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The importance of oscillations and deterministic chaos in natural biological systems has been discussed for several decades and was originally based on discrete-time population growth models (May 1974). Recently, all types of nonlinear dynamics were shown for experimental communities where several species interact. Yet, there are no data exhibiting the whole range of nonlinear dynamics for single-species systems without trophic interactions. Up until now, ecological experiments and models ignored the intracellular dimension, which includes multiple nonlinear processes even within one cell type. Here, we show that dynamics of single-species systems of protists in continuous experimental chemostat systems and corresponding continuous-time models reveal typical characteristics of nonlinear dynamics and even deterministic chaos, a very rare discovery. An automatic cell registration enabled a continuous and undisturbed analysis of dynamic behavior with a high temporal resolution. Our simple and general model considering the cell cycle exhibits a remarkable spectrum of dynamic behavior. Chaos-like dynamics were shown in continuous single-species populations in experimental and modeling data on the level of a single type of cells without any external forcing. This study demonstrates how complex processes occurring in single cells influence dynamics on the population level. Nonlinearity should be considered as an important phenomenon in cell biology and single-species dynamics and also, for the maintenance of high biodiversity in nature, a prerequisite for nature conservation.

Percolomonads (Heterolobosea) are aquatic heterotrophic flagellates frequently found in saline waters up to hypersaline environments. We isolated and cultivated seven strains of percolomonad flagellates from marine waters and sediments as well as from a hypersaline inland lake in the Atacama Desert. Morphological characterizations, comprising light and scanning electron microscopy, revealed only slight differences between the strains mainly limited to the cell shape, length of flagella, and length of the ventral feeding groove. Phylogenetic analyses of the 18S and 28S rDNA genes showed the formation of three fully supported clades within the Percolomonadida: the Percolomonadidae, the Barbeliidae fam. nov. and the Lulaidae fam. nov. We describe two new families (Barbeliidae fam. nov., Lulaidae fam. nov.), a new genus (Nonamonas gen. nov.), and five new species (Percolomonas adaptabilis sp. nov., Lula levis sp. nov., Barbelia pacifica sp. nov., Nonamonas montiensis gen. et sp. nov., Nonamonas santamariensis gen. et sp. nov.). Salinity experiments showed that P. adaptabilis sp. nov. from the Atlantic was better adapted to high salinities than all other investigated strains. Moreover, comparisons of our cultivation-based approach with environmental sequencing studies showed that P. adaptabilis sp. nov. seems to be globally distributed in marine surface waters while other species seem to be more locally restricted.

"Spumella"-like flagellates describes similar or even indiscernible colourless non-scaled chrysophytes which are important bacterivores common in different aquatic ecosystems. Recently, phylogenetic analyses revealed a high taxonomic diversity of these flagellates leading to the description of several new genera and species. Our present work on the functional group of pelagic bacterivorous chrysomonads from different water bodies resulted in an extended taxonomic analysis among chrysophytes unveiling yet undescribed genera and species pointing to the high hidden diversity of bacterivores in the pelagial of freshwaters. On the basis of phylogenetic analyses, we describe four new genera Atacamaspumella, Chlorospumella, Pseudapoikia, and Vivaspumella and a new species of the recently described genus Poteriospumella. Beside this, we redescribe the species Ochromonas vasocystis Doflein, 1923 to Poteriospumella vasocystis comb. nov. substantiated on the high sequence similarity with Poteriospumella lacustris Boenigk et Findenig and Poteriospumella maldiviensis nov. sp.

The heterotrophic nanoflagellate genus Cafeteria has been found to be ubiquitously distributed in the marine realm. We could isolate and cultivate ten strains morphologically similar to Cafeteria from various types of environment, including the deep sea, brackish waters and also meso- to hypersaline inland waters. Molecular analyses (18S rDNA, 28S rDNA) of newly isolated strains from the marine realm resulted in four more Cafeteria burkhardae strains from the deep North Atlantic Ocean and one new species (C. baltica sp. nov.) isolated from brackish waters of the Baltic Sea. Two strains isolated from the Atacama Desert belong to two new species (C. atacamiensis sp. nov. and C. paulosalfera sp. nov.), one other strain could not yet be assigned. Morphological characterizations of these strains obtained by high resolution microscopy revealed only small differences to already described species. However, molecular analyses showed a clear separation of the different Cafeteria species. We exposed several strains to different salt concentrations (2-150 PSU) to investigate their salinity tolerance. Only the marine strains of C. burkhardae were able to survive at salinities up to 150 PSU, indicating the possibility to inhabit a broader spectrum of habitats including hypersaline environments besides the deep sea with its high hydrostatic pressure.

Heterotrophic protists (unicellular eukaryotes) form a major link from bacteria and algae to higher trophic levels in the sunlit ocean. Their role on the deep seafloor, however, is only fragmentarily understood, despite their potential key function for global carbon cycling. Using the approach of combined DNA metabarcoding and cultivation-based surveys of 11 deep-sea regions, we show that protist communities, mostly overlooked in current deep-sea foodweb models, are highly specific, locally diverse and have little overlap to pelagic communities. Besides traditionally considered foraminiferans, tiny protists including diplonemids, kinetoplastids and ciliates were genetically highly diverse considerably exceeding the diversity of metazoans. Deep-sea protists, including many parasitic species, represent thus one of the most diverse biodiversity compartments of the Earth system, forming an essential link to metazoans., Alexandra Schoenle et al. use DNA metabarcoding and cultivation-based surveys of 11 regions in the Pacific and Atlantic Oceans encompassing the bathyal, abyssal, and hadal zones to identify the role of deep-sea heterotrophic protists relative to other trophic levels. Their results demonstrate that protistan species richness was locally specific and greater than that of metazoans and that measured in pelagic samples, and that deep-sea protist communities are relatively distinct from their pelagic counterparts.

