Drivers Of Species Composition Research Articles

Beta diversity patterns in Andean rodents: current and historical factors as drivers of turnover and nestedness

Abstract The Andes are known to be one of the most species-rich regions on Earth, with their origination promoting formation of new habitats and acting as climatic barriers. This orogenic event was structured latitudinally with 3 major segments having different formation times and with different environmental and historical conditions. The Andean region has been historically important in the evolutionary history of small mammals and is considered the region of Earth with the highest values of rodent species richness and turnover. Therefore, the region provides an excellent opportunity to evaluate mechanisms involved in structuring rodent assemblages and beta diversity patterns. Here, we assessed spatial patterns of rodent beta diversity along the Andes Mountains, with specific goals of examining patterns of turnover and nestedness as 2 recognized processes that drive species composition. We explored the role of spatial, climate, and historical factors on beta diversity patterns using geographic range maps for 432 rodent species to construct a species presence–absence matrix. We used piecewise regressions to identify latitudinal breakpoints in turnover and nestedness, and conducted multiple regressions and variation-partitioning approaches to assess the contribution of each set of postulated drivers in shaping beta diversity patterns. Beta diversity decreases from north to south. The highest turnover was located at the Northern Andes (5°N and 10°S) and western-central Andes (20°S and 35°S). Nestedness was higher below 40°S and in some areas of the western-central Andes between 15° and 20°S. Turnover and nestedness registered breakpoints coincident with the Central and Southern Andes limit. Turnover explained most of the beta diversity for all segments, although within the Southern Andes there was also some contribution of nestedness. In all segments, beta diversity was mainly explained by spatial factors with minor contributions from spatially structured climate (Northern Andes), interaction of all factors (Central Andes), and the interaction between current climate and history (Southern Andes).

Drivers of species composition in arable‐weed communities of the Austrian–Hungarian borderland region: What is the role of the country?

AbstractQuestionsDue to their high ecological and agronomical variability, borderland regions offer an excellent opportunity to study assembly patterns. In this study we compared the influence of various factors on summer annual weed communities consisting of both native and introduced species.LocationThe borderland region of Austria and Hungary.MethodsWe assessed the abundance of weed species in 300 fields of six summer annual crops, and collected information on 26 background variables for each plot. We applied redundancy analysis (RDA) to estimate multivariate species responses and variation partitioning to compare the relative importance of three groups of variables (environmental variables, management variables, and country as a singleton group), and we also checked for statistical association between country and the predictors of the other two groups.ResultsThe full RDA model explained 22.02% of the variance in weed species composition. Variation partitioning showed that environment and management had similarly high (~8%) influence on weeds, while country had a modest yet substantial (~1%) effect, and there was relatively little overlap between the variance attributable to the three groups. Comparing the individual variables, country ranked third (after preceding crop, and actual crop). The effects of 15 further variables were also significant, including seven management, and seven environmental variables, as well as the location of the sampling plots within the fields. Comparisons between the countries showed that farming type, preceding crops, tillage system, tillage depth and field size were significantly different between the countries.ConclusionsCountry exhibited a small but significant influence on weed community composition, which could not be explained with easily accessible management and environmental variables. This suggests that the distinct historical agronomical background of the two countries, possibly involving some legacies of the former Iron Curtain period, still has an impact on the weed species composition of arable fields.

Regional flea and host assemblages form biogeographic, but not ecological, clusters: evidence for a dispersal-based mechanism as a driver of species composition.

