The processing of organic matter is a central ecosystem function in freshwater ecosystems that allows the integration of terrestrial plant material into aquatic heterotrophic food webs. From an energetic perspective, many temperate aquatic systems fundamentally depend on such allochthonous inputs. Current research established that this central ecosystem function may be linked to population density of few yet dominant invertebrate detritivores, showing a negative density-dependent relationship. Here, we extended the current knowledge and experimentally assessed the processing of leaf litter by freshwater amphipods, using broad density gradients, interspecific competition, and laboratory and field experiments. We used two species of dominant amphipods, namely the native Gammarus fossarum and the non-native Gammarus roeselii, varying their density by two orders of magnitude, both in single-species and two-species treatments. Results from 252 mesocosms in the lab and 97 mesocosms in a natural stream show that per capita leaf litter processing rates are strongly negatively dependant on population density in monocultures. Interspecific competition in two-species treatments corroborated the negative density-dependent ecosystem function and highlighted the functional redundancy of the two detritivore species. We identified a flattening in the processing rates at previously unreported but well-defined breakpoints. These breakpoints may reflect minimal metabolic requirements at which survival takes precedence over any other process, such as interference competition. The breakpoints were consistent across both species, indicating that they must process leaf litter equalling one fifth to one fourth of their dry body weight per day as a minimal nutritional threshold. Our results suggest the need for integrating nonlinear density-dependencies and breakpoint population densities in key ecosystem functions in aquatic ecosystem modelling alongside classical biodiversity–ecosystem functioning relationships. Depending on the population density the corresponding ecosystem functioning could differ significantly. This corroborates the need to better understand biodiversity–abundance relationships when protecting aquatic ecosystems.

99 Schweizer Bäche wurden anhand der beiden Bioindikatoren Fische und Makrozoobenthos untersucht. Die Mehrzahl der Bäche zeigt wesentliche Defizite beim ökologischen Zustand. So fehlen beispielsweise in über 70% der Gewässer ein Teil der pestizidempfindlichen Insektenlarven und andere Kleinlebewesen. Statistische Auswertungen deuten darauf hin, dass sich eine beeinträchtigte Ökomorphologie sowie ein grosser Anteil Ackerland, Obstkulturen und Reben im Einzugsgebiet besonders negativ auf das Vorkommen dieser empfindlichen Kleinlebewesen auswirken.

Understanding spatial patterns of biodiversity within the context of long-term climatic shifts is of high importance, particularly in the face of contemporary climate change. In comparison to aboveground taxa, subterranean organisms respond to changing climates with generally much lower dispersal and recolonization potential, yet possible persistence in refugial groundwater habitats under ice-shields. However, knowledge on general and geographically large-scale effects of glaciation on contemporary groundwater biodiversity patterns is still very limited. Here, we tested how Late Pleistocene glaciation influenced the diversity and distribution of 36 groundwater amphipod species in Alpine and peri-Alpine regions, characterized by extensive glaciation cycles, and how its legacy explains contemporary diversity patterns. We based our analysis on an unprecedented density of ~ 1000 systematic sampling sites across Switzerland. Using presence–absence data, we assessed biodiversity and species' ranges, and calculated for each site within-catchment distance to the Last Glacial Maximum (LGM) glacier extent. We then applied a sliding window approach along the obtained distance gradient from LGM ice-covered to ice-free sites to compute biodiversity indices reflecting local richness, regional richness, and differentiation, respectively. We found a strong signal of the LGM ice extent on the present-day distribution of groundwater amphipods. Our findings revealed pronounced species turnover and spatial envelopes of individual species' occurrences in formerly ice-covered, ice-free, or transitional zones, respectively. While local richness remained constant and low along the LGM distance gradient, groundwater communities in LGM ice-covered areas were more similar to each other and had lower gamma diversities and decreased occurrence probabilities per sliding window compared to communities in Pleistocene ice-free areas. These results highlight the significant impact of Pleistocene glaciation on shaping biodiversity patterns of subterranean communities and imprinting contemporary distribution of groundwater organisms.

Aim: groundwater ecosystems harbour a unique biodiversity, but remain poorly studied, mainly due to difficulties in accessibility and imperfect species detection. Consequently, knowledge on the state and change of groundwater biodiversity remains highly deficient. In the context of global warming and excessive groundwater extraction, understanding groundwater from an ecosystem-perspective, including organism diversity and distribution, is essential. This study presents the largest ever systematic assessment of groundwater amphipods, which are a key component of European groundwater biodiversity.
Location: Switzerland (41,285 km²), including data from 906 sampling sites.
Taxon: groundwater amphipods, genera Niphargus and Crangonyx (Crustacea, Amphipoda).
Methods: we applied a highly standardized citizen science approach to collect repeated groundwater fauna samples in collaboration with municipal drinking water providers. Using detection–nondetection data of the genetically identified groundwater amphipod species, we assessed the overall species diversity of both rare and common species. The distribution of commonly found species was predicted using multispecies occupancy modelling.
Results: we retrieved 3882 samples from 906 sites, yielding 2350 groundwater amphipod individuals. We identified a remarkable species diversity, comprising few commonly and many rarely found species. Considering commonly found species, we identified distinct distribution ranges, low local species richness and a predominance of negative co-occurrences. In contrast, a major portion of species were found rarely (generally at just one or two sites each), distributed uniformly throughout the study area and unrelated to common species' recognized hotspots. Many of these rarely found species are not yet formally described.
Main conclusions: Our results give robust emphasis on the rare occurrence and narrow distribution of many groundwater dwellers. Our systematic and standardized sampling data of groundwater amphipods suggest that rarity is particularly prominent and inherent to groundwater organisms. We emphasize the need of systematic data to integrate rare groundwater species in biodiversity assessments, especially in times of global change.