Amphibian clutches are colonized by diverse but poorly studied communities of micro-organisms. One of the most noted ones is the unicellular green alga, Oophila amblystomatis, but the occurrence and role of other micro-organisms in the capsular chamber surrounding amphibian clutches have remained largely unstudied. Here, we undertook a multi-marker DNA metabarcoding study to characterize the community of algae and other micro-eukaryotes associated with agile frog (Rana dalmatina) clutches. Samplings were performed at three small ponds in Germany, from four substrates: water, sediment, tree leaves from the bottom of the pond, and R. dalmatina clutches. Sampling substrate strongly determined the community compositions of algae and other micro-eukaryotes. Therefore, as expected, the frog clutch-associated communities formed clearly distinct clusters. Clutch-associated communities in our study were structured by a plethora of not only green algae, but also diatoms and other ochrophytes. The most abundant operational taxonomic units (OTUs) in clutch samples were taxa from Chlamydomonas, Oophila, but also from Nitzschia and other ochrophytes. Sequences of Oophila “Clade B” were found exclusively in clutches. Based on additional phylogenetic analyses of 18S rDNA and of a matrix of 18 nuclear genes derived from transcriptomes, we confirmed in our samples the existence of two distinct clades of green algae assigned to Oophila in past studies. We hypothesize that “Clade B” algae correspond to the true Oophila, whereas “Clade A” algae are a series of Chlorococcum species that, along with other green algae, ochrophytes and protists, colonize amphibian clutches opportunistically and are often cultured from clutch samples due to their robust growth performance. The clutch-associated communities were subject to filtering by sampling location, suggesting that the taxa colonizing amphibian clutches can drastically differ depending on environmental conditions.

Land-use intensification is a major driver of biodiversity loss. However, understanding how different components of land use drive biodiversity loss requires the investigation of multiple trophic levels across spatial scales. Using data from 150 agricultural grasslands in central Europe, we assess the influence of multiple components of local- and landscape-level land use on more than 4,000 above- and belowground taxa, spanning 20 trophic groups. Plot-level land-use intensity is strongly and negatively associated with aboveground trophic groups, but positively or not associated with belowground trophic groups. Meanwhile, both above- and belowground trophic groups respond to landscape-level land use, but to different drivers: aboveground diversity of grasslands is promoted by diverse surrounding land-cover, while belowground diversity is positively related to a high permanent forest cover in the surrounding landscape. These results highlight a role of landscape-level land use in shaping belowground communities, and suggest that revised agroecosystem management strategies are needed to conserve whole-ecosystem biodiversity., Nature Communications, 12 (1), ISSN:2041-1723

New phagotrophic euglenoid species from marine surface waters and the deep sea were isolated and described by light and scanning electron microscopy and 18S rDNA sequencing: Keelungia nitschei, Petalomonas acorensis and Ploeotia costaversata. The morphological characteristics of Keelungia nitschei agree with Keelungia pulex besides the slightly truncated anterior front of the cell of our strain. Phylogenetic analysis indicated low sequence similarity between K. nitschei and K. pulex (87.3%). Ploeotia costaversata clustered within the Ploeotia costata clade with a sequence similarity of 96.1% to P. costata strain Tam. Ultrastructural characteristics of our strain revealed helically twisted strips towards both poles of the protoplast. 18S rDNA phylogenies showed that Petalomonas acorensis is related to the clade of Petalomonas cantuscygni/Scytomonas saepesedens with the highest sequence similarity of 81.2% to P. cantuscygni. Six pellicle strips are visible, while two of them reach only the middle of the cell and four (two longitudinal, two helically twisted) join at the posterior front of the cell. Pressure experiments showed that the deep-sea strain K. nitschei was better adapted to high hydrostatic pressures (up to 500 bar) at 4 °C than the two surface water strains. All three strains increased the database (18S rDNA) of the underrepresented group of phagotrophic euglenids.

Percolomonads are common freshwater, marine and hypersaline tetraflagellated organisms. Current phylogenetic analyses of eukaryotes comprise only two species of this underinvestigated family. Here, we studied the morphology, salinity tolerance and 18S rDNA gene-based phylogeny of seven percolomonad cultures. We describe three new genera and five novel species of Percolomonadida based on phylogenetic distances and morphological characteristics: Barbelia atlantica, B. abyssalis, Lula jakobsenorum, Nakurumonas serrata and Percolomonas doradorae. The new species show features typical for percolomonads, one long flagellum for skidding, three shorter flagella of equal length and a ventral feeding groove. The new species comprise organisms living in marine and athalassic hypersaline waters with salinity ranging from 10 to 150 PSU. Based on these novel taxa, the taxonomy and phylogeny of Percolatea was extended and further resolved.