We used data on the species composition of regional assemblages of fleas and their small mammalian hosts from 6 biogeographic realms and applied a novel method of step-down factor analyses (SDFA) and cluster analyses to identify biogeographic (across the entire globe) and ecological (within a realm across the main terrestrial biomes) clusters of these assemblages. We found that, at the global scale, the clusters of regional assemblage loadings on SDFA axes reflected well the assemblage distribution, according to the biogeographic realms to which they belong. At the global scale, the cluster topology, corresponding to the biogeographic realms, was similar between flea and host assemblages, but the topology of subtrees within realm-specific clusters substantially differed between fleas and hosts. At the scale of biogeographic realms, the distribution of regional flea and host assemblages did not correspond to the predominant biome types. Assemblages with similar loadings on SDFA axes were often situated in different biomes and vice versa. The across-biome, within-realm distributions of flea vs host assemblages suggested weak congruence between these distributions. Our results indicate that dispersal is a predominant mechanism of flea and host community assembly across large regions.

NO2 air pollution drives species composition, but tree traits drive species diversity of urban epiphytic lichen communities

Lichens serve as important bioindicators of air pollution in cities. Here, we studied the diversity of epiphytic lichens in the urban area of Munich, Bavaria, southern Germany, to determine which factors influence species composition and diversity. Lichen diversity was quantified in altogether 18 plots and within each, five deciduous trees were investigated belonging to on average three tree species (range 1–5). Of the 18 plots, two were sampled in control areas in remote areas of southern Germany. For each lichen species, frequency of occurrence was determined in 10 quadrats of 100 cm2 on the tree trunk. Moreover, the cover percentage of bryophytes was determined and used as a variable to represent potential biotic competition. We related our diversity data (species richness, Shannon index, evenness, abundance) to various environmental variables including tree traits, i.e. bark pH levels and species affiliation and air pollution data, i.e. NO2 and SO2 concentrations measured in the study plots. The SO2 levels measured in our study were generally very low, while NO2 levels were rather high in some plots. We found that the species composition of the epiphytic lichen communities was driven mainly by NO2 pollution levels and all of the most common species in our study were nitrophilous lichens. Low NO2 but high SO2 values were associated with high lichen evenness. Tree-level lichen diversity and abundance were mainly determined by tree traits, not air pollution. These results confirm that ongoing NO2 air pollution within cities is a major threat to lichen diversity, with non-nitrophilous lichens likely experiencing the greatest risk of local extinctions in urban areas in the future. Our study moreover highlights the importance of large urban green spaces for species diversity. City planners need to include large green spaces when designing urban areas, both to improve biodiversity and to promote human health and wellbeing.

Correction to: Indirect facilitation drives species composition and stability in drylands

The original version of this article unfortunately contained a mistake. The 2nd affiliation of Sonia Kefi was missing

Unraveling plant strategies in tidal marshes by investigating plant traits and environmental conditions

AbstractAimsTidal marsh vegetation along estuaries is exposed to strong environmental gradients that determine which species — enabled through specific traits — can establish and persist. With these ecosystems under anthropogenic pressure, in‐depth knowledge on the conservation of remaining tidal wetlands and restoration potentials is needed. In this study we elaborate the habitat conditions in the natural vegetation of the Elbe estuary and analyze: (a) which abiotic factors drive species composition; and (b) which species traits are key to the plants’ strategies in this specific ecotone.LocationThree sites in the Elbe estuary (river kilometer 671–703).MethodsWe collected data on soil nutrients, inundation period and wave height and sampled traits of the 17 most abundant plant species, which we analyzed by RLQ (three‐table analysis, including environment variables [R], species abundance [L] and a species traits table [Q]).ResultsWe detected a strong ˈwave–disturbance–inundation gradientˈ, which separated sparsely vegetated low‐lying plots receiving high wave impact and exposed to long inundation periods, from high‐lying plots showing dense vegetation and aerated soils. Close to the shore, plants showed low organ density and high investment into rhizomes, with a correlation of 0.72 between mass fraction of rhizome and phosphorus content of rhizome tissue. On higher elevations, plants strongly expressed traits relating to competition, like high allocation to stem biomass. Further, we found species with high leaf chlorophyll content showing low specific leaf area values and a negative correlation with the nutrient gradient.ConclusionThe results of our study are particularly relevant for restoration measures in order to re‐establish healthy marsh vegetation. However, changes in the environmental conditions, for instance stronger wave energies by a higher shipping frequency in the river channel and sea level rise may trigger changes in species composition through plant trait adaptations, for example by demanding a stem flexibility beyond what would be structurally feasible.

