Groundwater dependent ecosystems (GDEs) include valuable ecosystems such as springs, wetlands, rivers, lakes and lagoons. The protection of these systems and services they provide is highlighted by international agreements, i.e. Ramsar convention on wetlands, and regional legislation, i.e. the European Water Framework Directive. Groundwater provides water, nutrients and a relatively stable temperature. However, the role of groundwater in surface ecosystems is not fully understood. The ecosystem can depend on groundwater directly or indirectly, and the reliance can be continuous, seasonal or occasional. This has implications for the vulnerability of ecosystems, as some may be easily affected by external pressure. Conceptual models and quantitative assessments of how groundwater interacts with the environment are needed. GDEs are also threatened by different land use activities and climate change. Hence, we need to understand how GDEs are affected by changes in

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DNA sequences offer powerful tools for describing the members and interactions of natural communities. In this study, we establish the to-date most comprehensive library of DNA barcodes for a terrestrial site, including all known macroscopic animals and vascular plants of an intensively studied area of the High Arctic, the Zackenberg Valley in Northeast Greenland. To demonstrate its utility, we apply the library to identify nearly 20 000 arthropod individuals from two Malaise traps, each operated for two summers. Drawing on this material, we estimate the coverage of previous morphology-based species inventories, derive a snapshot of faunal turnover in space and time and describe the abundance and phenology of species in the rapidly changing arctic environment. Overall, 403 terrestrial animal and 160 vascular plant species were recorded by morphology-based techniques. DNA barcodes (CO1) offered high resolution in discriminating among the local animal taxa, with 92% of morphologically distinguishable taxa assigned to unique Barcode Index Numbers (BINs) and 93% to monophyletic clusters. For vascular plants, resolution was lower, with 54% of species forming monophyletic clusters based on barcode regions rbcLa and ITS2. Malaise catches revealed 122 BINs not detected by previous sampling and DNA barcoding. The insect community was dominated by a few highly abundant taxa. Even closely related taxa differed in phenology, emphasizing the need for species-level resolution when describing ongoing shifts in arctic communities and ecosystems. The DNA barcode library now established for Zackenberg offers new scope for such explorations, and for the detailed dissection of interspecific interactions throughout the community.

Summary1. Metacommunity research relies largely on proxies for inferring the effect of dispersal on local community structure. Overland and watercourse distances have been typically used as such proxies. A good proxy for dispersal should, however, take into account more complex landscape features that can affect an organism's movement and dispersal. The cost distance approach does just that, allowing determining the path of least resistance across a landscape. 2. Here, we examined the distance decay of assemblage similarity within a subarctic stream insect metacommunity. We tested whether overland, watercourse and cumulative cost distances performed differently as correlates of dissimilarity in assemblage composition between sites. We also investigated the effect of body size and dispersal mode on metacommunity organization. 3. We found that dissimilarities in assemblage composition correlated more strongly with environmental than physical distances between sites. Overland and watercourse distances showed similar correlations to assemblage dissimilarity between sites, being sometimes significantly correlated with biological variation of entire insect communities. In metacommunities deconstructed by body size or dispersal mode, contrary to our expectation, passive dispersers showed a slightly stronger correlation than active dispersers to environmental differences between sites, although passive dispersers also showed a stronger correlation than active dispersers to physical distances between sites. The strength of correlation between environmental distance and biological dissimilarity also varied slightly among the body size classes. 4. After controlling for environmental differences between sites, cumulative cost distances were slightly better correlates of biological dissimilarities than overland or watercourse distances between sites. However, quantitative differences in correlation coefficients were small between different physical distances. 5. Although environmental differences typically override physical distances as determinants of the composition of stream insect assemblages, correlations between environmental distances and biological dissimilarities are typically rather weak. This undetermined variation may be attributable to dispersal processes, which may be captured using better proxies for the process. We suggest that further modifying the measurement of cost distances may be a fruitful avenue, especially if complemented by more direct natural history information on insect dispersal behaviour and distances travelled by them.

We studied variation in benthic macrocrustacean and insect assemblages in relation to spring habitat characteristics in six springs located in a single groundwater area in south-west Finland. We defined five habitat types in the studied springs according to water flow and benthic substrate characteristics: minerogenic brooks, organogenic brooks, helocrenes, floating moss carpets and limnocrene pools. Most studied invertebrate orders, as well as individual taxa, showed differences in relative abundances between the habitat types, but the most common taxa occurred in all springs and habitat types. The studied macroinvertebrates were most abundant in the moss carpet sites and least abundant in the pool sites, but the difference was not statistically significant. We did not observe significant differences in mean taxonomic richness per sample between habitat classes. The observed taxonomic richness in pooled samples of habitat classes was highest in moss carpet habitat and lowest in pool habitat, and the rarefied richness estimate was lowest in pool habitat. Benthic macrocrustacean and insect assemblages varied more between habitat types than between individual springs. In an Nonmetric Multidimensional Scaling ordination analysis, spring brook sites were separated from the moss carpet and pool sites, whereas helocrene sites were widely scattered among sites in other habitat classes. The strongest ecological gradients were related to water flow and the presence of minerogenic substrate, separating lentic and lotic habitats. Abundances of moss and coarse detritus accounted for most of the within-class variation. We identified several indicator species for minerogenic and organogenic brooks and for moss carpet and pool habitats, but none for the helocrenes. We found several occurrences of two crenobiont insect species considered threatened in Finland. We suggest that combined studies on macroinvertebrate and bryophyte assemblages would be a powerful approach in assessing the biodiversity of springs.