Climatically promoted taxonomic homogenization of macroinvertebrates in unaffected streams varies along the river continuum.

Biotic homogenization appears to be a global consequence of anthropogenic change. However, the underlying environmental factors contributing to homogenization are difficult to identify because their effects usually interact and confound each other. This can be the reason why there is very little evidence on the role of climate warming in homogenization. By analysing macroinvertebrate assemblages in 65 streams that were as close to natural conditions as possible, we avoided the confounding effects of common anthropogenic stressors. This approach resulted in revealing a significant effect of increased temperature (both summer and winter) on changes in macroinvertebrate compositional over the past two decades. However, homogenization was significant only at opposite ends of the river continuum (submontane brooks, low-altitude rivers). Surprisingly, species of native origin predominated overall, increasing in frequency and abundance ("winners"), while only a minority of species declined or disappeared ("losers"). We hypothesise that undisturbed conditions mitigate species declines and thus homogenization, and that the temperature increase has so far been beneficial to most native species. Although we may have only captured a transitional state due to extinction debt, this underscores the importance of maintaining ecological conditions in stream to prevent species loss due to climate change.

Conservation and restoration of Central European fens by mowing: A consensus from 20 years of experimental work.

While the importance of conservation mowing for mesic grasslands is generally accepted, its use for fens and fen grasslands interspersed within agricultural land is still controversial. Although fens may persist naturally, ongoing environmental changes increase productivity and accelerate succession. These processes can be mitigated through conservation management with appropriate settings. However, long-term management experiments are challenging and provide only locally valid results. Here, we analysed vegetation data (bryophytes and vascular plants) from seven management experiments (spanning 3-20 years) conducted in Central European poor, moderately-rich, and calcareous spring fens (Czech Republic, Slovakia). Two of these experiments examined the effects of restoration of abandoned fens, while five experiments examined changes in mowing regimes in managed fens (cessation, intensification, delay to autumn, and litter removal). Data were analysed using unidimensional and multidimensional methods separately for the initial, extended, and entire period. Mowing had a statistically significant effect on species composition except for the shortest (3-year) experiment. Litter removal did not compensate for mowing. Mowing twice or delayed mowing significantly affected the species composition of calcareous fens. In all cases, cessation of mowing significantly reduced the richness of species, especially those of conservation importance. In contrast, any mowing of abandoned fens increased species richness. The effects of mowing intensification or cessation on species richness and composition of a restored calcareous fen were evident in the first 2-3 years. Other effects were initially weak or nonsignificant but later became stronger, such as mowing delay and restoration removal of litter, which became significant only after nearly 20 years. We found that cessation or restoration of mowing usually triggers a rapid vegetation change, whereas it can take decades to detect the response caused by changes in mowing timing. Importantly, mowing can stabilise or even restore vegetation of fen ecosystems that have been weakened by their fragmentation in the temperate agricultural landscapes.

Ecological niche divergence between extant and glacial land snail populations explained.

The presence of Last Glacial Maximum (LGM) biotic communities without modern counterparts is well known. It is particularly evident in central European fossil LGM land snails whose assemblages represent an odd mix of species that are currently limited to either xeric or wetland habitats. Here we document a genetically verified discovery of the modern calcareous wetland species Pupilla alpicola on Iceland, where it is limited to dry grasslands. This species also represents a common European LGM fossil, and its new records from Iceland help explain puzzling shifts of some glacial land snails of xeric grassland habitats to open wetlands today. Similarities between the climates of modern Iceland and LGM Eurasia suggest that this species did not become limited to wetlands in continental Europe until after the Late Pleistocene-Holocene climate transition. These results are a strong reminder that assumptions of ecological uniformity must be questioned and that the quality and robustness of palaeoecological reconstructions is dependent upon adequate knowledge of the full autecological range of species over time.

Land snail community patterns related to regional habitat conservation status of European spring fens.