A stream by any other name: does β-diversity differ between Nearctic and Neotropical streams?

Beta diversity typically increases with decreasing latitude, with explanations of this pattern often based on tropical origins of most clades. Black flies (Diptera: Simuliidae) are at odds with this trend, having a northern origin. Increasing β-diversity with decreasing latitude might also be a product of changes in causal mechanisms structuring local assemblages among biogeographical realms. We tested the hypotheses that (i) β-diversity of black flies in streams should be similar between the Nearctic and Neotropical realms, and (ii) causal processes influencing species composition do not differ between these areas. To this end, we examined two Nearctic (USA) and two Neotropical data sets (Ecuador, Venezuela). Results did not show distinct differences in β-diversity between Nearctic and Neotropical streams. Although stream conditions varied among regions, we provide evidence that several causal mechanisms of community structure are shared. We suggest that potentially similar causal mechanisms driving species composition contributed to the lack of distinct differences in β-diversity between Nearctic and Neotropical streams. In addition, dispersal appears to influence the β-diversity of larval black flies, a pattern different from many previous studies of stream invertebrates. The influence of taxon origins on β-diversity across biogeographic realms should be further investigated.

Indirect facilitation drives species composition and stability in drylands

Dryland ecosystems are likely to respond discontinuously to gradual changes in environmental conditions. Direct facilitation between plants, whereby plants improve the local environmental conditions for others, has been shown to be a mechanism contributing to these discontinuous ecosystem transitions. Theoretical models describing dryland vegetation dynamics often consider a single plant species and one type of facilitation, namely direct facilitation. However, another type of facilitation–indirect facilitation–is widespread in dryland ecosystems as well; it is performed by plants protected against grazing, the nurses, when this protection extends to other plants growing in their neighbourhood that are deprived of such protection, the protegees. Little is known about the long-term effects of indirect facilitation on dryland dynamics. Here, we developed and analysed a theoretical model including two species–a nurse and a protegee–and indirect facilitation through grazing. We investigated the effects of indirect facilitation on species composition, species spatial clustering and the stability of dryland ecosystems. We showed that indirect facilitation through grazing enables the stable coexistence of the nurse and the protegee and allows the reversibility of the protegee extinction. Surprisingly, the strength of indirect facilitation affected neither the total nor the interspecific vegetation clustering. Our study highlights that spatially explicit grazing protection may affect species composition and the stability of dryland ecosystems and gives hints about how species interactions translate into spatial clustering.

Environmental variables driving species composition in subarctic springs in the face of climate change

AbstractQuestionsWhich environmental variables are most important in determining plant species composition in subarctic springs? Do observed patterns differ between typical wetland and general matrix‐derived species?LocationHelocrenic (seepage) springs, Northern Norway.MethodsWe sampled 49 helocrenic spring sites, measuring environmental variables (water temperature, water pH, electrical conductivity, discharge volume, geographic position) and recording all species present. We performed a partial canonical correspondence analysis (pCCA) to determine the relative importance of water quality, spatial, and climatic variables for patterns in species composition and to compare the differences in these patterns between wetland and matrix‐derived species.ResultsWe found that climatic and water quality variables were almost equally important in determining species composition in subarctic springs, with climatic variables explaining 26.62% of variation in species composition and water quality variables explaining 26.14%. Spatial variables explained the least variation (21.53%). When looking at the variables individually, altitude (10.93%) and mean summer temperature (9.25%) explained the most variation. The trend was the same for matrix‐derived species and wetland species, with climatic variables explaining the most variation (matrix‐derived: 27.26%; wetland: 24.42%), followed by water quality (matrix‐derived: 26.40%; wetland: 24.13%) and spatial variables (matrix‐derived: 24.87%; wetland: 16.27%). The main difference between matrix‐derived species and typical wetland species was that the spatial variables explained less variation for wetland species.ConclusionsThe close relationship of species composition (total vegetation as well as separated into wetland and matrix‐derived species) with climatic and water quality conditions indicates a sensitivity of subarctic springs to future climate change. In combination with altitude, which was found to be the most important individual variable, it is likely that the future distribution of spring species tracking climate change will be limited by the occurrence of suitable spring habitats, especially at high altitudes.

