Data integration methods to account for spatial niche truncation effects in regional projections of species distribution.

Many species distribution models (SDMs) are built with precise but geographically restricted presence-absence data sets (e.g., a country) where only a subset of the environmental conditions experienced by a species across its range is considered (i.e., spatial niche truncation). This type of truncation is worrisome because it can lead to incorrect predictions e.g., when projecting to future climatic conditions belonging to the species niche but unavailable in the calibration area. Data from citizen-science programs, species range maps or atlases covering the full species range can be used to capture those parts of the species' niche that are missing regionally. However, these data usually are too coarse or too biased to support regional management. Here, we aim to (1) demonstrate how varying degrees of spatial niche truncation affect SDMs projections when calibrated with climatically truncated regional data sets and (2) test the performance of different methods to harness information from larger-scale data sets presenting different spatial resolutions to solve the spatial niche truncation problem. We used simulations to compare the performance of the different methods, and applied them to a real data set to predict the future distribution of a plant species (Potentilla aurea) in Switzerland. SDMs calibrated with geographically restricted data sets expectedly provided biased predictions when projected outside the calibration area or time period. Approaches integrating information from larger-scale data sets using hierarchical data integration methods usually reduced this bias. However, their performance varied depending on the level of spatial niche truncation and how data were combined. Interestingly, while some methods (e.g., data pooling, downscaling) performed well on both simulated and real data, others (e.g., those based on a Poisson point process) performed better on real data, indicating a dependency of model performance on the simulation process (e.g., shape of simulated response curves). Based on our results, we recommend to use different data integration methods and, whenever possible, to make a choice depending on model performance. In any case, an ensemble modeling approach can be used to account for uncertainty in how niche truncation is accounted for and identify areas where similarities/dissimilarities exist across methods.

Eco-genetic additivity of diploids in allopolyploid wild wheats.

Underpinnings of the distribution of allopolyploid species (hybrids with duplicated genome) along spatial and ecological gradients are elusive. As allopolyploid speciation combines the range of genetic and ecological characteristics of divergent diploids, allopolyploids initially show their additivity and are predicted to evolve differentiated ecological niches to establish in face of their competition. Here, we use four diploid wild wheats that differentially combined into four independent allopolyploid species to test for such additivity and assess the impact of ecological constraints on species ranges. Divergent genetic variation from diploids being fixed in heterozygote allopolyploids supports their genetic additivity. Spatial integration of comparative phylogeography and modelling of climatic niches supports ecological additivity of locally adapted diploid progenitors into allopolyploid species which subsequently colonised wide ranges. Allopolyploids fill suitable range to a larger extent than diploids and conservative evolution following the combination of divergent species appears to support their expansion under environmental changes.

Greater topoclimatic control of above- versus below-ground communities.

Assessing the degree to which climate explains the spatial distributions of different taxonomic and functional groups is essential for anticipating the effects of climate change on ecosystems. Most effort so far has focused on above-ground organisms, which offer only a partial view on the response of biodiversity to environmental gradients. Here including both above- and below-ground organisms, we quantified the degree of topoclimatic control on the occurrence patterns of >1,500 taxa and phylotypes along a c. 3,000 m elevation gradient, by fitting species distribution models. Higher model performances for animals and plants than for soil microbes (fungi, bacteria and protists) suggest that the direct influence of topoclimate is stronger on above-ground species than on below-ground microorganisms. Accordingly, direct climate change effects are predicted to be stronger for above-ground than for below-ground taxa, whereas factors expressing local soil microclimate and geochemistry are likely more important to explain and forecast the occurrence patterns of soil microbiota. Detailed mapping and future scenarios of soil microclimate and microhabitats, together with comparative studies of interacting and ecologically dependent above- and below-ground biota, are thus needed to understand and realistically forecast the future distribution of ecosystems.

Less favourable climates constrain demographic strategies in plants.