Human activities have enormous impact on current biodiversity distribution across all spatial scales. Despite the numerous studies showing the difference between preserved and impaired sites, only little is known about the regional scale. Therefore, we selected four European regions differing in habitat conservation status (HCS) to explore if the variation in land snail communities reflects regional differences. We collected quantitative land snail samples at 169 isolated spring fen sites and measured environmental parameters. The species richness of habitat specialists expressed low variation and weak associations with local conditions in the two regions of adequate HCS, presumably because of their common occurrence throughout most sites. In contrast, the richness of matrix-derived species, i.e. predominantly habitat generalists, was highly variable in these two regions and also tightly associated with local conditions, especially moisture. In both the intermediate and the inadequate HCS region, these associations were much weaker as the fens are less extreme and allow for penetration of matrix-derived species. Population densities of Vertigo geyeri, an umbrella species internationally protected by the EU Habitats Directive, were highest in the two adequate HCS regions. Species composition was primarily controlled by moisture in the regions of adequate HCS, while in the remaining regions, those predictors that are less easily jeopardized by human impact, such as climate, water chemistry and terrain topography, gained importance. In the inadequate HCS region, none of the analysed predictors was associated with the main compositional gradient, suggesting a complete disruption of community-environment relationships. Our results suggest that the species richness and community responses to natural gradients might be substantially modified by human impact, although the effect of some other region-specific factors cannot be easily disentangled because of inevitably low number of studied regions.

Habitat extremity and conservation management stabilise endangered calcareous fens in a changing world.

Calcareous fens represent an endangered type of peatlands, acting as refugia for stress-tolerant species in the currently changing landscapes. The resurveys across many regions have reported their recent disappearance or deterioration despite both the extreme habitat conditions (carbonate richness, presence of calcareous tufa, nutrient limitation, high water level) and conservation management. To test the stability of their biotic communities in different environmental and management configurations, we repeatedly sampled molluscs (terrestrial and aquatic), vascular plants, and bryophytes at 30 calcareous fens in the Inner Western Carpathians (Slovakia, Poland) after 13-17 years of warm summers and land-use changes. We found a small yet statistically significant effect of sampling period (old versus new survey) on the species composition of all three groups of organisms when the effect of various positions of sites along ecological gradients was controlled for. The compositional changes, interpreted with the help of Ellenberg Indicator Values, suggest an incipient succession towards grasslands and shrublands, driven by decreasing soil moisture and increasing nutrient availability. Although the number of habitat specialists did not change, the number of matrix-derived vascular plant and bryophyte species significantly increased, with six ubiquitous species of productive habitats being significantly more represented currently, while the richness of aquatic molluscs significantly decreased. Fens in which potentially strongly competitive plant species were less stressed because of less intense management and lower habitat extremity were more prone to such succession. There was no single factor that could predict the magnitude of composition changes; instead, tested factors were found to act synergistically. Conservation management was predominantly important for bryophytes, while extreme habitat conditions were predominantly important for terrestrial snails. We suggested a way how nature conservancy authorities can prioritise the management needs by applying an abiotic indicator system, with less environmentally extreme fens requiring more intense conservation management.

Small-scale Variation of Testate Amoeba Assemblages: the Effect of Site Heterogeneity and Empty Shell Inclusion.

Studies on testate amoeba species distribution at small scales (i.e., single peatland sites) are rare and mostly focus on bogs or mineral-poor Sphagnum fens, leaving spatial patterns within mineral-rich fens completely unexplored. In this study, two mineral-rich fen sites of contrasting groundwater chemistry and moss layer composition were selected for the analysis of testate amoeba compositional variance within a single site. At each study site, samples from 20 randomly chosen moss-dominated plots were collected with several environmental variables being measured at each sampling spot. We also distinguished between empty shells and living individuals to evaluate the effect of empty shell inclusion on recorded species distribution. At the heterogeneous-rich Sphagnum-fen, a clear composition turnover in testate amoebae between Sphagnum-dominated and brown moss-dominated samples was closely related to water pH, temperature and redox potential. We also found notable species composition variance within the homogeneous calcareous fen, yet it was not as high as for the former site and the likely drivers of community assembly remained unidentified. The exclusion of empty shells provided more accurate data on species distribution as well as their relationship with some environmental variables, particularly moisture. Small-scale variability in species composition of communities seems to be a worthwhile aspect in testate amoeba research and should be considered in future sampling strategies along with a possible empty shell bias for more precise understanding of testate amoeba ecology and paleoecology.

When is a "cryptic" species not a cryptic species: A consideration from the Holarctic micro-landsnail genus Euconulus (Gastropoda: Stylommatophora).