Defining the baseline for river restoration: comparing carabid beetle diversity of natural and human-impacted riparian habitats

Near-natural rivers and riparian ecosystems can represent biodiversity hotspots harbouring many highly specialised, rare and endangered species. During the past centuries, these habitats have been heavily degraded by anthropogenic use, and therefore river restoration is one of the most striking fields of action that is legally defined by the European Union Water Framework Directive. Successful restoration depends on realistic and specified targets that should be defined beforehand and founded on status quo surveys. We present a comparison of carabid beetle communities in riparian habitats of natural and managed river sites of the Mulde River in the Biosphere Reserve Middle Elbe. This endeavour is part of a unique multi-level revitalisation project. Pitfall trapping in 2016 and 2017 yielded 111 carabid species with many species of conservation concern in natural and managed habitats. However, Simpson diversity and functional diversity were lower in the latter. Both habitats harboured specific species assemblages with characteristic indicator species. Additionally, the trap location on slip-off slopes or cut banks was a significant driver of species composition. Our results indicate high ecological development potentials for the Mulde River, but restoration should consider differences between slip-off slopes and cut-off banks. We postulate that future restoration will foster population increases as well as a wider distribution of rare and endangered riparian habitat specialists.

Distinct taxonomic and phylogenetic patterns of plant communities on acid and limestone soils in subtropical and tropical China

AbstractQuestionUnderstanding the mechanisms determining species composition is one of the central goals in ecology. The limestone landscape of southwestern China is one of the most widely known landscapes developed on carbonate bedrock in the world. We explored the following question: What are the patterns of species composition and phylogenetic structures of the communities developed on the acid and limestone soils?LocationSouthwestern China.MethodsUsing 536 plots on acid and limestone soils, we identified distinct vegetation types on both soil types based on hierarchical clustering and ordination. Variation partitioning partitioned the effects of climate, geographic distance and soil on species composition. Generalized linear models determined the effects of climate and soil on phylogenetic diversities.ResultsThe typical vegetation on acid soils is evergreen broad‐leaved forests characterized by Castanopsis species, while typical vegetation on limestone soils is mixed evergreen and deciduous forests characterized by a high proportion of deciduous trees and calciphytes. Soil type was much more important in driving species composition than climate and spatial distance. The standardized mean pairwise phylogenetic distance (ses.MPD) and mean nearest taxon distance (ses.MNTD) were significantly lower on limestone soils than on acid soils, indicating a more clustered phylogenetic structure. The effect of soil on ses.MPD was stronger than on ses.MNTD, and ses.MNTD increased with increasing annual temperature and precipitation seasonality on acid soils but decreased on limestone soils, suggesting that soil affected phylogenetic structure at relatively deep branches, while climate combined with soil mainly affects the relatively recent branches, as ses.MPD measures phylogenetic structure at deep branches and ses.MNTD that at shallow branches.ConclusionsThis study highlights the importance of soil in shaping community structure at a regional scale, and it also reveals the unique features of limestone vegetation and supports the validity of limestone vegetation types in the Chinese vegetation classification system.