Implementing and optimizing biodiversity monitoring is crucial given the current, worldwide biodiversity decline. Compared to other ecosystems, monitoring of biodiversity is lagging behind in groundwater ecosystems, both because of sparse taxonomic knowledge and methodological constraints. We here assessed temporal variation in the occurrence and abundance of macroinvertebrates collected systematically from shallow groundwater aquifers of Switzerland to establish general principles on seasonality and repeatability of assessment outcomes. We found no seasonal abundance pattern for obligate groundwater amphipods and isopods, indicating temporal consistency. In contrast, other macroinvertebrates (predominantly stygophiles and stygoxenes) showed pronounced seasonality in their detection rate. However, we found variability in detection rates across groundwater amphipod species and especially across sampling sites. For groundwater communities, characterized by narrowly-distributed and rare species, our results highlight the need for tailored and extensive sampling strategies. When setting up monitoring programs on groundwater fauna, detection probability, temporal autocorrelation, and standardization of sampling effort should be carefully considered. Applying novel, systematic approaches, can offer promising methodologies for understanding and conserving groundwater ecosystems.

Terrestrial and aquatic ecosystems are tightly linked, with direct implications for applied resource management and conservation. It is well known that human land use change and intensification of terrestrial systems can have large impacts on surface freshwater ecosystems. Contrastingly, the study and understanding of such land use impacts on groundwater communities is lagging behind. Both the impact strength of land use on groundwater communities and the spatial extents at which such interlinkages are operating are largely unknown, despite our reliance on groundwater for drinking water extraction as a key ecosystem service. Here, we analyzed groundwater amphipod occurrence from several hundred shallow groundwater aquifers used for drinking water extraction across a region of varying agricultural intensity and human population density in Switzerland. Despite drinking water extraction sites being generally built at locations with expected minimal aboveground impacts on water quality, we found a direct correlation between land use type and intensity within the surrounding catchment area and the locally measured nitrate concentrations, which is a direct proxy for drinking water quality. Furthermore, groundwater amphipods were more likely to be found at sites with higher forest coverage than at sites with higher crop and intensive pasture coverages, clearly indicating a tight connection between aboveground land use and groundwater biodiversity. Our results indicate that land use type effects on groundwater communities are most relevant and pronounced to spatial scales of about 400–1000 m around the groundwater sampling site. Importantly, the here identified spatial scale is 1.2- to 3-fold exceeding the average extent of currently defined groundwater protection zones. We postulate that incorporating an ecosystem perspective into groundwater management strategies is needed for effective protection of groundwater quality and biodiversity.

Subterranean ecosystems (comprising terrestrial, semi-aquatic, and aquatic components) are increasingly threatened by human activities; however, the current network of surface-protected areas is inadequate to safeguard subterranean biodiversity. Establishing protected areas for subterranean ecosystems is challenging. First, there are technical obstacles in mapping three-dimensional ecosystems with uncertain boundaries. Second, the rarity and endemism of subterranean organisms, combined with a scarcity of taxonomists, delays the accumulation of essential biodiversity knowledge. Third, establishing agreements to preserve subterranean ecosystems requires collaboration among multiple actors with often competing interests. This perspective addresses the challenges of preserving subterranean biodiversity through protected areas. Even in the face of uncertainties, we suggest it is both timely and critical to assess general criteria for subterranean biodiversity protection and implement them based on precautionary principles. To this end, we examine the current status of European protected areas and discuss solutions to improve their coverage of subterranean ecosystems.

Groundwater is a vital ecosystem of the global water cycle, hosting unique biodiversity and providing essential services to societies. Despite being the largest unfrozen freshwater resource, in a period of depletion by extraction and pollution, groundwater environments have been repeatedly overlooked in global biodiversity conservation agendas. Disregarding the importance of groundwater as an ecosystem ignores its critical role in preserving surface biomes. To foster timely global conservation of groundwater, we propose elevating the concept of keystone species into the realm of ecosystems, claiming groundwater as a keystone ecosystem that influences the integrity of many dependent ecosystems. Our global analysis shows that over half of land surface areas (52.6%) has a medium-to-high interaction with groundwater, reaching up to 74.9% when deserts and high mountains are excluded. We postulate that the intrinsic transboundary features of groundwater are critical for shifting perspectives towards more holistic approaches in aquatic ecology and beyond. Furthermore, we propose eight key themes to develop a science-policy integrated groundwater conservation agenda. Given ecosystems above and below the ground intersect at many levels, considering groundwater as an essential component of planetary health is pivotal to reduce biodiversity loss and buffer against climate change.

Die noch wenig bekannte Grundwasserfauna ist ein bedeutender Bestandteil der Biodiversität und trägt zum Funktionieren der Grundwasserökosysteme bei. Mittels Proben aus der ganzen Schweiz wurde erstmals ein Überblick über die Vielfalt und Biomasse von Makroinvertebraten in Grundwasserökosystemen der Schweiz erarbeitet. Eine hohe Diversität an ober- und unterirdischen Makroinvertebraten im Grundwasser weist auf die Wechselwirkungen zwischen Oberfläche und Grundwasser hin.

The genus Niphargus is the most diverse subterranean amphipod genus in the western Palearctic region but, owing to the presence of cryptic species and homoplasy, its taxonomy and biogeographic scenarios are complex, making molecular methods essential to understand its evolution. We conducted a study combining dna-based taxonomy with traditional morphotaxonomy to investigate Niphargus bihorensis Schellenberg, 1940, known from the Western Alps and Carpathians. We redescribed the type material of N. bihorensis from Bihor County, Romania, and revealed the presence of a cryptic species, N. absconditus n. sp., in the same karstic area (Pădurea Craiului Mountains). Additionally, the Alpine populations previously attributed to N. bihorensis turned out to belong to a new, not so closely related species, N. tizianoi n. sp. Phylogenetic analyses based on a concatenated dataset of one mitochondrial and two nuclear markers suggest that the N. bihorensis species complex belongs to a strongly supported clade, together with several species distributed from Switzerland to Iran.