Protists are the most diverse eukaryotes on our planet and metabarcoding has revealed an enormous diversity even from deep-sea environments. A range of different species has also been isolated from the deep sea and some have proven able to survive and even grow under deep-sea conditions. However, little is known about how the community structure of benthic protists changes from sublittoral down to abyssal depths. This is especially important regarding island and seamount communities which are surrounded by deep-sea areas potentially isolating them. Using a combination of live-counting and cultivation techniques, we investigated the abundance and taxonomic composition of benthic protist communities in sediments from sublittoral to abyssal depths around three islands and two seamounts of the Azores’ archipelago in the North Atlantic. Protist abundance decreased significantly and community composition changed with increasing depth. While some species were found at all depths, others were only detected in sublittoral or lower bathyal depths, indicating that some benthic taxa are limited in their distribution to a certain depth, whereas others are also present at the deep-sea floor. The proportion of unidentified specimens increased with depths pointing towards a high number of so far undetected species in the deep-sea realm.

Recent studies suggested that the diversity of microbial eukaryotes in hypersaline environments is widely underestimated. Placidids are a group of heterotrophic stramenopile flagellates that are frequently found in these environments, but up to now only very few species were isolated and fully described, mostly from marine or brackish water sites. In this study, we extend the known diversity of Placididea by three new genera (Allegra, Haloplacidia, and Placilonga) compromising nine new species, isolated from athalassic, mostly hypersaline environments (Allegra dunaii, Allegra atacamiensis, Allegra hypersalina, Haloplacidia cosmopolita, Suigetsumonas keniensis) and marine waters (Placilonga atlantica, Placidia azorensis, Placidia abyssalis, Wobblia pacifica) including a description of their morphology and molecular phylogeny. In total, 36 strains were comparatively analysed. Studies from athalassic waters revealed an especially high number of different genotypes. A multigene analysis based on a ten genes dataset revealed a clear separation into marine, athalassic and brackish water clades. Several representatives were found to cope with hypersaline conditions from 20 to 250 PSU, even up to 284 PSU, suggesting that they may form a halotolerant group.

Locomotion is considered to be the main form of expression of ciliate behaviour regarding their overall life activity. But how ciliates behave under deep-sea conditions is still unclear. Data on the occurrence of ciliates in the deep sea are scarce and mostly based on molecular studies. We isolated three different ciliates,Aristerostomasp.,Euplotes dominicanusandPseudocohnilembus persalinusfrom two stations located in abyssal depths of the North Atlantic Ocean (≥ 4000 m; 15° 55.89′ N, 68° 53.34′ W; 23° 33.23′ N, 48° 5.04′ W) during the deep-sea expedition with the research vessel R/V Meteor (Cruise M139, 08.07.–08.08.2017). We observed their behaviour directly under high hydrostatic pressures up to 500 bar. The three ciliate species behaved normally up to a pressure of 200 bar, but showed disturbances of the normal behaviour at higher pressures. For all three isolated deep-sea ciliates, additional long-term survival experiments were carried out for 6 days at 200, 350 and 430 bar. Several specimens showed an ability to survive the entire experimental time interval at the highest pressure and to recover from pressure release (returning to their normal movement) indicating their barotolerance. Our results suggest that ciliates are active in the deep sea even in regions deeper than 2000 m and might be an important part of the deep-sea microbial food web.

The deep sea (>200 m depth) encompasses >95% of the world’s ocean volume and represents the largest and least explored biome on Earth (

International audience; The dark ocean and the underlying deep seafloor together represent the largest environment on this planet, comprising about 80% of the oceanic volume and covering more than two- thirds of the Earth’s surface, as well as hosting a major part of the total biosphere. Emerging evidence suggests that these vast pelagic and benthic habitats play a major role in ocean biogeochemistry and represent an ”untapped reservoir“ of high genetic and metabolic microbial diversity. Due to its huge volume, the water column of the dark ocean is the largest reservoir of organic carbon in the biosphere and likely plays a major role in the global carbon budget. The dark ocean and the seafloor beneath it are also home to a largely enigmatic food web comprising little-known and sometimes spectacular organisms, mainly prokaryotes and protists. This review considers the globally important role of pelagic and benthic protists across all protistan size classes in the deep-sea realm, with a focus on their taxonomy, diversity, and physiological properties, including their role in deep microbial food webs. We argue that, given the important contribution that protists must make to deep-sea biodiversity and ecosystem processes, they should not be overlooked in biological studies of the deep ocean.

1.Rapid growth of the world's human population has increased pressure on landscapes to deliver high levels of multiple ecosystem services, including food and fibre production, carbon storage, biodiversity conservation and recreation. However, we currently lack general principles describing how to achieve this landscape multifunctionality. 2.We combine theoretical simulations and empirical data on 14 ecosystem services measured across 150 grasslands in three German regions. In doing so, we investigate the circumstances under which spatial heterogeneity in a driver of ecosystem functioning (an ‘ecosystem‐driver,’ e.g. the presence of keystone species, land‐use intensification or habitat types) increases landscape‐level ecosystem multifunctionality. 3.Simulations based on theoretical data demonstrated that relationships between heterogeneity and landscape multifunctionality are highly variable and can range from non‐significant to strongly positive. Despite this variability, we could identify criteria under which heterogeneity‐landscape multifunctionality relationships were most strongly positive: this happened when multiple ecosystem services responded contrastingly (both positively and negatively) to an ecosystem‐driver. 4.These findings were confirmed using empirical data, which showed that heterogeneity in land‐use intensity promoted landscape multifunctionality in cases where functions with both positive (e.g. plant biomass) and negative (e.g. flower cover) responses to land use intensification were included. For example, the simultaneous provisioning of ecosystem functions related to forage production (generally profiting from land‐use intensification), biodiversity conservation and recreation (generally decreasing with land‐use intensification) was highest in landscapes consisting of sites varying in land‐use intensity. 5.Synthesis and applications. Our findings show that there are general principles governing landscape multifunctionality. A knowledge of these principles may support land management decisions. For example, knowledge of relationships between ecosystem services and their drivers, such as land use type, can help estimate the consequences of increasing or decreasing heterogeneity for landscape‐level ecosystem service supply, although interactions between landscape units (e.g. the movement of pollinators) must also be considered.

