Correlative species distribution models have long been the predominant approach to predict species' range responses to climate change. Recently, the use of dynamic models is increasingly advocated for because these models better represent the main processes involved in range shifts and also simulate transient dynamics. A well-known problem with the application of these models is the lack of data for estimating necessary parameters of demographic and dispersal processes. However, what has been hardly considered so far is the fact that simulating transient dynamics potentially implies additional uncertainty arising from our ignorance of short-term climate variability in future climatic trends. Here, we use endemic mountain plants of Austria as a case study to assess how the integration of decadal variability in future climate affects outcomes of dynamic range models as compared to projected long-term trends and uncertainty in demographic and dispersal parameters. We do so by contrasting simulations of a so-called hybrid model run under fluctuating climatic conditions with those based on a linear interpolation of climatic conditions between current values and those predicted for the end of the 21st century. We find that accounting for short-term climate variability modifies model results nearly as differences in projected long-term trends and much more than uncertainty in demographic/dispersal parameters. In particular, range loss and extinction rates are much higher when simulations are run under fluctuating conditions. These results highlight the importance of considering the appropriate temporal resolution when parameterizing and applying range-dynamic models, and hybrid models in particular. In case of our endemic mountain plants, we hypothesize that smoothed linear time series deliver more reliable results because these long-lived species are primarily responsive to long-term climate averages.