Amphipods are among the most abundant macroinvertebrates in freshwater ecosystems of the Palaearctic and crucial for ecosystem functioning. Furthermore, their high sensitivity to environmental change and pollutants makes them widely used model organisms in environmental sciences and ecotoxicology. In field studies and surveys across Eurasia, species of the genus Gammarus are commonly used, yet laboratory-based studies and ecotoxicological tests are often restricted to the in most parts of the world non-native Hyalella azteca, as Gammarus is much harder to breed and maintain under laboratory conditions. However, for direct comparisons and extrapolations of results of field- vs. laboratory-based studies, the use of the same species would be desirable. Here, we investigated different settings with respect to feeding, shelter and day length to successfully increase survival, juvenile production and their respective growth and survival, and ultimately multi-generation breeding of the amphipod Gammarus fossarum. Amphipod populations persisted and reproduced successfully under optimized husbandry conditions for 12 months and were partially maintained for another year in populations up to a few hundred individuals. Specifically, supplementing diet with protein-rich food sources as well as the provisioning of shelters improved survival rate of G. fossarum significantly. However, we found no significant effect of different day length treatments on the overall relative reproductive activity or on the total amphipod abundance maintained. We conclude that G. fossarum can be kept and reared under standardized conditions. Despite the longer generation times of G. fossarum and higher effort required for maintenance compared to H. azteca, direct ecological relevance and comparability of results to natural systems may justify its future use and development as a study organism for environmental sciences and ecotoxicology.

Despite being the most important source of liquid freshwater on the planet, groundwater is severely threatened by climate change, agriculture, or industrial mining. It is thus extensively monitored for pollutants and declines in quantity. The organisms living in groundwater, however, are rarely the target of surveillance programmes and little is known about the fauna inhabiting underground habitats. The difficulties accessing groundwater, the lack of expertise, and the apparent scarcity of these organisms challenge sampling and prohibit adequate knowledge on groundwater fauna. Environmental DNA (eDNA) metabarcoding provides an approach to overcome these limitations but is largely unexplored. Here, we sampled water in 20 communal spring catchment boxes used for drinking water provisioning in Switzerland, with a high level of replication at both filtration and amplification steps. We sequenced a portion of the COI mitochondrial gene, which resulted in 4917 ASVs, yet only 3% of the reads could be assigned to a species, genus, or family with more than 90% identity. Careful evaluation of the unassigned reads corroborated that these sequences were true COI sequences belonging mostly to diverse eukaryotic groups, not present in the reference databases. Principal component analyses showed a strong correlation of the community composition with the surface land-use (agriculture vs. forest) and geology (fissured rock vs. unconsolidated sediment). While incomplete reference databases limit the assignment of taxa in groundwater eDNA metabarcoding, we showed that taxonomy-free approaches can reveal large hidden diversity and couple it with major land-use drivers, revealing their imprint on chemical and biological properties of groundwater.

Groundwater is the physically largest freshwater ecosystem, yet one of the least explored habitats on earth, both because of accessing difficulties and the scarcity of the organisms inhabiting it. Here, we demonstrate how a two-fold approach provides complementary information on the occurrence and diversity of groundwater amphipods. Firstly, we used a citizen science approach in collaboration with municipal water providers who sampled groundwater organisms in their spring catchment boxes over multiple weeks, followed by DNA barcoding. Secondly, we collected four 10 L water samples at each site, in one sampling event, for environmental DNA (eDNA) metabarcoding. We found that citizen science was very effective in describing the distribution and abundance of groundwater amphipods. Although the single time-point of eDNA sampling did not detect as many amphipods, it allowed the assessment of the entire groundwater community, including microorganisms. By combining both methods, we found different amphipod species co-occurring with distinct sequences from the eDNA-metabarcoding dataset, representing mainly micro-eukaryotic species. We also found a distinct correlation between the diversity of amphipods and the overall biodiversity of groundwater organisms detected by eDNA at each site. We thus suggest that these approaches can be used to get a better understanding of subterranean biodiversity.

Subterranean fauna is an important contributor to the global fauna, but it is still understudied and a large part of its taxonomy is not yet resolved. One species complex with unresolved taxonomy is the groundwater amphipod Niphargus ruffoi, endemic to the Alpine chain. Here, we used new samples from across the Alpine arc to review the taxonomic status of the entire clade, including the species N. ruffoi and Niphargus arolaensis. We sequenced four genetic markers from the collected specimens, assessed the phylogenetic position of N. ruffoi within the genus, and studied the structure of this species complex using four molecular species delimitation methods. We tested for recombination using the alignments of the concatenated nuclear rDNA genes. The phylogenetic analyses revealed high support for the monophyly of the studied species complex, defining two lineages (i.e., N. arolaensis and N. ruffoi) within the clade. Molecular species delimitation methods suggested that N. arolaensis is a single species, while N. ruffoi should be considered as a species complex of three (using ITS) to eight (using COI) putative species. Moreover, we found a discrepancy between the different nuclear ribosomal DNA markers, indicating a possible recombination with fragments of 28S DNA of N. ruffoi s. lat. present in the genome of N. arolaensis. For the above-mentioned reasons, the internal phylogenetic structure of N. ruffoi s. lat. could not be fully resolved. Moreover, no clear morphological evidence supported the molecular species delimitation. Consequently, no taxonomic changes were proposed. We postulate that this complex scenario was influenced by Pleistocene climate oscillations with subsequent fragmentation events and secondary contacts, making this an interesting study system to investigate the evolution and biogeography of Alpine clades.