By measuring the change in soil protist communities, the effect of human land use on grasslands can be monitored to promote sustainable ecosystem functioning. Protists form the active link in the rhizosphere between the plant roots and higher trophic organisms; however, only few morphological species and their ecological values have yet been described in this context. To investigate the communicability between morphological and molecular databases used in the molecular barcoding of protists and in the biomonitoring of grassland soil, the present high-throughput sequencing (HTS) study (N=150) covered the area of central Europe (mesoscale) known to be well studied for ciliated protists. HTS delivered 2,404 unique reads identifying taxa in all major ciliophoran classes but exact reference matches were few. The study identified clear discrepancies between databases for well-studied taxa, where molecular databases contained multiple gene variants for single morphospecies of dominant taxa. Gene variants presented own biogeography - the eukaryotic microdiversity along gradients (e.g., land-use intensity, soil water). It is possible that many of the so called novel phylogenetic lineages and hidden diversity pointed out in environmental surveys could be evidence for the severe lack of molecular data for already known and morphologically described species, present in morphological databases.

Ornamental species of the aquarium pet trade contribute more and more to the introduction of previously exotic species in Europe. Among these, freshwater crustaceans play an increasingly larger role. Especially in thermally altered streams they may become invasive; however, field data on population ecology of invasive crustaceans are very rare. The aim of this study was to analyze the population dynamics of the invasive freshwater shrimp Neocaridina davidi (indigenous to China and other Asian countries) in a year-round warm-water habitat in the Gillbach stream (North Rhine-Westphalia, Germany), where this species forms a stable population. The species reached abundances between 0.23–1.39 individuals per m². N. davidi is highly productive and detectable in the whole Gillbach stream and in the lower Erft River, too. Egg ratio and cohort development were documented for a period of 27 months. Among adults, between 45 and 59% were females (average 50.5%). Life span was about 12 months in the field and up to 21 months under laboratory conditions. Development times under field and laboratory conditions were similar, ovigerous females appeared about 7 months after hatching. The population in the Gillbach stream was multivoltine with about two generations per year. Only 2–3% of eggs reached adulthood in the field most probably due to intensive predation by indigenous and invasive fish species. One specimen could already be detected in the Lower River Rhine. N. davidi is omnivorous and well adapted to different freshwater habitats. The risk of further spreading to other European river systems is obvious.

Freshwater sponges play a major role in freshwater ecological system as important filter-feeding organisms and bioindicators. There are only few data about their ecological diversity and population genetic structure available, though a deeper knowledge is needed to propose proper conservation and effective management. The aim of this study was to assess data on distribution patterns of freshwater sponges to study the connectivity of genotypes of Ephydatia fluviatilis in a river system. We sampled specimens from River-Sieg system (River Agger and River Sieg, Germany). We hypothesized that strong anthropogenic influence would cause a uniform distribution of population structures. The genetic structure of E. fluviatilis populations was analysed with a set of eleven microsatellite loci from seven locations in River-Sieg system. Besides of E. fluviatilis, three other species co-occurred (Ephydatia mulleri, Spongilla lacustris, Eunapius fragilis). In contrast to our hypothesis, we observed an overall correlation between genetic and geographic distances among populations of this sessile species, which follows a clear isolation-by-distance pattern. A significant microsatellite polymorphism and high levels of genetic divergence between populations (FST) in upstream reaches were present. These results will provide important information for conservation management of populations with limited dispersal ability in connected river systems.

Although the abyssal seafloor represents the most common benthic environment on Earth, eukaryotic microbial life at abyssal depths is still an uncharted territory. This is in striking contrast to their potential importance regarding the material flux and bacteria consumption in the deep sea. Flagellate genotypes determined from sedimentary DNA deep‐sea samples might originate from vital deep‐sea populations or from cysts of organisms sedimented down from surface waters. The latter one may have never been active under deep‐sea conditions. We wanted to analyze the principal ability of cultivable heterotrophic flagellates of different phylogenetic groups (choanoflagellates, ancyromonads, euglenids, kinetoplastids, bicosoecids, chrysomonads, and cercozoans) to survive exposure to high hydrostatic pressure (up to 670 bar). We summarized our own studies and the few available data from literature on pressure tolerances of flagellates isolated from different marine habitats. Our results demonstrated that many different flagellate species isolated from the surface waters and deep-sea sediments survived drastic changes in hydrostatic pressure. Barophilic behavior was also recorded for several species isolated from the deep sea indicating their possible genetic adaptation to high pressures. This is in accordance with records of heterotrophic flagellates present in environmental DNA surveys based on clone libraries established for deep‐sea environments.

