Summary 1. The spatial patterns of groundwater biodiversity in Europe remain poorly known, yet their knowledge is essential to understand local variation in groundwater assemblages and to develop sound conservation policies. We explore here the broad‐scale distribution of groundwater biodiversity across Europe, focussing on obligate subterranean species. 2. We compiled published distributional data of obligate subterranean aquatic taxa for six European countries (Belgium, France, Italy, Portugal, Slovenia and Spain), and conducted a detailed biological survey of six regions (one in Belgium, two in France, one in Italy, one in Slovenia and one in Spain). Based on this data set, we mapped spatial patterns of biodiversity in Europe on a cell grid with 0.2 × 0.2 ° resolution. 3. As of mid‐2006, the total number of described stygobiotic species in the six countries was 930 and the total number of genera with at least one described stygobiotic species was 191. The total number of sampling sites where at least one stygobiont had been collected was 4709, distributed in 1228 of the 4668 grid cells covering the study area. 4. Groundwater stygobiotic biodiversity was dominated by Crustacea with 757 species in 122 genera. Insects were represented by only two species of a single genus of dytiscid beetles restricted to south‐eastern France. 5. The geographic distribution of stygobionts was extremely heterogeneous. Stygobionts were recorded in 26% of the 4668 grid cells and only 33 cells had more than 20 stygobiotic species. These 33 ‘hot‐cells’ of groundwater species richness clustered in seven hotspots. 6. Endemicity was very high, with 43% of the total number of stygobiotic species restricted to a single cell, i.e. <500 km2. 7. Hotspots defined by rarity, number of genera, number of genera with only one species known in Europe, or number of monospecific genera differed markedly in ranking from those based on species richness. However, a core of four hotspots emerged in all cases: one stretching across Slovenia and northeastern Italy, one in the French Pyrenees, one in the Cévennes in southern France and one in the Rhine River valley in northeastern France. 8. Unevenness in stygobiont distribution cannot be explained solely by unevenness in sampling effort. This is indicated in particular by the fact that our comprehensive sampling survey roughly matched the level of taxonomic richness of the studied regions based on previously published information. 9. With sampling effort continuing, a twofold or higher increase in species richness can be expected in several Mediterranean areas, with a potential to discover up to 50% more new species than are currently known in the region.

Summary 1. Species assemblages of diatoms, rotifers, chydorids, planktonic crustaceans and chironomids were studied in 235 alpine lakes in the Alps, Pyrenees, Tatras (Western Carpathians), Retezat (Southern Carpathians) and Rila Mountains (Balkans). 2. For all taxonomic groups we found a hierarchical structure in the community assemblage using distinct scales of lake clustering (number of k‐means groups) based on species composition similarity (Hellinger distance). We determined the optimal partition in assemblage types (i.e. number of lake clusters) for each taxonomic group by maximising the sum of the taxon indicative value (IndVal) and performed discriminant analyses, using environmental variables not conditioned by geographical patterns. Relevant environmental variables differed among and within taxonomic groups. Therefore the assemblages respond to a complex environmental mosaic, with the exception of diatom assemblages, which followed an acid–base gradient. 3. The significant environmental variables could be grouped into four general factors: lake size, tropho‐dynamic status, acid–base balance and ice‐cover duration (i.e., altitudinal gradient). Lake size was significant for the highest number of assemblage types; however, the most significant factor differed among taxonomic groups: acid–base balance for diatoms, lake size for rotifers, ice‐cover duration for chydorids and planktonic crustaceans and tropho‐dynamic status for chironomids. No single environmental typology accounted for the assemblage structure of all taxonomic groups. 4. However, defining ecological thresholds as values within environmental gradients at which the rate of change in assemblages is accelerated relative to points distant from that threshold, we were able to find specific threshold values for each of the four main general environmental factors identified, which were relevant across several taxonomic groups: 3 ha for lake area; 0.6 mg L−1 for dissolved organic carbon; 190 days for ice‐cover duration and 200 μeq L−1 for acid neutralising capacity. Above and below these values ecosystem organisation change substantially. They have direct applications in establishing lake typologies for environmental quality and biodiversity conservation programmes, and in improving predictions about global change impacts.