Haplotaxis Hoffmeister, 1843 is a poorly known genus: it is rarely collected because it usually inhabits groundwater, and many species are only known from immature specimens. Even the type species, Haplotaxis gordioides (Hartmann in Oken, 1819) remains poorly defined because of the absence of reference types. Most of the Haplotaxis species have been placed in synonymy with H. gordioides since the end of the 19^th century, a situation that has remained essentially unchanged until now. As a result, the species is supposedly present on all continents, except Antarctica. This observation is all the more questionable as the aquatic subterranean environment is nowadays well known to harbour many species with restricted distribution, due to the strong hydrogeographic isolation and the low dispersal abilities of its inhabitants.
In this study we assessed the hypothesis of H. gordioides as a single species with wide distribution versus a complex of cryptic species with narrow distribution. We used a DNA-barcoding approach based on the COI mitochondrial marker of 46 Haplotaxis specimens collected in Switzerland, mostly as part of a countrywide sampling campaign to study groundwater macroinvertebrates.
Preliminary results suggested that H. gordioides is a complex of at least 6 cryptic species in Switzerland, which has important implications both for the knowledge of the exact identity of the type species and for the synonymy of most of the species described in the 19^th century. However, as it is based on a single-locus approach, this study should be seen as the first step in an integrative taxonomic process that should include additional biological material, the study of complementary markers (especially nuclear), and the morphological study of specimens.

À ce jour, dix espèces exotiques ont pu s’établir en Suisse, dont cinq sont considérées comme envahissantes. Certaines de ces espèces ont, en partie, un régime alimentaire de prédateur ou se nourrissent en filtrant l’eau; leurs populations peuvent atteindre des densités très élevées. Jusqu’à présent, les espèces exotiques ne sont toutefois apparues que dans de grand lacs et cours et plans d’eau.

Until now, ten alien amphipod species have become established in Switzerland, of which five are considered invasive. These species are partly predatory or are filter-feeding and can reach very high densities. Until now they have only been found in larger bodies of water, however.

In der Schweiz konnten sich bislang zehn gebietsfremde Arten etablieren. Von diesen zählen fünf zu den invasiven Arten. Einige dieser Arten ernähren sich teilweise räuberisch oder leben filtrierend und können sehr hohe Dichten erreichen. Allerdings sind die gebietsfremden Arten bisher nur in grösseren Gewässern aufgetreten.

Quaternary climate fluctuations can affect speciation in regional biodiversity assembly in two non-mutually exclusive ways: a glacial species pump, where isolation in glacial refugia accelerates allopatric speciation, and adaptive radiation in underused adaptive zones during ice-free periods. We detected biogeographic and genetic signatures associated with both mechanisms in the assembly of the biota of the European Alps. Age distributions of endemic and widespread species within aquatic and terrestrial taxa (amphipods, fishes, amphibians, butterflies and flowering plants) revealed that endemic fish evolved only in lakes, are highly sympatric, and mainly of Holocene age, consistent with adaptive radiation. Endemic amphipods are ancient, suggesting preglacial radiation with limited range expansion and local Pleistocene survival, perhaps facilitated by a groundwater-dwelling lifestyle. Terrestrial endemics are mostly of Pleistocene age and are thus more consistent with the glacial species pump. The lack of evidence for Holocene adaptive radiation in the terrestrial biome is consistent with faster recolonization through range expansion of these taxa after glacial retreats. More stable and less seasonal ecological conditions in lakes during the Holocene may also have contributed to Holocene speciation in lakes. The high proportion of young, endemic species makes the Alpine biota vulnerable to climate change, but the mechanisms and consequences of species loss will likely differ between biomes because of their distinct evolutionary histories.

Erstmals wurde die Verbreitung und Artenvielfalt der Grundwasserflohkrebse im Einzugsgebiet der Töss, unter Einbezug von Bürgerwissenschaft, räumlich hoch aufgelöst untersucht. Bei der durch die Eawag und die Universität Zürich durchgeführten Studie wurden vier in dieser Region bisher noch nicht nachgewiesene Arten gefunden. Auch konnte gezeigt werden, dass die Daten zur Fundwahrscheinlichkeit, die von Wasserversorgungen erhoben wurden, mit denen von Forschenden vergleichbar sind.

Knowledge on the diversity and distribution of subterranean organisms is still scattered, even in faunistically relatively well-researched countries such as Switzerland. This is mostly due to the restricted access to these subterranean habitats. Better knowledge on these organisms is needed, because they contribute substantially to overall biodiversity of a region, often contain unique elements of biodiversity, and can potentially be indicative of the ecological status of subterranean ecosystems that are providing important ecosystem services such as drinking water. Past research on subterranean organisms has often used highly specialised sampling techniques and expert knowledge. Here, we show that inclusion of non-professionals can be an alternative and highly promising sampling strategy. We retrieved citizen science-based samples from municipal groundwater wells across Switzerland, mainly from the Swiss Plateau. Opportunistic samples from 313 sites revealed a previously undocumented groundwater fauna including organisms from different major invertebrate groups, with a dominance of crustaceans. Here, we studied amphipods of the genus Niphargus. Among all 363 individuals sampled, we found in total eight nominal species. Two of them, namely N. fontanus and N. kieferi, are reported for Switzerland for the first time. We also found four further phylogenetic lineages that are potentially new species to science. One of them is here formally described as Niphargus arolaensis sp. nov. The description is based on molecular and morphometric data. Our study proves the suitability of citizen science to document subterranean diversity, supports groundwater conservation efforts with data, and raises awareness for the relevance and biodiversity of groundwater amphipods among stakeholders.