A Constraint-based model of Dynamic Island Biogeography: environmental history and species traits predict hysteresis in populations and communities

A Constraint-based model of Dynamic Island Biogeography: environmental history and species traits predict hysteresis in populations and communities We present a conceptual model that shows how hysteresis can emerge in dynamic island systems given simple constraints on trait-mediated processes. Over time, many islands cycle between phases of increasing and decreasing size and connectivity to a mainland species pool. As these phases alternate, the dominant process driving species composition switches between colonization and extinction. Both processes are mediated by interactions between organismal traits and environmental constraints: colonization probability is affected by a species’ ability to cross the intervening matrix between a population source and the island; population persistence (or extinction) is driven by the minimum spatial requirements for sustaining an isolated population. Because different suites of traits often mediate these two processes, similar environmental conditions can lead to differences in species compositions at two points of time. Thus, the Constraint-based model of Dynamic Island Biogeography (C-DIB) illustrates the possible role of hysteresis—the dependency of outcomes not only on the current system state but also the system’s history of environmental change—in affecting populations and communities in insular systems. The model provides a framework upon which additional considerations of lag times, biotic interactions, evolution, and other processes can be incorporated. Importantly, it provides a testable framework to study the physical and biological constraints on populations and communities across diverse taxa, scales, and systems.

Environment or management? Relative importance for floristic composition of sub‐Mediterranean hay meadows in Central Italy

AbstractQuestionsWhat are the separate and combined effects of natural and management factors on floristic assemblage in sub‐Mediterranean hay meadows? How are hay meadow dynamics affected by the environment excluding management?LocationCentral Apennines, Italy.MethodsVegetation was studied in 66 hay meadows by random plots, recording four groups of explanatory variables: management, soil, topography, and space. Using partial ordinations, the explanatory variables of the variation in floristic composition and plant traits were divided into sets. Then, to emphasize the relationship between purely environmental variables and vegetation, we considered only the partial ordination executed with soil and topographic variables using management and spatial data as covariables.ResultsThe soil variables had the highest marginal explanatory power. Organic matter was the main driver of floristic composition, indicating a gradient associated also with soil moisture. The pure effect of spatial variables was very important, reflecting historical processes. On the contrary, management and topography variables sets were relatively weak predictors. Partial ordination showed that two topographic variables, slope and height above the plain, had an important weight in the hay meadow floristic assemblage, influencing soil fertility.ConclusionsThis analysis (a) confirms the determining function of soil fertility and moisture in driving plant species composition and traits; (b) focuses on the marginal power of current management when past traditional agro‐pastoral activity undergoes changes in mode and intensity; (c) highlights the role of spatial variables and site history; and (d) improves knowledge about topographical factors as drivers of species composition. The maintenance of hay meadow mosaics requires preservation of grasslands in complete topographical catenae and prevention of the abandonment of parcels on steep slopes. Considering the spatial variability of the studied communities, it is necessary to include a high number of geographically and ecologically different areas in a conservation network.

Discriminating among forest communities based on taxonomic, phylogenetic and trait distances

Quantifying the compositional differences among communities is central to answering some of the most challenging questions in community ecology. Traditional species-based estimates of community dissimilarity convey little information regarding the biological heterogeneity of species. More refined phylogenetic- and functional-based measures can improve the understanding of ecological mechanisms that drive species composition. However, a generalized framework, which unifies taxonomic, phylogenetic, and functional information of communities is still lackling. We present a new general framework for assessing the biological dissimilarity among communities based on species frequencies and biological (including taxonomic, phylogenetic, and functional) distances between species. We used the observations collected in a 30-hectare forest plot in northeastern China to illustrate the application of the new approach and its ability to discriminate communities along spatial and environmental gradients. The results suggested that both spatial and environmental gradients play significant roles in driving the species composition of forest communities. Compared with spatial gradients, local environmental conditions had a greater influence. Conclusion: The ability to measure differences among communities, based on species frequency and biological distances is useful for estimating effects of habitat heterogeneity, for understanding the mechanism of community assembly, and for assessing disturbance effects or species invasions at local or global scales. The Avalanche approach presented in this study represents an effective framework for comparing different measures of biological dissimilarity in one compatible system.

Priority Effects and Nonhierarchical Competition Shape Species Composition in a Complex Grassland Community.