Genomic data for less than one quarter of ∼1.8 million named species on earth exist in public databases like GenBank. Little information exists on the estimated one million small sized (1–100 μm) heterotrophic nanoflagellates and ciliates and their taxa-area relationship. We analyzed environmental DNA from 150 geo-referenced grassland plots representing topographical and land-use ranges typical for Central Europe. High through-put barcoding allowed the identification of operational taxonomic units (OTUs) at species level, with high pairwise identity to reference sequences (≥99.7%), but also the identification of sequences at the genus (≥97%) and class (≥80%) taxonomic level. Species richness analyses revealed, on average, 100 genus level OTUs (332 unique individual read (UIR) and 56 class level OTUs per gram of soil sample in the mesoscale (1–1 000 km). Database shortfalls were highlighted by increased uncertain taxonomic lineages at lower resolution (≥80% sequence identity). No single barcode occurred ubiquitously across all sites. Taxa-area relationships indicated that OTUs spread over the entire mesoscale were more similar than in the local scale and increased land-use (fertilization, mowing and grazing) promoted taxa-area separation. Only a small fraction of sequences strictly matched reference library sequences, suggesting a large protistan “dark matter” in soil which warrants further research.

Microbial eukaryotes play an important role in biogeochemical cycles not only in productive surface waters but also in the deep sea. Recent studies based on metagenomics report deep-sea protistan assemblages totally different from continental slopes and shelf waters. To give an overview about the ciliate fauna recorded from the deep sea we summarized the available information on ciliate occurrence in the deep sea. Our literature review revealed that representatives of the major phylogenetic groups of ciliates were recorded from the deep sea (> 1000 m depth): Karyorelictea, Heterotrichea, Spirotrichea (Protohypotrichia, Euplotia, Oligotrichia, Choreotrichia, Hypotrichia), Armophorea (Armophorida), Litostomatea (Haptoria), Conthreep (Phyllopharyngea incl. Cyrtophoria, Chonotrichia, Suctoria; Nassophorea incl. Microthoracida, Synhymeniida, Nassulida; Colpodea incl. Bursariomorphida, Cyrtolophosidida; Prostomatea; Plagiopylea incl. Plagiopylida, Odontostomatida; Oligohymenophorea incl. Peniculia, Scuticociliatia, Hymenostomatia, Apostomatia, Peritrichia, Astomatia). Species occurring in both habitats, deep sea and shallow water, are rarely found to our knowledge to date. This indicates a high deep-sea specific ciliate fauna. Our own studies of similar genotypes (SSU rDNA and cox1 gene) revealed that two small scuticociliate species ( Pseudocohnilembus persalinus and Uronema sp. ) could be isolated from surface as well as deep waters (2687 m, 5276 m, 5719 m) of the Pacific. The adaptation to deep-sea conditions was investigated by exposing the ciliate isolates directly or stepwise to different hydrostatic pressures ranging from 1 to 550 atm at temperatures of 2 °C and 13 °C. Although the results indicated no general barophilic behavior, all four isolated strains survived the highest established pressure. A better survival at 550 atm could be observed for the lower temperature. Among microbial eukaryotes, ciliates should be considered as a diverse and potentially important component of deep-sea microeukaryote communities.

Spatial distribution patterns of protists in the deep sea are not yet understood and due to the remoteness of the deep-sea biome this is a challenging task. Here, we investigated the structure and dispersal of protist communities in abyssal sediments at three sites in the Caribbean Sea (15°53.21′N/68°55.73′W), the northwestern Atlantic (23°33.23′N/48°05.04′W) and the central Atlantic Ocean (10°20.38′N/36°57.76′W), focusing especially on small horizontal (~1 m) and vertical (1 cm) spatial scales. High-throughput sequencing of the V9 SSU rDNA, revealed diverse protist communities dominated by diplonemids and dinoflagellates. Only 12% of OTUs were shared between the three deep-sea regions (4000–5134 m deep), indicating patterns of protistan distribution at large spatial scales in deep-sea sediments. The analyses of sediment samples taken from cores of the same Multi-Corer deployment, separated by ~1 m, showed differences in the community structure, indicating a distinct microbial eukaryotic diversity even at a local horizontal scale. At a vertical scale, in different sediment depth layers (upper 2 mm layer, 1 cm, 2 cm, 3 cm) of the same core, only 4–11% of OTUs were shared. We could show, that already at small spatial scales protist communities varied strongly, indicating an underestimated complexity of deep-sea microbial food webs.

Detailed knowledge of the significance of shrimps in freshwater food webs is very limited. However, determination of the current potential invasion of shrimps into European freshwater systems requires information on their ecology and feeding behaviour. Atyaephyra desmarestii has established stable populations in Western and Central Europe, while the ornamental species Neocaridina davidi was released in 2009 into a small tributary of the Erft River (North Rhine Westphalia, Germany), where it has thrived. Both species use leaf-litter as a significant food source. In this study, we assessed a reproducible method to compare the preferences of this two shrimp species for decaying leaves of four different species of deciduous tree: alder (Alnus glutinosa), Italian poplar (Populus x canadensis), pedunculate oak (Quercus robur) and goat willow (Salix caprea). We also determined the relevance of A. desmarestii and N. davidi in leaf-litter breakdown. Adults of both species showed a significant preference for leaves of alder and Italian poplar, whereas juvenile individuals did not favour any particular leaf species. A. desmarestii and N. davidi adults exhibited higher night-time than daytime activity. Diurnal consumption rates were determined for N. davidi. It consumed 51.0% leaf litter dry weight per body dry weight per day. Alnus and Salix leaves (including biofilm) made up the majority of the diet of Neocaridina, followed by Populus and Quercus leaves. Our results demonstrate the distinct relevance of leaf-litter in the diet of freshwater shrimps, and their role in leaf-litter breakdown. While the invasion potential of A. desmarestii seems to be relatively low, at least for now, N. davidi has thus far been a very successful invader. This is supported by its high feeding rates on leaf litter of the regional vegetation. Since there is no indigenous shrimp species in the study area, the potential implications of the invasion process merit further investigation.