Theory predicts that the distribution of genetic diversity in a landscape is strongly dependent on the connectivity of the metapopulation and the dispersal of individuals between patches. However, the influence of explicit spatial configurations such as dendritic landscapes on the genetic diversity of metapopulations is still understudied, and theoretical corroborations of empirical patterns are largely lacking. Here, we used microsatellite data and stochastic simulations of two metapopulations of freshwater amphipods in a 28,000 km² riverine network to study the influence of spatial connectivity and dispersal strategies on the spatial distribution of their genetic diversity. We found a significant imprint of the effects of riverine network connectivity on the local and global genetic diversity of both amphipod species. Data from 95 sites showed that allelic richness significantly increased towards more central nodes of the network. This was also seen for observed heterozygosity, yet not for expected heterozygosity. Genetic differentiation increased with instream distance. In simulation models, depending on the mutational model assumed, upstream movement probability and dispersal rate, respectively, emerged as key factors explaining the empirically observed distribution of local genetic diversity and genetic differentiation. Surprisingly, the role of site-specific carrying capacities, for example by assuming a direct dependency of population size on local river size, was less clear cut: while our best fitting model scenario included this feature, over all simulations, scaling of carrying capacities did not increase data-model fit. This highlights the importance of dispersal behaviour along spatial networks in shaping population genetic diversity.

Eine Pilotstudie der Eawag hat an über 300 Standorten im Schweizer Mittelland die Grundwasserfauna untersucht, wobei die Rohwässer in Brunnenstuben durch die Wasserversorgungen selbst beprobt wurden. Anschliessend wurden die gefundenen Organismen morphologisch und teils genetisch bestimmt. An über 60% der Stellen wurden Organismen aus 18 biologischen Ordnungen gefunden. Bei den Grundwasserflohkrebsen wurden neue Arten für die Schweiz und auch für die Wissenschaft entdeckt.

Environmental change can alter species' abundances within communities consistently; for example, increasing all abundances by the same percentage, or more idiosyncratically. Here, we show how comparing effects of temperature on species grown in isolation and when grown together helps our understanding of how ecological communities more generally respond to environmental change. In particular, we find that the shape of the feasibility domain (the parameter space of carrying capacities compatible with positive species' abundances) helps to explain the composition of experimental microbial communities under changing environmental conditions. First, we introduce a measure to quantify the asymmetry of a community’s feasibility domain using the column vectors of the corresponding interaction matrix. These column vectors describe the effects each species has on all other species in the community (hereafter referred to as species' multidimensional effects). We show that as the asymmetry of the feasibility domain increases the relationship between species' abundance when grown together and when grown in isolation weakens. We then show that microbial communities experiencing different temperature environments exhibit patterns consistent with this theory. Specifically, communities at warmer temperatures show relatively more asymmetry; thus, the idiosyncrasy of responses is higher compared with that in communities at cooler temperatures. These results suggest that while species’ interactions are typically defined at the pairwise level, multispecies dynamics can be better understood by focusing on the effects of these interactions at the community level.

Amphipods are a highly diverse group of aquatic invertebrates with >10,000 described species globally. About 20% of these are freshwater species, with half of them found in the West Palaearctic. Amphipods inhabit almost all freshwater ecosystems, including lakes, rivers, streams, as well as cave and groundwater systems, and can be the dominant macroinvertebrates in these habitats. They exhibit essential roles in the functioning of aquatic ecosystems, contributing to leaf litter breakdown and serving as important prey for fish. Furthermore, they are commonly used as indicator taxa for biomonitoring and in ecotoxicological studies. Many amphipod communities are currently undergoing rapid changes due to various drivers of global change, and some amphipod species are among the most successful nonnative invasive invertebrates.
Despite their ecological and economic significance, the knowledge on amphipods in Switzerland was hitherto limited, and until now no checklist, distribution maps, or broadscale estimates on genetic, functional, and morphological diversity of all amphipod species in Switzerland existed. All hitherto available literature on the ecology, faunistics, and taxonomy of amphipods in Central Europe is either outdated by many decades (Chevreux & Fage, 1925; Schellenberg, 1942), does not cover all species (Ginet, 1995; Eggers & Martens, 2001, Piscart & Bollache, 2012), or is only targeting neighboring countries (Vornatscher, 1965; Karaman, 1993; Eggers & Martens, 2001; Piscart & Bollache, 2012, Zettler & Zettler, 2017). This lack of appropriate baseline information on the ecology, distribution and faunistics of amphipods in Switzerland is a serious limitation for fundamental ecological research, impedes appropriate conservation strategies, and limits the use and application of amphipods as indicator taxa.

Alpine landscapes are being transformed through the rapid recession of glaciers, resulting in the development of numerous non-glacial headwater streams inhabited by a diverse assemblage of macroinvertebrates. We examined spatial patterns in biodiversity and rarity of macroinvertebrates in 41 non-glacial streams from five glacierized catchments in the Swiss Alps undergoing rapid glacial recession over the last decades. Water physico-chemistry and food resources (periphyton, benthic organic matter) varied widely among streams within each catchment, while no significant differences occurred among catchments. Variability in community composition was similar among streams within each catchment but differed among catchments, reflecting differences in catchment-scale species pools due to biogeographical context and season. Overall, 101 taxa from ca 33,000 individuals collected were identified in the streams with 7–33 taxa found in individual streams. Some 64% of the taxa comprised less than 5% of the total abundances at the streams (rare in abundance) with 78% of the taxa being represented by less than 5% of the most common taxon (Baetis sp.), whereas 47% of the taxa were found in less than 10% of the streams (rare in distribution). No taxon was found at all sites (maximum presence at 85% of the sites), while 15% of the taxa were found at 50% of the sites or more. However, analyzing the rank-abundance distribution showed that rarity was less prevalent than previously shown in other ecosystems. The results indicated that community assembly of alpine headwater streams is a complex interaction between environmental properties (habitat filtering), habitat stability coupled with dispersal (source sink dynamics), and time since deglaciation (island biogeography). Integrating these processes is essential towards understanding ongoing colonization events in headwater streams of alpine catchments as glaciers continue to recede.