Naive use of molecular data may lead to ambiguous conclusions, especially within the context of "cryptic" species. Here, we integrated molecular and morphometric data to evaluate phylogenetic relationships in the widespread terrestrial micro-snail genus, Euconulus. We analyzed mitochondrial (16S + COII) and nuclear (ITS1 + ITS2) sequence across 94 populations from Europe, Asia and North America within the nominate species E. alderi, E. fulvus and E. polygyratus, and used the southeastern USA E. chersinus, E. dentatus, and E. trochulus as comparative outgroups. Phylogeny was reconstructed using four different reconstruction methods to identify robust, well-supported topological features. We then performed discriminant analysis on shell measurements between these genetically-identified species-level clades. These analyses provided evidence for a biologically valid North American "cryptic" species within E. alderi. However, while highly supported polyphyletic structure was also observed within E. fulvus, disagreement in placement of individuals between mtDNA and nDNA clades, lack of morphological differences, and presence of potential hybrids imply that these lineages do not rise to the threshold as biologically valid cryptic species, and rather appear to simply represent a complex of geographically structured populations within a single species. These results caution that entering into a cryptic species hypothesis should not be undertaken lightly, and should be optimally supported along multiple lines of evidence. Generally, post-hoc analyses of macro-scale features should be conducted to attempt identification of previously ignored diagnostic traits. If such traits cannot be found, i.e. in the case of potentially "fully cryptic" species, additional criteria should be met to propound a cryptic species hypothesis, including the agreement in tree topology among both mtDNA and nDNA, and little (or no) evidence of hybridization based on a critical analysis of sequence chromatograms. Even when the above conditions are satisfied, it only implies that the cryptic species hypothesis is plausible, but should optimally be subjected to further careful examination.

Spring-fen habitat islands in a warming climate: Partitioning the effects of mesoclimate air and water temperature on aquatic and terrestrial biota.

Climate warming and associated environmental changes lead to compositional shifts and local extinctions in various ecosystems. Species closely associated with rare island-like habitats such as groundwater-dependent spring fens can be severely threatened by these changes due to a limited possibility to disperse. It is, however, largely unknown to what extent mesoclimate affects species composition in spring fens, where microclimate is buffered by groundwater supply. We assembled an original landscape-scale dataset on species composition of the most waterlogged parts of isolated temperate spring fens in the Western Carpathian Mountains along with continuously measured water temperature and hydrological, hydrochemical, and climatic conditions. We explored a set of hypotheses about the effects of mesoclimate air and local spring-water temperature on compositional variation of aquatic (macroinvertebrates), semi-terrestrial (plants) and terrestrial (land snails) components of spring-fen biota, categorized as habitat specialists and other species (i.e. matrix-derived). Water temperature did not show a high level of correlation with mesoclimate. For all components, fractions of compositional variation constrained to temperature were statistically significant and higher for habitat specialists than for other species. The importance of air temperature at the expense of water temperature and its fluctuation clearly increased with terrestriality, i.e. from aquatic macroinvertebrates via vegetation (bryophytes and vascular plants) to land snails, with January air temperature being the most important factor for land snails and plant specialists. Some calcareous-fen specialists with a clear distribution centre in temperate Europe showed a strong affinity to climatically cold sites in our study area and may hence be considered as threatened by climate warming. We conclude that prediction models solely based on air temperature may provide biased estimates of future changes in spring fen communities, because their aquatic and semiterrestrial components are largely affected by water temperature that is modified by local hydrological and landscape settings.

Contrasting diversity of testate amoebae communities in Sphagnum and brown-moss dominated patches in relation to shell counts.

Ecological studies of peatland testate amoebae are generally based on totals of 150 individuals per sample. However, the suitability of this standard has never been assessed for alkaline habitats such as spring fens. We explored the differences in testate amoeba diversity between Sphagnum and brown-moss microhabitats at a mire site with a highly diversified moss layer which reflects the small-scale heterogeneity in groundwater chemistry. Relationships between sampling efficiency and sample completeness were explored using individual-based species accumulation curves and the effort required to gain an extra species was assessed. Testate amoeba diversity differed substantially between microhabitats, with brown mosses hosting on average twice as many species and requiring greater shell totals to reach comparable sample analysis efficiency as for Sphagnum. Thus, for samples from alkaline conditions an increase in shell totals would be required and even an overall doubling up to 300 individuals might be considered for reliable community description. Our small-scale data are likely not robust enough to provide an ultimate solution for the optimization of shell totals. However, the results proved that testate amoebae communities from acidic and alkaline environments differ sharply in both species richness and composition and they might call for different methodological approaches.

Unimodal latitudinal pattern of land-snail species richness across northern Eurasian lowlands.