Correlative species distribution models are based on the observed relationship between species' occurrence and macroclimate or other environmental variables. In climates predicted less favourable populations are expected to decline, and in favourable climates they are expected to persist. However, little comparative empirical support exists for a relationship between predicted climate suitability and population performance. We found that the performance of 93 populations of 34 plant species worldwide - as measured by in situ population growth rate, its temporal variation and extinction risk - was not correlated with climate suitability. However, correlations of demographic processes underpinning population performance with climate suitability indicated both resistance and vulnerability pathways of population responses to climate: in less suitable climates, plants experienced greater retrogression (resistance pathway) and greater variability in some demographic rates (vulnerability pathway). While a range of demographic strategies occur within species' climatic niches, demographic strategies are more constrained in climates predicted to be less suitable.

Numerical ragweed pollen forecasts using different source maps: a comparison for France.

One of the key input parameters for numerical pollen forecasts is the distribution of pollen sources. Generally, three different methodologies exist to assemble such distribution maps: (1) plant inventories, (2) land use data in combination with annual pollen counts, and (3) ecological modeling. We have used six exemplary maps for all of these methodologies to study their applicability and usefulness in numerical pollen forecasts. The ragweed pollen season of 2012 in France has been simulated with the numerical weather prediction model COSMO-ART using each of the distribution maps in turn. The simulated pollen concentrations were statistically compared to measured values to derive a ranking of the maps with respect to their performance. Overall, approach (2) resulted in the best correspondence between observed and simulated pollen concentrations for the year 2012. It is shown that maps resulting from ecological modeling that does not include a sophisticated estimation of the plant density have a very low predictive skill. For inventory maps and the maps based on land use data and pollen counts, the results depend very much on the observational site. The use of pollen counts to calibrate the map enhances the performance of the model considerably.

Will climate change increase the risk of plant invasions into mountains?

Mountain ecosystems have been less adversely affected by invasions of non-native plants than most other ecosystems, partially because most invasive plants in the lowlands are limited by climate and cannot grow under harsher high-elevation conditions. However, with ongoing climate change, invasive species may rapidly move upwards and threaten mid-, and then high-elevation mountain ecosystems. We evaluated this threat by modeling the current and future habitat suitability for 48 invasive plant species in Switzerland and New South Wales, Australia. Both regions had contrasting climate interactions with elevation, resulting in possible different responses of species distributions to climate change. Using a species distribution modeling approach that combines data from two spatial scales, we built high-resolution species distribution models (≤ 250 m) that account for the global climatic niche of species and also finer variables depicting local climate and disturbances. We found that different environmental drivers limit the elevation range of invasive species in each of the two regions, leading to region-specific species responses to climate change. The optimal suitability for plant invaders is predicted to markedly shift from the lowland to the montane or subalpine zone in Switzerland, whereas the upward shift is far less pronounced in New South Wales where montane and subalpine elevations are already suitable. The results suggest that species most likely to invade high elevations in Switzerland will be cold-tolerant, whereas species with an affinity to moist soils are most likely to invade higher elevations in Australia. Other plant traits were only marginally associated with elevation limits. These results demonstrate that a more systematic consideration of future distributions of invasive species is required in conservation plans of not yet invaded mountainous ecosystems.

Influence of climate on the presence of colour polymorphism in two montane reptile species.

The coloration of ectotherms plays an important role in thermoregulation processes. Dark individuals should heat up faster and be able to reach a higher body temperature than light individuals and should therefore have benefits in cool areas. In central Europe, montane local populations of adder (Vipera berus) and asp viper (Vipera aspis) exhibit a varying proportion of melanistic individuals. We tested whether the presence of melanistic V. aspis and V. berus could be explained by climatic conditions. We measured the climatic niche position and breadth of monomorphic (including strictly patterned individuals) and polymorphic local populations, calculated their niche overlap and tested for niche equivalency and similarity. In accordance with expectations, niche overlap between polymorphic local populations of both species is high, and even higher than that of polymorphic versus monomorphic montane local populations of V. aspis, suggesting a predominant role of melanism in determining the niche of ectothermic vertebrates. However, unexpectedly, the niche of polymorphic local populations of both species is narrower than that of monomorphic ones, indicating that colour polymorphism does not always enable the exploitation of a greater variability of resources, at least at the intraspecific level. Overall, our results suggest that melanism might be present only when the thermoregulatory benefit is higher than the cost of predation.