Metazoans normally avoid anaerobic environments, at least when they are combined with toxic stress due to hydrogen sulfide. In Lake Speldrop, a small but deep gravel pit lake at the Lower Rhine, Daphnia galeata was found not only to dominate the zooplankton community, but was also regularly found in anoxic and even sulfidic layers during summer. We conducted field experiments with a newly developed “Zooplankton In-situ Incubator”, simulating vertical migrations of D. galeata. When daphnids were exposed to sulfidic conditions, mortality increased with exposure time, revealing LT50-values between 129 and 42 min in relation to increasing concentrations of sulfide. Additionally, those experimental individuals originating from 12.5 m depth showed significantly higher mortality rates than those from 7.5 m depth. Further migration experiments showed that an interruption period of sulfidic exposition in less stressful environments reduced mortality rates significantly. Daphnids found in the hypolimnion belonged partly to moribund parts of the population; however, the majority of daphnids showed regular mowing activity and was able to withstand sulfidic conditions in the hypolimnion for a limited time. It is so far unclear what maybe the major ultimate factor for this type of short term migratory behavior, to seek for shelter or to use high amounts of sulfur bacteria as an alternative food resource.

We investigate the appearance of chaos in a microbial 3-species model motivated by a potentially chaotic real world system (as characterized by positive Lyapunov exponents (Becks et al., Nature 435, 2005). This is the first quantitative model that simulates characteristic population dynamics in the system. A striking feature of the experiment was three consecutive regimes of limit cycles, chaotic dynamics and a fixed point. Our model reproduces this pattern. Numerical simulations of the system reveal the presence of a chaotic attractor in the intermediate parameter window between two regimes of periodic coexistence (stable limit cycles). In particular, this intermediate structure can be explained by competition between the two distinct periodic dynamics. It provides the basis for stable coexistence of all three species: environmental perturbations may result in huge fluctuations in species abundances, however, the system at large tolerates those perturbations in the sense that the population abundances quickly fall back onto the chaotic attractor manifold and the system remains. This mechanism explains how chaos helps the system to persist and stabilize against migration. In discrete populations, fluctuations can push the system towards extinction of one or more species. The chaotic attractor protects the system and extinction times scale exponentially with system size in the same way as with limit cycles or in a stable situation.

Paramonosiga coloniensis sp. nov., Salpingoeca amphoriscia sp. nov., S. fluviatilis sp. nov. and S. loutrophoria sp. nov. are frequently found craspedid species in the River Rhine which have not yet been described, despite their high abundance. All new species are characterized based on a distinct morphology which is different from all up to now described species and on a molecular level based on transcriptome data. In addition, we give extended redescriptions of S. amphoridium and S. angulosa, based on SSU and LSU rDNA data and morphology. The six-gene phylogenetic analyses place all species into freshwater clades of the craspedids. The separation of the freshwater and marine clades of this group is becoming more distinct with every craspedid sequence added. The River Rhine is one of the largest rivers in Europe but its protist biodiversity is fairly undescribed, especially regarding choanoflagellates. We conclude that the biodiversity of craspedid choanoflagellates is broadly underestimated.

A new Euplotes species, isolated from abyssal depths (>4000 m) of the North Atlantic Ocean, was described based on morphology, ciliary pattern and molecular data. Euplotes dominicanus sp. n. is characterized by a small body size (29–40 × 17–27 μm in vivo), 18–22 adoral membranelles, 10 frontoventral, five transverse and two left marginal cirri and one caudal cirrus, five or six dorsolateral kineties with 7–9 dikinetids in mid-dorsolateral kinety (DK3), and dorsal silverline system of the double-eurystomus type. Phylogenetic analyses inferred from 18S rRNA sequences show that Euplotes dominicanus sp. n. is most closely related to E. curdsi, with a sequence similarity of 97.6 %. Euplotes dominicanus sp. n. was able to survive hydrostatic pressures up to 500 bar indicating its barotolerance. Metabarcoding data demonstrate the presence of E. dominicanus sp. n. in sediments of several deep-sea basins.

In general, acanthoecid choanoflagellates have been described to occur exclusively in brackish water to marine habitats. Only recently, two studies have proven their existence in inland waters. One of them has shown, that an acanthoecid species from a small lake (near Apia on the island of Upolu, Samoa) is strictly freshwater adapted, not able to tolerate even brackish water. In this study, we present the first euryoecious acanthoecid species, able to live and reproduce in freshwater as well as under hypersaline conditions. The new species, Enibas tolerabilis gen. et sp. nov. was isolated in 2017 from the Salar de Ascotan in the Altiplano at 3750 m a. s. l., Northern Chile. The salinity at the time of sampling was 6 PSU. A series of autecological experiments have revealed a salinity tolerance from freshwater up to 70 PSU. In our phylogenetic analysis, E. tolerabilis gen. et sp. nov. clustered within the family of Acanthoecidae, forming a well-supported sister clade together with two other, environmental choanoflagellate sequences. We erected a new genus, Enibas gen. nov. and described the morphology, molecular biology and autecology for E. tolerabilis gen. et sp. nov. which has a stephanoecid-like lorica morphology. We emphasize that the definition of the genus Stephanoeca, being polyphyletic, is in urgent need of revision as we showed that this morphology is present in both acanthoecid families.