Large-scale patterns of species richness and their causes are still poorly understood for most terrestrial invertebrates, although invertebrates can add important insights into the mechanisms that generate regional and global biodiversity patterns. Here we explore the general plausibility of the climate-based "water-energy dynamics" hypothesis using the latitudinal pattern of land-snail species richness across extensive topographically homogeneous lowlands of northern Eurasia. We established a 1480-km long latitudinal transect across the Western Siberian Plain (Russia) from the Russia-Kazakhstan border (54.5°N) to the Arctic Ocean (67.5°N), crossing eight latitudinal vegetation zones: steppe, forest-steppe, subtaiga, southern, middle and northern taiga, forest-tundra, and tundra. We sampled snails in forests and open habitats each half-degree of latitude and used generalized linear models to relate snail species richness to climatic variables and soil calcium content measured in situ. Contrary to the classical prediction of latitudinal biodiversity decrease, we found a striking unimodal pattern of snail species richness peaking in the subtaiga and southern-taiga zones between 57 and 59°N. The main south-to-north interchange of the two principal diversity constraints, i.e. drought stress vs. cold stress, explained most of the variance in the latitudinal diversity pattern. Water balance, calculated as annual precipitation minus potential evapotranspiration, was a single variable that could explain 81.7% of the variance in species richness. Our data suggest that the "water-energy dynamics" hypothesis can apply not only at the global scale but also at subcontinental scales of higher latitudes, as water availability was found to be the primary limiting factor also in this extratropical region with summer-warm and dry climate. A narrow zone with a sharp south-to-north switch in the two main diversity constraints seems to constitute the dominant and general pattern of terrestrial diversity across a large part of northern Eurasia, resulting in a subcontinental diversity hotspot of various taxa in this zone.

Diversity and biotic homogenization of urban land-snail faunas in relation to habitat types and macroclimate in 32 central European cities.

The effects of non-native species invasions on community diversity and biotic homogenization have been described for various taxa in urban environments, but not for land snails. Here we relate the diversity of native and non-native land-snail urban faunas to urban habitat types and macroclimate, and analyse homogenization effects of non-native species across cities and within the main urban habitat types. Land-snail species were recorded in seven 1-ha plots in 32 cities of ten countries of Central Europe and Benelux (224 plots in total). Each plot represented one urban habitat type characterized by different management and a specific disturbance regime. For each plot, we obtained January, July and mean annual temperature and annual precipitation. Snail species were classified into either native or non-native. The effects of habitat type and macroclimate on the number of native and non-native species were analysed using generalized estimating equations; the homogenization effect of non-native species based on the Jaccard similarity index and homogenization index. We recorded 67 native and 20 non-native species. Besides being more numerous, native species also had much higher beta diversity than non-natives. There were significant differences between the studied habitat types in the numbers of native and non-native species, both of which decreased from less to heavily urbanized habitats. Macroclimate was more important for the number of non-native than native species; however in both cases the effect of climate on diversity was overridden by the effect of urban habitat type. This is the first study on urban land snails documenting that non-native land-snail species significantly contribute to homogenization among whole cities, but both the homogenization and diversification effects occur when individual habitat types are compared among cities. This indicates that the spread of non-native snail species may cause biotic homogenization, but it depends on scale and habitat type.

The age of island-like habitats impacts habitat specialist species richness.

While the effects of contemporaneous local environment on species richness have been repeatedly documented, much less is known about historical effects, especially over large temporal scales. Using fen sites in the Western Carpathian Mountains with known radiocarbon-dated ages spanning Late Glacial to modern times (16 975-270 cal years before 2008), we have compiled richness data from the same plots for three groups of taxa with contrasting dispersal modes: (1) vascular plants, which have macroscopic propagules possessing variable, but rather low, dispersal abilities; (2) bryophytes, which have microscopic propagules that are readily transported long distances by air; and (3) terrestrial and freshwater mollusks, which have macroscopic individuals with slow active migration rates, but which also often possess high passive dispersal abilities. Using path analysis we tested the relationships between species richness and habitat age, area, isolation, and altitude for these groups. When only matrix-derived taxa were considered, no significant positive relation was noted between species richness and habitat size or age. When only calcareous-fen specialists were considered, however, habitat age was found to significantly affect vascular plant richness and, marginally, also bryophyte richness, whereas mollusk richness was significantly affected by habitat area. These results suggest that in inland insular systems only habitat specialist (i.e., interpatch disperser and/or relict species) richness is influenced by habitat age and/or area, with habitat age becoming more important as species dispersal ability decreases.