Climate change may reveal currently unavailable parts of species' ecological niches.

The ability of climatic niche models to predict species extinction risks can be hampered if niches are incompletely quantified. This can occur when niches are estimated considering only currently available climatic conditions, disregarding the fact that climate change can open up portions of the fundamental niche that are currently inaccessible to species. Using a new metric, we estimate the prevalence of potential situations of fundamental niche truncation by measuring whether current ecological niche limits are contiguous to the boundaries of currently available climatic conditions for 24,944 species at the global scale in both terrestrial and marine realms and including animals and plants. We show that 12,172 (~49%) species are showing niche contiguity, particularly those inhabiting tropical ecosystems and the marine realm. Using niche expansion scenarios, we find that 86% of species showing niche contiguity could have a fundamental niche potentially expanding beyond current climatic limits, resulting in lower-yet still alarming-rates of predicted biodiversity loss, particularly within the tropics. Caution is therefore advised when forecasting future distributions of species presenting niche contiguity, particularly towards climatic limits that are predicted to expand in the future.

To what extent can we predict variation of bryophyte and tracheophyte community composition at fine spatial scale along an elevation gradient?

Mounting evidence points to the need for high-resolution climatic data in biodiversity analyses under global change. As we move to finer resolution, other factors than climate, including other abiotic variables and biotic interactions play, however, an increasing role, raising the question of our ability to predict community composition at fine scales. Focusing on two lineages of land plants, bryophytes and tracheophytes, we determine the relative contribution of climatic, non-climatic environmental drivers, spatial effects, community architecture and composition of one lineage to predict community composition of the other lineage, and how our ability to predict community composition varies along an elevation gradient. The relationship between community composition of one lineage and 68 environmental variables at 2-25 m spatial resolution, architecture and composition of the other lineage, and spatial factors, was investigated by hierarchical and variance partitioning across 413 2x2m plots in the Swiss Alps. Climatic data, although significant, contributed less to the model than any other variable considered. Community composition of one lineage, reflecting both direct interactions and unmeasured (hidden) abiotic factors, was the best predictor of community composition of the other lineage. Total explained variance substantially varied with elevation, underlining the fact that the strength of the species composition-environment relationship varies depending on environmental conditions. Total variance explained increased towards high elevation up to 50 %, with an increasing importance of spatial effects and vegetation architecture, pointing to increasing positive interactions and aggregated species distribution patterns in alpine environments. In tracheophytes, an increase of the contribution of non-climatic environmental factors was also observed at high elevation, in line with the hypothesis of a stronger environmental control under harsher conditions. Further improvements of our ability to predict changes in plant community composition may involve the implementation of historical variables and higher-resolution climatic data to better describe the microhabitat conditions actually experienced by organisms.

SWECO25: a cross-thematic raster database for ecological research in Switzerland.

Standard and easily accessible cross-thematic spatial databases are key resources in ecological research. In Switzerland, as in many other countries, available data are scattered across computer servers of research institutions and are rarely provided in standard formats (e.g., different extents or projections systems, inconsistent naming conventions). Consequently, their joint use can require heavy data management and geomatic operations. Here, we introduce SWECO25, a Swiss-wide raster database at 25-meter resolution gathering 5,265 layers. The 10 environmental categories included in SWECO25 are: geologic, topographic, bioclimatic, hydrologic, edaphic, land use and cover, population, transportation, vegetation, and remote sensing. SWECO25 layers were standardized to a common grid sharing the same resolution, extent, and geographic coordinate system. SWECO25 includes the standardized source data and newly calculated layers, such as those obtained by computing focal or distance statistics. SWECO25 layers were validated by a data integrity check, and we verified that the standardization procedure had a negligible effect on the output values. SWECO25 is available on Zenodo and is intended to be updated and extended regularly.