The family Moitessieriidae is poorly known, as its members, inhabiting exclusively subterranean waters, are often known only from few minute, empty shells. Molecular studies on their relationships confirmed the distinctness of this family. Their monophyly, however, remained doubtful, since the Moitessieriidae did not form a distinct clade in the phylogenetic tree based on the most commonly applied mitochondrial cytochrome oxidase subunit I (COI), and the representative of the family Cochliopidae occupied a position among the moitessieriid clades. In the present paper two new nuclear loci, namely histone H3 gene and ribosomal internal transcribed spacer ITS2, have been applied to resolve the status of the Moitessieriidae. The resulting phylogenies show the Moitessieriidae as a distinct, well supported clade, confirming thus their monophyly.

Assessing individual components of biodiversity, such as local or regional taxon richness, and differences in community composition is a long‐standing challenge in ecology. It is especially relevant in spatially structured and diverse ecosystems. Environmental DNA (eDNA) has been suggested as a novel technique to detect taxa and therefore may allow to accurately measure biodiversity. However, we do not yet fully understand the comparability of eDNA‐based assessments to classical morphological approaches. We assessed may‐, stone‐, and caddisfly genera with two contemporary methods, namely eDNA sampling followed by molecular identification and kicknet sampling followed by morphological identification. We sampled 61 sites distributed over a large river network, allowing a comparison of various diversity measures from the catchment to site levels and providing insights into how these measures relate to network properties. We extended our data with historical morphological records of total diversity at the catchment level. At the catchment scale, identification based on eDNA and kicknet samples detected similar proportions of the overall and cumulative historically documented richness (gamma diversity), 42% and 46%, respectively. We detected a good overlap (62%) between genera identified from eDNA and kicknet samples at the regional scale. At the local scale, we found highly congruent values of local taxon richness (alpha diversity) between eDNA and kicknet samples. Richness of eDNA was positively related to discharge, a descriptor of network position, while kicknet was not. Beta diversity, a measure of dissimilarity between sites, was comparable for the two contemporary methods and is driven by species replacement and not by nestedness. Although eDNA approaches are still in their infancy and optimization regarding sampling design and laboratory work is still needed, our results indicate that it can capture different components of diversity, proving its potential utility as a new tool for large sampling campaigns across hitherto understudied complete river catchments.

The present thesis investigated the diversity and distribution of amphipods (Crustacea, Amphipoda) in Switzerland, based on an extensive collection and analysis of faunistic and ecological data. The thesis aimed to provide an updated and substantiated foundation for future ecological, faunistic and taxonomic research on these important crustaceans, with a particular focus on Switzerland. The thesis was conducted as integral part of the project Amphipod.CH that was initiated to intensify the study of amphipods in Switzerland.

Amphipods are an order of crustaceans that represent keystone species in a wide variety of both marine and freshwater ecosystems. As such they contribute considerably to the biodiversity and functioning of these ecosystems, for example by linking different trophic levels or as major decomposers. In many temperate freshwater ecosystems (including groundwater, caves, lakes and stream ecosystems), amphipods are among the most abundant macroinvertebrates. However, their communities are also rapidly changing due to their high sensitivity to environmental change and pollution as well as a high proportion of non-native amphipod species reaching new habitats, particularly in Europe and Switzerland. It is therefore crucial to understand their ecology and systematics in order to detect and understand such changes and take appropriate conservation measures.

In my thesis I focused on improving the knowledge and data on amphipods in Switzerland. Before the onset of the project, only limited information on amphipods had been available for the European Alps and Switzerland in particular. With several major European drainage areas originating in Switzerland, the region represents a biogeographically interesting area . In a first step, I reviewed for the thesis all previously published literature on amphipods and summarized this ecological and faunistic information for a general introduction to amphipods of Switzerland. This includes publications from the 19th century up to the most recent findings from 2017. I then described several new species for Switzerland or even new to science based on molecular methods and morphological features, some of these species are even endemic to Switzerland. Based on this work, I developed a fully illustrated species identification key. It covers all 44 species of amphipods known from Switzerland, is fully compliant with recent systematics and covers all drainage areas of Switzerland. Thereby, it is the first ever key to all amphipods of Switzerland. Finally, I analyzed amphipod communities using diversity analyses covering a large spatial extent to get a deeper insight into the importance of dispersal and habitat connectivity for community formation and biological invasions in dendritic networks. Overall, my thesis improves and synthesizes the knowledge on amphipods of Switzerland and will hopefully ease all future work on this important taxonomic group of freshwater macroinvertebrates.

Habitat connectivity crucially influences dispersal of organisms. It is especially seen as an important driver of the spatial structuring of biological communities in ecosystems that have intrinsic and general connectivity patterns, such as the universal dendritic structure of fluvial networks. These networks not only define dispersal of native species, but also represent corridors of biological invasions, making understanding network topology effects on invasion dynamics and subsequent diversity patterns of high interest. We studied amphipod community diversity and structure in the upper 27,882-km² drainage basin of the river Rhine in Central Europe, focusing on differences between native and non-native species. Overall, species richness increased along the network from headwaters to the outlet nodes. We found, however, contrasting patterns of native and non-native amphipod richness along the network, with headwater nodes representing refugia for native species and more downstream nodes being hotspots of biological invasions. Importantly, while species turnover (b-diversity) of native species increased with distance between nodes in the network, this was not the case for non-native species, indicating a much lower dispersal limitation of the latter. Finally, the overall amphipod community structure closely mirrored the topological modularity of the network, highlighting the network’s imprint on community structure. Our results underpin the importance of connectivity for community formation and the significance of rivers for biological invasions and suggest that empirically observed matches of diversity patterns in rivers predicted by null models are the long-term outcome of species invasions and species sorting.