Niche and fitness differences control the outcome of competition, but determining their relative importance in invaded communities—which may be far from equilibrium—remains a pressing concern. Moreover, it is unclear whether classic approaches for studying competition, which were developed predominantly for pairs of interacting species, will fully capture dynamics in complex species assemblages. We parameterized a population-dynamic model using competition experiments of two native and three exotic species from a grassland community. We found evidence for minimal fitness differences or niche differences between the native species, leading to slow replacement dynamics and priority effects, but large fitness advantages allowed exotics to unconditionally invade natives. Priority effects driven by strong interspecific competition between exotic species drove single-species dominance by one of two exotic species in 80% of model outcomes, while a complex mixture of nonhierarchical competition and coexistence between native and exotic species occurred in the remaining 20%. Fungal infection, a commonly hypothesized coexistence mechanism, had weak fitness effects and is unlikely to substantially affect coexistence. In contrast to previous work on pairwise outcomes in largely native-dominated communities, our work supports a role for nearly neutral dynamics and priority effects as drivers of species composition in invaded communities.

Environmental drivers of plant assemblages: are there differences between palustrine and lacustrine wetlands? A case study from the northern Apennines (Italy)

Mountain wetlands are among the most vulnerable habitats in the Mediterranean basin. Their conservation requires knowledge of plant species assemblages and their environmental drivers. In this study, we investigated what the main environmental factors driving species composition in mountain wetlands are. Differences in environmental control and floristic composition between palustrine and lacustrine wetlands were explored. We used a dataset of 168 vegetation plots (relevés), sampled at 45 mountain wetlands in the northern Apennines (central Italy). Direct ordination showed that water depth, geology type and altitude were the main factors responsible for species distribution. The most important gradient was linked to soil moisture, with hygrophilous species increasing with moisture levels. Indicator Species Analysis underlined a clear distinction in the distribution of aquatic plants between wetland subsystems. Geology and rainfall affected species assemblages in lacustrine and palustrine subsystems. Indirect ordination and Generalized Additive Models revealed that plant species and their attributes significantly changed in the wetland subsystems with an increase in hydrophytes with increasing rainfall in palustrine wetlands and a decrease in thermophilous species along an altitudinal gradient in lacustrine wetlands. Management and conservation guidelines for northern Apennines wetlands are suggested.

Teamwork makes the dream work: Disentangling cross-taxon congruence across soil biota in black pine plantations

Soil plays a fundamental role in many ecological processes, throughout a complex network of above- and below-ground interactions. This has aroused increasing interest in the use of correlates for biodiversity assessment and has demonstrated their reliability with respect to proxies based on environmental data alone. Although co-variation of species richness and composition in forests has been discussed in the literature, only a few studies have explored these elements in forest plantations, which are generally thought to be poor in biodiversity, being aimed at timber production. Based on this premise our aims were 1) to test if cross-taxon congruence across different groups of organisms (bacteria, vascular plants, mushrooms, ectomycorrhizae, mycelium, carabids, microarthropods, nematodes) is consistent in artificial stands; 2) to evaluate the strength of relationships due to the existing environmental gradients as expressed by abiotic and biotic factors (soil, spatial-topographic, dendrometric variables). Correlations between groups were studied with Mantel and partial Mantel tests, while variance partition analysis was applied to assess the relative effect of environmental variables on the robustness of observed relationships. Significant cross-taxon congruence was observed across almost all taxonomic groups pairs. However, only bacteria/mycelium and mushrooms/mycelium correlations remained significant after removing the environmental effect, suggesting that a strong abiotic influence drives species composition. Considering variation partitioning, the results highlighted the importance of bacteria as a potential indicator: bacteria were the taxonomic group with the highest compositional variance explained by the predictors used; furthermore, they proved to be involved in the only cases where the variance attributed solely to the pure effect of biotic or abiotic predictors was significant. Remarkably, the co-dependent effect of all predictors always explained the highest portion of total variation in all dependent taxa, testifying the intricate and dynamic interplay of environmental factors and biotic interactions in explaining cross-taxon congruence in forest plantations.