Monitoring of species' genetic diversity in Europe varies greatly and overlooks potential climate change impacts.

Genetic monitoring of populations currently attracts interest in the context of the Convention on Biological Diversity but needs long-term planning and investments. However, genetic diversity has been largely neglected in biodiversity monitoring, and when addressed, it is treated separately, detached from other conservation issues, such as habitat alteration due to climate change. We report an accounting of efforts to monitor population genetic diversity in Europe (genetic monitoring effort, GME), the evaluation of which can help guide future capacity building and collaboration towards areas most in need of expanded monitoring. Overlaying GME with areas where the ranges of selected species of conservation interest approach current and future climate niche limits helps identify whether GME coincides with anticipated climate change effects on biodiversity. Our analysis suggests that country area, financial resources and conservation policy influence GME, high values of which only partially match species' joint patterns of limits to suitable climatic conditions. Populations at trailing climatic niche margins probably hold genetic diversity that is important for adaptation to changing climate. Our results illuminate the need in Europe for expanded investment in genetic monitoring across climate gradients occupied by focal species, a need arguably greatest in southeastern European countries. This need could be met in part by expanding the European Union's Birds and Habitats Directives to fully address the conservation and monitoring of genetic diversity.

Predicting species distributions for conservation decisions.

Species distribution models (SDMs) are increasingly proposed to support conservation decision making. However, evidence of SDMs supporting solutions for on-ground conservation problems is still scarce in the scientific literature. Here, we show that successful examples exist but are still largely hidden in the grey literature, and thus less accessible for analysis and learning. Furthermore, the decision framework within which SDMs are used is rarely made explicit. Using case studies from biological invasions, identification of critical habitats, reserve selection and translocation of endangered species, we propose that SDMs may be tailored to suit a range of decision-making contexts when used within a structured and transparent decision-making process. To construct appropriate SDMs to more effectively guide conservation actions, modellers need to better understand the decision process, and decision makers need to provide feedback to modellers regarding the actual use of SDMs to support conservation decisions. This could be facilitated by individuals or institutions playing the role of 'translators' between modellers and decision makers. We encourage species distribution modellers to get involved in real decision-making processes that will benefit from their technical input; this strategy has the potential to better bridge theory and practice, and contribute to improve both scientific knowledge and conservation outcomes.

Dispersal routes reconstruction and the minimum cost arborescence problem.

We show that the dispersal routes reconstruction problem can be stated as an instance of a graph theoretical problem known as the minimum cost arborescence problem, for which there exist efficient algorithms. Furthermore, we derive some theoretical results, in a simplified setting, on the possible optimal values that can be obtained for this problem. With this, we place the dispersal routes reconstruction problem on solid theoretical grounds, establishing it as a tractable problem that also lends itself to formal mathematical and computational analysis. Finally, we present an insightful example of how this framework can be applied to real data. We propose that our computational method can be used to define the most parsimonious dispersal (or invasion) scenarios, which can then be tested using complementary methods such as genetic analysis.

Anthropogenic disturbance as a driver of microspatial and microhabitat segregation of cytotypes of Centaurea stoebe and cytotype interactions in secondary contact zones.

BACKGROUND AND AIMS: In a mixed-ploidy population, strong frequency-dependent mating will lead to the elimination of the less common cytotype, unless prezygotic barriers enhance assortative mating. However, such barriers favouring cytotype coexistence have only rarely been explored. Here, an assessment is made of the mechanisms involved in formation of mixed-ploidy populations and coexistence of diploid plants and their closely related allotetraploid derivates from the Centaurea stoebe complex (Asteraceae). METHODS: An investigation was made of microspatial and microhabitat distribution, life-history and fitness traits, flowering phenology, genetic relatedness of cytotypes and intercytotype gene flow (cpDNA and microsatellites) in six mixed-ploidy populations in Central Europe. KEY RESULTS: Diploids and tetraploids were genetically differentiated, thus corroborating the secondary origin of contact zones. The cytotypes were spatially segregated at all sites studied, with tetraploids colonizing preferentially drier and open microhabitats created by human-induced disturbances. Conversely, they were rare in more natural microsites and microsites with denser vegetation despite their superior persistence ability (polycarpic life cycle). The seed set of tetraploid plants was strongly influenced by their frequency in mixed-ploidy populations. Triploid hybrids originated from bidirectional hybridizations were extremely rare and almost completely sterile, indicating a strong postzygotic barrier between cytotypes. CONCLUSIONS: The findings suggest that tetraploids are later immigrants into already established diploid populations and that anthropogenic activities creating open niches favouring propagule introductions were the major factor shaping the non-random distribution and habitat segregation of cytotypes at fine spatial scale. Establishment and spread of tetraploids was further facilitated by their superior persistence through the perennial life cycle. The results highlight the importance of non-adaptive spatio-temporal processes in explaining microhabitat and microspatial segregation of cytotypes.