The amphipod genus Niphargus (Amphipoda: Niphargidae Bousfield, 1977) is the most species-rich genus of freshwater amphipods in the World. Species of this genus, which live almost exclusively in subterranean water, offer an interesting model system for basic and applied biodiversity science. Their use, however, is often limited due to the hitherto unresolved taxonomy within the whole genus. As a comprehensive taxonomic revision of the currently >425 Niphargus species is too demanding, it has been suggested that the taxonomy of the genus could be advanced in smaller steps, by reviewing regional faunas, that would eventually integrate into a global revision. In this study, we provide such a revision of Niphargus in Switzerland. First, we molecularly delimited, morphologically diagnosed, and formally described two new species, namely Niphargus luchoffmanni sp. n. and Niphargus tonywhitteni sp. n. Second, we updated and revised a checklist of Niphargus in Switzerland with new findings, and prepared a list of reference sequences for routine molecular identification, available at BOLD and GenBank. All available specimens of 22 known species from the area were morphologically examined, and their morphological variation was compiled in a data file of DEscription Language for TAxonomy, which can be used for automated generation of dichotomous or interactive keys. The data file is freely available at the World Amphipoda Database. Together, the checklist, the library of reference sequences, the DELTA file, but also a list of hitherto unresolved aspects are an important step towards a complete revision of the genus within a well-defined and biogeographically interesting area in Central Europe.

Losses and gains in species diversity affect ecological stability^1–7 and the sustainability of ecosystem functions and services^8–13. Experiments and models have revealed positive, negative and no effects of diversity on individual components of stability, such as temporal variability, resistance and resilience^{2,3,6,11,12,14}. How these stability components covary remains poorly understood¹⁵. Similarly, the effects of diversity on overall ecosystem stability¹⁶, which is conceptually akin to ecosystem multifunctionality^17,18, remain unknown. Here we studied communities of aquatic ciliates to understand how temporal variability, resistance and overall ecosystem stability responded to diversity (that is, species richness) in a large experiment involving 690 micro-ecosystems sampled 19 times over 40 days, resulting in 12,939 samplings. Species richness increased temporal stability but decreased resistance to warming. Thus, two stability components covaried negatively along the diversity gradient. Previous biodiversity manipulation studies rarely reported such negative covariation despite general predictions of the negative effects of diversity on individual stability components³. Integrating our findings with the ecosystem multifunctionality concept revealed hump- and U-shaped effects of diversity on overall ecosystem stability. That is, biodiversity can increase overall ecosystem stability when biodiversity is low, and decrease it when biodiversity is high, or the opposite with a U-shaped relationship. The effects of diversity on ecosystem multifunctionality would also be hump- or U-shaped if diversity had positive effects on some functions and negative effects on others. Linking the ecosystem multifunctionality concept and ecosystem stability can transform the perceived effects of diversity on ecological stability and may help to translate this science into policy-relevant information.

Amphipods are keystone species in many freshwater ecosystems. Understanding their distribution and diversity is crucial to ensure and preserve freshwater ecosystem functioning, particularly in the northern hemisphere. For the European Alps information on amphipods has been relatively limited until recently. We describe a new, widely distributed amphipod species, Gammarus alpinus sp. nov., found across the Alps and analyse its distribution, biogeography as well as genetic and morphological differentiation. Until now, this species has been reported as Gammarus lacustris. Based on genetic and morphometric measurements, we show that G. alpinus is highly divergent from G. lacustris. The latter has a circumpolar distribution, but may be absent from the Alps. The observed occurrence pattern of G. alpinus might be explained by a Pliocene range expansion and vicariance due to climate warming following the Quaternary glaciation. Historical drainage divides suggest a single recolonization route from a distinct southern refugium. While G. lacustris is widely distributed and not endangered at a global scale, G. alpinus is endemic to the Alps and its habitat is negatively affected by eutrophication, non-native species and possibly climate change.

Groundwater belongs to the spatially most extensive, but least explored freshwater systems. On a global scale, the species richness of several subterranean invertebrate taxa parallels species richness found in surface waters, while on a local scale species richness hardly exceeds 20 species. This results in a high contribution of groundwater ecosystems to regional β- and γ-diversity, and to a smaller degree to α-diversity, and deserves focused attention. In general, more species are to be found in large cave systems. The second largest cave system in Europe is Hölloch in Switzerland. In this paper we revised the taxonomic, phylogenetic and ecological diversity of the amphipod community in the Hölloch cave system. While previous records listed five geographically widespread species of the genus Niphargus for this cave system, we could not confirm the presence of any of those species, but rather found three highly distinct species new to science. In this paper we describe Niphargus styx sp. nov., Niphargus murimali sp. nov., and Niphargus muotae sp. nov., and suggest that previous records from that cave were probably misidentificationsf. Although amphipod species richness in this cave system seems to be lower than previously thought in terms of absolute numbers, the cave retained its regional and international importance in terms of nature conservation for multiple reasons. First, all newly described species are probably endemic to this cave system. Second, they are phylogenetically distantly related and exhibit moderate to high phylogenetic diversity. Third, the species, as inferred from their functional morphology, are also ecologically highly divergent. Based on geographic distribution of their nearest relatives, we hypothesize that the cave system was most likely independently colonized from North, West and South and that the pre-adapted ancestors occupied different ecological niches within the system.

Background: Worldwide, natural communities are invaded by non-native species, with potentially devastating effects on the native communities. A large part of past research aimed at finding traits and characteristics of the invading species or the invaded community explaining observed invasions. Only recently, the focus shifted on the spatial patterns during invasions per se. Empirical data, however, are limited, as invasions are often unique incidences of a complex spatio-temporal process. In order to identify generalities of invasion patterns, we studied 13 naturally replicated tributary streams draining into Lake Constance, and studied the occurrence of native and non-native amphipods along linear transects from the stream outlets to the upstream headwater reaches.
Results: We found repeated spatial patterns of community composition and the occurrence of native and non-native amphipod species across two different years. Specifically, occurrence as well as abundance of two non-native amphipod species decreased from the stream outlets at the lake site towards upstream headwater reaches. Populations of the most common native amphipod species were largest at the uppermost headwater reaches. All populations of this native species, however, showed significant signals of recent genetic bottlenecks, irrespective of the stream position and occurrence of non-native species. Contrary to our expectations, this native species also showed no longitudinal genetic differentiation within individual tributaries as postulated for headwater versus outlet populations.
Conclusions: Our results indicate that invasions of river-systems may overall follow predictable patterns on the level of spatial distributions and community composition. However, effects of invading organisms on the genetic diversity and genetic structure of native populations observed at larger scales may not necessarily be directly reflected at the scale of smaller tributaries.