Mollusc communities in Bulgarian fens: predictive power of the environment, vegetation, and spatial structure in an isolated habitat.

Mollusc communities of previously unexplored Bulgarian fens were studied in order to determine and generalise the patterns of species richness and composition along the mineral richness gradient. The aim was also to compare predictive values of the environment, vegetation and spatial structure. Altogether, 44 mollusc species were recorded at 40 treeless fen sites. Species richness varied from 0 to 18 species per site, and it was positively associated with the mineral gradient and negatively with altitude. However, the best predictor was obtained using plant species composition. All explanatory variables had higher effect on land snails than on the entire mollusc assemblage (including aquatic species). Species richness and abundance were significantly and positively correlated with the species composition turnover; the communities were highly nested, with poor sites having subsets of the fauna found in the richest. The main direction of mollusc species turnover was highly associated with that observed for vegetation, and the main gradient of plant species composition was able to explain nearly 20% of total variation in mollusc data. We found that spatial structure explained by far the highest proportion of independent variation, which reflected the high level of geographical isolation of Bulgarian fens and regional differences independent of any environmental variation. Our results demonstrate (1) the general role of mineral richness gradient for structuring mollusc communities in fens, (2) the pivotal indicator role of plant species composition in predicting species composition of mollusc communities, despite being trophically independent and (3) the effect of isolation and origins of the habitat on species composition: most species have wide geographical distributions within the habitat type, and geographical patterns within Bulgaria may have a stochastic element.

Deterministic assembly of land snail communities according to species size and diet.

1. We investigated whether coexisting snail species in 145 treeless fen communities in the Western Carpathian Mountains differed more in size and diet than would be expected by chance, as predicted for traits commonly associated with competition and differential resource acquisition under limiting similarity theory. 2. Contrary to expectations, coexisting snail species were no more different in body size than expected by chance under a null model. However, variation in body size played a significant role in structuring snail communities: coexisting snail species were significantly more similar with respect to body size. 3. We developed two new test statistics to expand our investigation of limiting similarity to include diet, a nominal trait. We tested whether communities of snails were characterized by a greater richness of diet, and whether different diets were represented more or less evenly within communities. Communities of snails were significantly less evenly distributed than expected by chance, with detritivores being over-represented relative to predatory strategies. 4. We also examined the effect of water pH and conductivity, herbaceous cover, and bryophyte and vascular plant richness, on these trends by examining how the effect size of our tests varied across these gradients. Convergence in species size increased with increasing habitat pH. Specifically, smaller snail species were over-represented in fen communities in general, and this effect was accentuated in increasingly calcareous fens. 5. Theory predicts that traits related strongly to environmental conditions are more likely to be convergent. Our findings support this suggestion, as small snail species have an advantage in tolerating freezing conditions over winter when refuges are limited. 6. These results add to the growing body of literature demonstrating that variation in body size and diet play a strong role in structuring communities, although frequently in ways not predicted by limiting similarity theory. Finally, our results increase our understanding of how species are assembled non-randomly into communities with respect to important traits.

Nested patterns in hyporheic meta-communities: the role of body morphology and penetrability of sediment.

Nestedness has been regarded as a common pattern of species distribution especially in terrestrial systems and vertebrate faunas. However, a significantly lower degree of nestedness has been reported for aquatic invertebrates. We analyzed the vertical distribution patterns of taxa in the upper 70 cm of the hyporheic zone. This biotope is abundantly inhabited by epigean fauna, which is morphologically pre-adapted to life within the limited space of sediment interstices. We tested the hypotheses that in the vertical profile of the hyporheic zone sediment acts as a physical barrier (filter), allowing only the morphologically pre-adapted and adapted (i.e., smaller, narrower, more flexible) taxa to penetrate to deeper layers. We demonstrated that this mechanism can promote a strongly nested and colonization-driven pattern at higher taxa levels. The sediment filter (1) constricted the body width: 0.50 mm appeared to be the upper limit for successful sediment colonization at the study site, and (2) favored elongated taxa against small sized taxa. We tested also the assumption that distribution of fine sediment affects the accessibility of hyporheic zone for fauna ("filter density") and thereby influences nestedness. However, we found that nestedness could be sufficiently explained by the depth itself. Our study offers a possible explanation of depth patterns in hyporeic meta-communities as a result of morphological characteristics promoting nestedness at higher taxa level.