Spatial predictions of phylogenetic diversity in conservation decision making.

Considering genetic relatedness among species has long been argued as an important step toward measuring biological diversity more accurately, rather than relying solely on species richness. Some researchers have correlated measures of phylogenetic diversity and species richness across a series of sites and suggest that values of phylogenetic diversity do not differ enough from those of species richness to justify their inclusion in conservation planning. We compared predictions of species richness and 10 measures of phylogenetic diversity by creating distribution models for 168 individual species of a species-rich plant family, the Cape Proteaceae. When we used average amounts of land set aside for conservation to compare areas selected on the basis of species richness with areas selected on the basis of phylogenetic diversity, correlations between species richness and different measures of phylogenetic diversity varied considerably. Correlations between species richness and measures that were based on the length of phylogenetic tree branches and tree shape were weaker than those that were based on tree shape alone. Elevation explained up to 31% of the segregation of species rich versus phylogenetically rich areas. Given these results, the increased availability of molecular data, and the known ecological effect of phylogenetically rich communities, consideration of phylogenetic diversity in conservation decision making may be feasible and informative.

Resumen: Durante mucho tiempo se ha argumentado que la consideración de las relaciones genéticas entre especies es un paso importante hacia la medición más precisa de la diversidad biológica, en lugar de solo basarse en la riqueza de especies. Algunos investigadores han correlacionado medidas de la diversidad filogenética y de la riqueza de especies en una serie de sitios y sugieren que los valores de la diversidad filogenética no difieren suficientemente de los valores de riqueza de especies para justificar su inclusión en la planificación de la conservación. Comparamos las predicciones de riqueza de especies y 10 medidas de diversidad filogenética mediante la creación de modelos de distribución de 168 especies de una familia de plantas muy rica en especies, Proteaceae. Cuando utilizamos cantidades promedio de terrenos protegidos para comparar áreas seleccionadas con base en la riqueza de especies con áreas seleccionadas con base en la diversidad filogenética, las correlaciones entre riqueza de especies y las diferentes medidas de diversidad filogenética variaron considerablemente. Las correlaciones entre riqueza de especies y medidas que se basaron en la longitud de las ramas de los árboles filogenéticos y la forma del árbol fueron más débiles que las que se basaron solamente en la forma del árbol. La elevación explicó hasta 31% de la segregación de áreas ricas en especies versus las áreas filogenéticamente ricas. Dados estos resultados, la mayor disponibilidad de datos moleculares, y el efecto ecológico conocido de las comunidades filogenéticamente ricas, la consideración de la diversidad filogenética en la toma de decisiones de conservación puede ser factible e informativa.

Distance to native climatic niche margins explains establishment success of alien mammals.

One key hypothesis explaining the fate of exotic species introductions posits that the establishment of a self-sustaining population in the invaded range can only succeed within conditions matching the native climatic niche. Yet, this hypothesis remains untested for individual release events. Using a dataset of 979 introductions of 173 mammal species worldwide, we show that climate-matching to the realized native climatic niche, measured by a new Niche Margin Index (NMI), is a stronger predictor of establishment success than most previously tested life-history attributes and historical factors. Contrary to traditional climatic suitability metrics derived from species distribution models, NMI is based on niche margins and provides a measure of how distant a site is inside or, importantly, outside the niche. Besides many applications in research in ecology and evolution, NMI as a measure of native climatic niche-matching in risk assessments could improve efforts to prevent invasions and avoid costly eradications.