Dispersal increases ecological selection by increasing effective community size.

Selection and drift are universally accepted as the cornerstones of evolutionary changes. Recent theories extend this view to ecological changes, arguing that any change in species composition is driven by deterministic fitness differences among species (enhancing selection) and/or stochasticity in birth and death rates of individuals within species (enhancing drift). These forces have contrasting effects on the predictability of ecological dynamics, and thus understanding the factors affecting their relative importance is crucial for understanding ecological dynamics. Here we test the hypothesis that dispersal increases the relative importance of ecological selection by increasing the effective size of the community (i.e., the size relevant for competitive interactions and drift). According to our hypothesis, dispersal increases the effective size of the community by mixing individuals from different localities. This effect diminishes the relative importance of demographic stochasticity, thereby reducing drift and increasing the relative importance of selective forces as drivers of species composition. We tested our hypothesis, which we term the "effective community size" hypothesis, using two independent experiments focusing on annual plants: a field experiment in which we manipulated the magnitude of dispersal and a mesocosm experiment in which we directly manipulated the effective size of the communities. Both experiments, as well as related model simulations, were consistent with the hypothesis that increasing dispersal increases the role of selective forces as drivers of species composition. This finding has important implications for our understanding of the fundamental forces affecting community dynamics, as well as the management of species diversity, particularly in patchy and fragmented environments.

What are the principal environmental filters driving species composition and succession on mineralogically different spoil heaps?

ABSTRACTThe relationship between selected environmental variables and plant species composition was studied on two mineralogically different spoil heaps (Hg and Cu) in Central Slovakia with contrasting reclamation approaches. Data on plant species composition were collected by stratified random sampling in defined physiognomic vegetation types. A detrended correspondence analysis showed that most of the variability in species composition was related to the succession gradient from open communities with a low cover of vascular plants to forest vegetation, and to the moisture gradient. Variance partitioning by canonical correspondence analysis revealed that most of the variability in plant composition was related to the content of various heavy metals (27.8% at the Hg-spoil heap and 28.3% at the Cu-spoil heap), but a significant relationship was found only for Mn. Other significant factors comprised soil moisture, pH and P content for the Hg-spoil heap and soil temperature and Ca content for the Cu-spoil heap. Although heavy metal content explained most of the variability in species composition, the relationship was caused by the correlation of heavy metal content with other environmental variables rather than by a direct causal relationship.

Forest snail diversity and its environmental predictors along a sharp climatic gradient in southern Siberia

Diversity patterns of forest snail assemblages have been studied mainly in Europe. Siberian snail faunas have different evolutionary history and colonization dynamics than European faunas, but studies of forest snail diversity are almost missing from Siberia. Therefore, we collected snails at 173 forest sites in the Russian Altai and adjacent areas, encompassing broad variation in climate and forest types. We found 51 species, with a maximum of 15 and an average of seven species per site. The main gradient in species composition was related to soil pH, a variable that also positively correlates with snail abundances. The second gradient was associated with climate characteristics of winter. We observed significant differences in both species richness and composition among six forest types defined based on vegetation classification. Hemiboreal continental forests were the poorest of these types but hosted several species characteristic of European full-glacial stages of the Late Pleistocene. A high snow cover in Temperate coniferous and mixed forests, protecting the soil from freezing, allowed the frost-sensitive large-bodied (>10 mm) species to inhabit this forest type. In contrast to most of the European snail assemblages studied so far we found that the factors responsible for the variation in species richness differed from those driving species composition. This may be attributed to the sharp climatic gradient and the presence of the cold-adapted species typical of the Pleistocene cold stages. We suggest that southern Siberian forests hosting these species can serve as modern analogues of full-glacial forests in periglacial Central and Eastern Europe.