Many experiments have shown that loss of biodiversity reduces the capacity of ecosystems to provide the multiple services on which humans depend. However, experiments necessarily simplify the complexity of natural ecosystems and will normally control for other important drivers of ecosystem functioning, such as the environment or land use. In addition, existing studies typically focus on the diversity of single trophic groups, neglecting the fact that biodiversity loss occurs across many taxa and that the functional effects of any trophic group may depend on the abundance and diversity of others. Here we report analysis of the relationships between the species richness and abundance of nine trophic groups, including 4,600 above- and below-ground taxa, and 14 ecosystem services and functions and with their simultaneous provision (or multifunctionality) in 150 grasslands. We show that high species richness in multiple trophic groups (multitrophic richness) had stronger positive effects on ecosystem services than richness in any individual trophic group; this includes plant species richness, the most widely used measure of biodiversity. On average, three trophic groups influenced each ecosystem service, with each trophic group influencing at least one service. Multitrophic richness was particularly beneficial for 'regulating' and 'cultural' services, and for multifunctionality, whereas a change in the total abundance of species or biomass in multiple trophic groups (the multitrophic abundance) positively affected supporting services. Multitrophic richness and abundance drove ecosystem functioning as strongly as abiotic conditions and land-use intensity, extending previous experimental results to real-world ecosystems. Primary producers, herbivorous insects and microbial decomposers seem to be particularly important drivers of ecosystem functioning, as shown by the strong and frequent positive associations of their richness or abundance with multiple ecosystem services. Our results show that multitrophic richness and abundance support ecosystem functioning, and demonstrate that a focus on single groups has led to researchers to greatly underestimate the functional importance of biodiversity.

Freshwater snails are widely distributed in running water systems. Most of them are feeding on biofilms attached to substrata. We used gastropods with different morphology and potentially different traits to analyse the effect of water currents and substratum roughness on the interactions between grazers and biofilms. The gastropods were exposed to naturally grown biofilms from the River Rhine that differed in age and in their artificial substratum roughness (mimicked by abrasive paper). The experiments were conducted in endless channels with paddle wheels where the current velocity could be set steplessly. The freshwater pulmonates Ancylus fluviatilis and Physella acuta as well as the freshwater prosobranchs Bithynia tentaculata and Potamopyrgus antipodarum were used as model organisms in this investigation. The time the snails remained on the test area related to the substratum roughness, and current velocity was used to compare the performance of the four snail species on different aged biofilms. In one experiment, the locomotive activity of snails was traced. Gastropods left substrata without biofilms and did not show different responses to biofilms of diverse ages with significant differences in their biochemical contents. Grazing by snails altered the biofilms in their chlorophyll- a content, ash-free dry mass, and ash mass. A current velocity above 0.12 m s −1 drove B. tentaculata , P. acuta , and P. antipodarum off the test areas, only A. fluviatilis remained unaffected by the applied current velocities (up to 0.23 m s −1 ). The substratum roughness had no direct influence on the retention time of snails and their locomotive activity on the substrata, whereas the chlorophyll- a content of the biofilm strongly influenced snail activity. Substratum roughness had only an indirect influence on the behaviour of snails contrary to studies on marine snail species.

Unicellular eukaryotes, also called protists, are potentially fast evolving organisms. The small size of protists, their fast reproduction rate and ability to form cysts as well as their adaptability to extreme conditions allow them to associate to endemic animals, plants, saline lakes and soil even in extremely arid systems. These properties make unicellular eukaryotes ideal model organisms to combine studies on evolutionary processes of very different groups of organisms and across very different time scales comprising even geological ones. The hyperarid Atacama Desert offers a study area unique on Earth, where a predominantly arid climate was present for millions of years. A comprehensive analysis of the diversity of unicellular eukaryotes in different habitats (endemic desert plant phyllosphere, the gut of endemic darkling beetles, isolated hypersaline waters) revealed a dataset distinct and divergent from other regions on Earth. We used standard isolation and cultivation protocols to elucidate divergence patterns for a variety of very different and independent taxonomic groups of protists such as gregarines and ciliates among alveolates, placidids among stramenopiles and choanoflagellates among opisthokonts. The ability to rapidly adapt to extreme environments, which enhance a fast divergence rate at high UV radiation, has only been reported for prokaryotes up to now. The establishment of arid to hyperarid conditions in the Atacama Desert about 20 Ma ago has obviously led to an isolation of protist populations followed by a radiation of species. There are only a few regions on Earth with similar extreme salinity conditions reducing the chance of an exchange between protist populations. Divergence patterns in unicellular eukaryotes in very different phylogenetic groups independently mirror the effect of geological evolution and climate variability during the Neogene.

