Summary The relevance of environmental and spatial processes for species distributions varies among environments and types of metacommunities. Here, for the first time, we use modern statistical approaches to test the contribution of these two processes in structuring ecologically unique and threatened biotas of insular spring fens. We applied two species categorisations, common/rare and generalists/specialists, to disentangle the roles of dispersal capacity and habitat specialisation. In accordance with current understanding of headwater ecosystems, we found that environmental processes played a major role in most of the spring fen taxonomic and functional groups. However, we observed significant spatial structure in passive dispersers (Clitellata, a class of annelid worms), common species and habitat specialists. Spatial processes played the leading role in structuring the metacommunity of passively dispersing specialists. In contrast, all analysed insect groups, even those known to be poor dispersers, were able to reach virtually all favourable sites. We conclude that dispersal mode (active versus passive) and, to a lesser extent, habitat specialisation are the main factors determining the mechanism of spring fen metacommunity structuring.

Aims The aim was to identify the main drivers of aquatic macroinvertebrate species richness in spring‐fen habitats (i.e. groundwater seepage wetlands) because these habitats are among the most threatened temperate biodiversity hotspots. Location Isolated spring fens in the western Carpathian Mountains. Methods Assemblages of Tricladida, Clitellata, Mollusca, Ephemeroptera, Plecoptera, Trichoptera and Diptera were sampled at 48 fens distributed evenly along a gradient of water mineral richness and grouped according to habitat specialization and dispersal mode. Several physicochemical parameters that have been reported to be the main drivers of species distribution were recorded for each site, along with radiocarbon measurements of the absolute age of the sites. The numbers of species in the taxonomic groups analysed were modelled as a function of the predictors via multiple linear regressions. Results We achieved a notably higher sampling efficiency than had been used in previous spring aquatic macroinvertebrate studies. In total, we collected and identified 255 species within 331 taxa from > 235,000 individuals. The number of species in all taxonomic groups increased with water discharge but the highest predictive power was obtained with water redox potential, which explained as much as 55.7% of the specialist species richness variance. We found contrasting and systematic differences in the importance of predictors related mainly to the level of species habitat specialization. Species richness of spring‐fen specialists was strongly determined by the main environmental gradient of change in groundwater chemistry, while generalists primarily reflected habitat stability linked to higher water discharge, habitat size and absolute age. Main conclusion Isolated island‐like spring fens can harbour unusually species‐rich assemblages of aquatic macroinvertebrates, the species richness of which is shaped by contrasting mechanisms dependent mainly on habitat specialization and also partly on dispersal mode. The richness of habitat specialist species seen at calcareous fens indicates their conservation priority.

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Groundwater chemistry is a major determinant of assemblages of various taxonomic groups in spring fens, but its effect on insect assemblages has not been proved yet. We investigated dipteran assemblages at 17 isolated spring fens, which encompass the whole mineral richness gradient from rich (calcareous) to poor (highly acidic) sites, and analyzed faunal patterns at two contrasting mesohabitats: flowing water and standing water. The effect of water chemistry, substratum features, discharge, and temperature on the dipteran assemblages were assessed using PERMANOVA and GAM. Highly diverse dipteran assemblages (156 taxa) were closely related to the mineral richness gradient at both mesohabitats, showing a continual and nearly complete species exchange along the gradient, while their total abundance and taxa density did not change significantly. The assemblages included both habitat generalists and taxa specifically associated with acidic, moderate, or calcareous conditions. The mineral richness gradient was also reflected by changes in substratum properties, thus creating a complex environmental gradient that we suggest is the main environmental gradient structuring aquatic assemblages in spring fens.

We examined responses of macroinvertebrate assemblages to environmental and temporal variations along spring source-spring brook transects in two fen habitats, sharply differing in groundwater chemistry, and compared the patterns among individual taxonomical groups. We hypothesised a different importance of environmental heterogeneity and seasonal changes primarily linked to strong tufa precipitation, which causes stronger environmental filtering in the calcareous fen. In concordance, we observed that assemblages of the more homogenous calcareous fen primarily changed over time, due to seasonal shifts in source availability and favourable conditions. Their spatial distribution was determined by the amount of CPOM, tufa crusts and temperature variation, but a substantial part of the assemblage exhibited spatial uniformity (Plecoptera, Clitellata, and especially Trichoptera and Diptera). The assemblages of the more heterogeneous Sphagnum-fen were primarily driven by water pH and substrate and the season was a notably weaker predictor. We found that different macroinvertebrate groups can display various responses to the measured variables shaping the overall pattern obtained based on the whole community. Further, greater environmental heterogeneity can result in temporally stable species distribution patterns even at very small spatial scales within a single site.