1.A survey of c. 350 remote high altitude and high latitude lakes from 11 different mountain regions was undertaken to explore species distribution across Europe at a scale not previously attempted. 2. Lakes were sampled for planktonic crustaceans, rotifers, littoral invertebrates and subfossil chironomids, diatoms and cladocerans. Each lake was characterised in terms of water chemistry, morphology, catchment attributes and geographical location. 3. Separate TWI NSPAN TWI NSPAN analyses were undertaken on diatom, chironomid, planktonic crustacean, littoral invertebrate and cladoceran (chydorids only) data to classify sites according to taxonomic composition. For most datasets there was a spatial component to the classification with distinct geographical groups emerging -Norway and Scotland, Finland and Central ⁄ Eastern Europe. 4. Constrained ordination methods were employed to examine how species responded to a range of environmental factors, which were aggregated into a series of component groups -proximal environment (the chemical, trophic and physical attributes of the lake), catchment characteristics and geographical location. Several key environmental gradients were identified, which explained significant levels of the variance across several of the biological groups including dissolved organic carbon (chironomids, planktonic crustaceans), temperature (chironomids and littoral invertebrates), chloride ⁄ sea-salt (littoral invertebrates, diatoms and rotifers), lake morphology (all groups), calcium ⁄ pH (diatoms), nitrate (chydorids, littoral invertebrates, rotifers and planktonic crustaceans) and fish (littoral invertebrates). In some cases these statistical relationships are likely to represent direct ecological constraints and, in others, it is probable that the environmental variable is acting as a surrogate for some other attribute or process. 5. Variance partitioning was undertaken to quantify how much of the variation in each biological group could be uniquely attributed to variables representing the proximal environment, catchment characteristics and geographical location. For most groups the Correspondence: Martin Kernan, Environmental

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Morphological study of Alona protzi Hartwig, 1900, Alona phreatica Dumont, 1983 and Alona smirnovi Petkovski & FloAner, 1972 reveals close affinities with Alona labrosa Vasiljeva & Smirnov, 1969. We separate these four species from the polyphyletic Alona Baird, 1843 (Anomopoda: Chydoridae). United under Phreatalona gen. nov., these taxa share primitive features on the limbs, together with specializations to a rheic life mode. Phreatalona contains some of the only true hyporheic taxa within the Cladocera. Endemism in two ancient lakes (P. smirnovi and P. labrosa) and a preference for river sediments in Europe (P. phreatica and P. protzi) suggest a long isolation from typical littoral/benthic biotopes. We discuss close links with southern vicariant Nicsmirnovius, the position of these (hypo)rheic chydorids within the subfamily and their affinities with Acroperus. We remark an independent evolution of external (habitus, postabdomen) vs. internal (limb) morphology in the protzi-complex. Phreatalona is likely tertiary in origin, evolving from a littoral alonine adapting to rheic and finally hyporheic environments. Baikal endemic P. labrosa is likely the most primitive species of the genus. We discuss adaptations and evolution in the hyporheic and the effect on dispersal and biogeography of Phreatalona

SUMMARY1. Estimates of species richness obtained from exhaustive field inventories over large spatial scales are expensive and time-consuming. For this reason, efficiency demands the use of indicators as 'surrogates' of species richness. Biodiversity indicators are defined herein as a limited suite of taxonomic groups the species richness of which is correlated with the species richness of all other taxonomic groups present in the survey area. 2. Species richness in ground water was assessed at different spatial scales using data collected from six regions in Europe. In total, 375 stygobiotic species were recorded across 1157 sites and 96 aquifers. The taxonomic groups collected from more than one site and with more than two species (Oligochaeta, Gastropoda, Cyclopoida, Harpacticoida, Ostracoda, Isopoda, Amphipoda, Bathynellacea and Acari) were used to develop nonparametric models to predict stygobiotic biodiversity at the aquifer scale. 3. Pair-wise correlations between taxonomic groups were low, i.e. variation in species richness of a single taxonomic group did not usually reflect variation of the other groups. In contrast, multiple regressions calculated between species richness of any combination of taxa and extra-group species richness along the six regions resulted in a number of significant relationships. 4. These results suggest that some taxonomic groups (mainly Copepoda and Amphipoda and, to a lesser extent, Oligochaeta and Gastropoda) combined in different ways across the regions, were good biodiversity indicators in European groundwater ecosystems. However, the uneven distribution of taxonomic groups prevented selection of a common set of indicators for all six regions. Faunal differences among regions are presumably related to both historical and ecological factors, including palaeogeography, palaeoecology, geology, aquifer fragmentation and isolation, and, less clearly, anthropogenic disturbance.

A series of sampling sites in 6 caves was established in order to assess the importance of the epikarst as a habitat of high diversity of meiofauna. Special attention was given to the stygobiotic species of copepods (Crustacea), which was the most abundant group of animals in water percolating from the epikarst. In total, 37 species of Copepoda were identified from nearly 4000 individuals. To understand epikarst as a habitat, 14 structural and physico-chemical factors were analysed and compared for two types of habitats, i.e., trickles and pools filled with percolation water. Results of canonical correspondence analysis indicated that the significant factors influencing faunal composition in trickles were the thickness of the cave ceiling and the concentrations of sodium, nitrate, and potassium ions, while in pools correlations between individual species and pH, temperature, volume, conductivity and thickness of the cave ceiling were statistically significant. Aquatic copepod assemblages in epikarst encompass species of different taxonomic affinities, living in severe environmental conditions. The pattern of the system reflects both high taxonomic diversity and environmental severity.