Multiple introductions boosted genetic diversity in the invasive range of black cherry (Prunus serotina; Rosaceae).

BACKGROUND AND AIMS: Black cherry (Prunus serotina) is a North American tree that is rapidly invading European forests. This species was introduced first as an ornamental plant, then it was massively planted by foresters in many countries, but its origins and the process of invasion remain poorly documented. Based on a genetic survey of both native and invasive ranges, the invasion history of black cherry was investigated by identifying putative source populations and then assessing the importance of multiple introductions on the maintenance of gene diversity. METHODS: Genetic variability and structure of 23 populations from the invasive range and 22 populations from the native range were analysed using eight nuclear microsatellite loci and five chloroplast DNA regions. KEY RESULTS: Chloroplast DNA diversity suggests there were multiple introductions from a single geographic region (the north-eastern United States). A low reduction of genetic diversity was observed in the invasive range for both nuclear and plastid genomes. High propagule pressure including both the size and number of introductions shaped the genetic structure in Europe and boosted genetic diversity. Populations from Denmark, The Netherlands, Belgium and Germany showed high genetic diversity and low differentiation among populations, supporting the hypothesis that numerous introduction events, including multiple individuals and exchanges between sites, have taken place during two centuries of plantation. CONCLUSIONS: This study postulates that the invasive black cherry has originated from east of the Appalachian Mountains (mainly the Allegheny plateau) and its invasiveness in north-western Europe is mainly due to multiple introductions containing high numbers of individuals.

Shift in cytotype frequency and niche space in the invasive plant Centaurea maculosa.

Polyploidy is often assumed to increase the spread and thus the success of alien plant species, but few empirical studies exist. We tested this hypothesis with Centaurea maculosa Lam., a species native to Europe and introduced into North America approximately 120 years ago where it became highly invasive. We analyzed the ploidy level of more than 2000 plants from 93 native and 48 invasive C. maculosa populations and found a pronounced shift in the relative frequency of diploid and tetraploid cytotypes. In Europe diploid populations occur in higher frequencies than tetraploids and only four populations had both cytotypes, while in North America diploid plants were found in only one mixed population and thus tetraploids clearly dominated. Our results showed a pronounced shift in the climatic niche between tetraploid populations in the native and introduced range toward drier climate in North America and a similar albeit smaller shift between diploids and tetraploids in the native range. The field data indicate that diploids have a predominately monocarpic life cycle, while tetraploids are often polycarpic. Additionally, the polycarpic life-form seems to be more prevalent among tetraploids in the introduced range than among tetraploids in the native range. Our study suggests that both ploidy types of C. maculosa were introduced into North America, but tetraploids became the dominant cytotype with invasion. We suggest that the invasive success of C. maculosa is partly due to preadaptation of the tetraploid cytotype in Europe to drier climate and possibly further adaptation to these conditions in the introduced range. The potential for earlier and longer seed production associated with the polycarpic life cycle constitutes an additional factor that may have led to the dominance of tetraploids over diploids in the introduced range.

Prediction of plant species distributions across six millennia.

The usefulness of species distribution models (SDMs) in predicting impacts of climate change on biodiversity is difficult to assess because changes in species ranges may take decades or centuries to occur. One alternative way to evaluate the predictive ability of SDMs across time is to compare their predictions with data on past species distributions. We use data on plant distributions, fossil pollen and current and mid-Holocene climate to test the ability of SDMs to predict past climate-change impacts. We find that species showing little change in the estimated position of their realized niche, with resulting good model performance, tend to be dominant competitors for light. Different mechanisms appear to be responsible for among-species differences in model performance. Confidence in predictions of the impacts of climate change could be improved by selecting species with characteristics that suggest little change is expected in the relationships between species occurrence and climate patterns.