Chironomids generally dominate macroinvertebrate assemblages in alpine streams. This study compared chironomid assemblages inhabiting 51 non-glacial streams in six alpine catchments in the Swiss Alps in 2012. Total abundances ranged from 3 to ca. 1900 individuals per sample with 42 taxa identified overall. Common taxa differed among catchments and among streams within a catchment (5 taxa comprised > 95 % of the total abundances among streams). Taxon richness varied from 18 to 29 between catchments and between 2 and 16 between individual streams. Beta diversity (B_w) ranged from 1.27 to 3.03 between streams within catchments. The most common taxa included Diamesa spp. (mostly D. zernyi-cinerella group), Pseudodiamesa spp. (mostly P. branickii), Pseudokiefferiella parva, Corynoneura sp, Eukiefferiella spp., Orthocladius frigidus, Parametriocnemus stylatus, and Micropsectra spp. Nineteen taxa (45 %) were found at < 5 % of the streams and 8 taxa (19 %) were exclusive to one stream alone. These results indicate a highly diverse and spatially-structured chironomid assemblage with a prevalence towards rarity in these high elevation alpine catchments.

We investigated environmental characteristics and macroinvertebrate trophic structures of alpine streams in the glaciated Dinwoody catchment of the Wind River Range, Wyoming, USA. Study sites were above 3100 m a.s.l. and all above treeline except for one site representing a forested headwater stream. Physico-chemistry grouped sites by sub-basin and glacial influence. Seston levels were higher in the forested stream and a side-slope tributary (> 227 mg m^-3) than the other streams (< 100 mg m^–3). Periphyton levels were higher (> 2.7 mg m^–2) in streams in the Heap Steep sub-basin fed by a remnant hanging glacier (active ice field) than in the other streams (< 1.0 mg m^–2). Macroinvertebrate abundances (range = 140 – 2400 individuals per 5-min kick sample) and taxa richness were generally low (2–8 taxa at most sites), although being higher in lower-basin sites and the forested stream than at sites nearest the glaciers. Macroinvertebrate trophic structures were simple, consisting of primary consumers and predators, including two riparian spiders. Macroinvertebrate δ¹⁵N values ranged from ca. –2.54 to –5.41, whereas δ¹³C values ranged from –24.23 to –16.08. Individual taxa varied widely in δ¹³C among sites, suggesting carbon sources differed among sites. Carbon isotope ratios also increased from upper sites to lower sites in the same subbasin, suggesting a glacial carbon source at sites near the glacier to an instream carbon source at lower glacial sites lacking substantial riparian inputs. The upper glacial sites also had carbon isotope ratios similar to that of the forested headwater stream. These data suggest a highly heterogeneous spatial structure in alpine landscapes concomitant with the longitudinal development of glacial streams that is reflected in the trophic structure of local macroinvertebrates.

Amphipods are key organisms in many freshwater systems and contribute substantially to the diversity and functioning of macroinvertebrate communities. Furthermore, they are commonly used as bioindicators and for ecotoxicological tests. For many areas, however, diversity and distribution of amphipods is inadequately known, which limits their use in ecological and ecotoxicological studies and handicaps conservation initiatives. We studied the diversity and distribution of amphipods in Switzerland (Central Europe), covering four major drainage basins, an altitudinal gradient of>2,500 m, and various habitats (rivers, streams, lakes and groundwater). We provide the first provisional checklist and detailed information on the distribution and diversity of all amphipod species from Switzerland. In total, we found 29 amphipod species. This includes 16 native and 13 non-native species, one of the latter (Orchestia cavimana) reported here for the first time for Switzerland. The diversity is compared to neighboring countries. We specifically discuss species of the genus Niphargus, which are often receiving less attention. We also found evidence of an even higher level of hidden diversity, and the potential occurrence of further cryptic species. This diversity reflects the biogeographic past of Switzerland, and suggests that amphipods are ideally suited to address questions on endemism and adaptive radiations, post-glaciation re-colonization and invasion dynamics as well as biodiversity-ecosystem functioning relationships in aquatic systems.

Many glaciers on the globe have significantly receded during the last decades in response to climate change. Most studies in this context have focused on biotic assemblages in glacial-fed (kryal) streams, suggesting a high potential loss in macroinvertebrate diversity in kryal streams as glacier-flow contributions diminish. Alpine basins also comprise a large number of non-glaciated (krenal, rhithral) streams likely inhabited by macroinvertebrate assemblages different than those common to kryal streams. We examined macroinvertebrate assemblages in 41 non-glacial tributaries within 5 different glacial catchments in the Swiss Alps. The data were used to determine their contribution and role in the overall biodiversity of glacial catchments. We found an astonishing high diversity of macroinvertebrates in these tributaries with 64 taxa in 12 orders being identified overall and up to 24 different taxa found at individual sites. The abundance, taxonomic richness (α-diversity) and assemblage composition (β-diversity) differed highly between tributaries, suggesting their role as habitat for non-glacial specialists and as potential refugia for alpine species in general. The tributaries showed high individuality in physico-chemical factors as well as in taxonomic composition. Most of the taxa found could be defined as rare when comparing abundances, highlighting the importance of an analytical approach including all taxa. These results indicate that non-glacial tributaries contribute significantly to
the biodiversity of alpine running waters and may play an important role in sustaining alpine biodiversity under landscape transformation resulting from climate change.