Cafeteria is one of the most common and ecologically significant genera of heterotrophic nanoflagellates in marine plankton. We could isolate and cultivate 29 strains morphologically similar to Cafeteria obtained from surface waters and the deep sea all over the world's ocean. Morphological characterization obtained by high resolution microscopy revealed only small differences between the strains. Sequencing the type material of the type species C. roenbergensis (CCAP 1900/1) and molecular analyses (18S rDNA, 28S rDNA) of newly isolated strains resulted in a revision and separation of the Cafeteriaceae into two known species (C. roenbergensis, C. mylnikovii) and six new species (C. maldiviensis, C. biegae, C. loberiensis, C. chilensis, C. graefeae, C. burkhardae). Many deposited Cafeteria sequences at GenBank and most of our own sequences clustered within one clade (C. burkhardae) with a p-distance of 5% to strain CCAP 1900/1. Only C. maldiviensis clustered together with the type species C. roenbergensis. While C. burkhardae seems to have a cosmopolitan distribution, the distribution of the other species seems to be more restricted. A strain from the Angola Basin had a p-distance of 10% to Cafeteria species and clustered separately within the Anoecales requiring the erection of a new genus, Bilabrum gen. nov., with B. latius sp. nov. as type species.

With this study we aim to extend the knowledge on the biogeography of craspedid choanoflagellates with additional data from extreme environments. Up to now, very little is known about choanoflagellates from extreme saline environments, as most studies have focused on marine and freshwater habitats. Though previously investigated high saline ice biota communities have indicated a possible adaptation to environments with high salt concentrations. Hypersaline endorheic basins, so-called salt flats or salares from the Atacama Desert in Northern Chile provide an intense environment regarding fluctuating and extreme salinities, which allow for studies on evolutionary adaptations of protists to hypersaline conditions. This study focused on choanoflagellate species isolated from different salt flats, their morphological characteristics using light and electron microscopy, molecular marker genes (SSU and LSU rDNA) and their salinity tolerance. Here, we described four new craspedid choanoflagellate species, highly adapted to the hypersaline environment of the Atacama Desert. This study extends our knowledge on choanoflagellate phylogeny and ecology and can become the basis for further molecular studies to understand the mechanisms of adaptations. Additionally, we emphasize the need of adding additional data such as autecological characteristics to amend species definitions, which is only possible from cultivated strains. This data would support the use of molecular data originating from metagenomic analyses also in an ecological context.

Biofilms, the complex communities of microbiota that live in association with aquatic interfaces, are considered to be hotspots of microbial life in many aquatic ecosystems. Although the importance of attached algae and bacteria is widely recognized, the role of the highly abundant biofilm-dwelling micrograzers (i.e., heterotrophic protists and small metazoans) is poorly understood. Studies often highlight the resistance of bacterial biofilms to grazing within the microbial food web and therefore argue that the micrograzers have a modulating role (i.e., have effects on biofilm phenotype) rather than a direct trophic role within biofilms. In the present review, we show that this view comes too short, and we establish a conceptual framework of biofilm food webs consisting of three major elements. (1) Energy pathways and subsidization from plankton. As inhabitants of interfaces, biofilm-dwelling grazers potentially access both planktonic organisms and surface-associated organisms. They can play an important role in importing planktonic production into the biofilm food web and thus in the coupling of the planktonic and benthic food webs. Nevertheless, specialized grazers are also able to utilize significant amounts of autochthonous biofilm production. (2) Horizontal complexity of the basal food web. While bacteria and algae within biofilms are edible in general, food quality and accessibility of both bacteria and algae can differ considerably between different prey phenotypes occurring during biofilm formation with respect to morphology, chemical defense, and nutrient stoichiometry. Instead of considering bacteria and algae within biofilms to be generally resistant to feeding by micrograzers, we suggest considering a horizontal food-quality axis to be at the base of biofilm food webs. This food quality gradient is probably associated with increasing costs for the micrograzers. (3) Vertical food web complexity and food chain length. In addition to the consumption of bacteria and algae, many predatory micrograzers exist within biofilm food webs. With the help of video microscopy, we were able to demonstrate the existence of a complex food web with several trophic levels within biofilms. Our conceptual framework should assist in integrating food web concepts and processes into whole-biofilm budgets and in understanding food-web-related interactions within biofilms.

Quantitative aspects of the life cycle of the freshwater shrimpAtyaephyra desmarestiiwere used to investigate its life span at the edge of its distribution area in Central Europe (Upper Rhine) for a period of three years. We estimated the proportion of the population surviving a second hibernation period. The appearance of ovigerous females in late spring was followed by a strong decline in the number of adults, the appearance of a high percentage of juveniles in late summer (0.58 ind./m2), and their subsequent disappearance from the shore in winter. A small semivoltine cohort survived a second winter. The average life span was 12-15 (max. 29) months under laboratory conditions. In molecular studies (COX I), the Upper Rhine and Southern France populations clustered within the North Africa/West-Central Europe species clade. Due to its low productivity, this invasive species is unlikely to cause management problems.

One of the most prominent manifestations of the ongoing climate warming is the retreat of glaciers and ice sheets around the world. Retreating glaciers result in the formation of new ponds and lakes, which are available for colonization. The gradual appearance of these new habitat patches allows us to determine to what extent the composition of asexual Daphnia (water flea) populations is affected by environmental drivers vs. dispersal limitation. Here, we used a landscape genetics approach to assess the processes structuring the clonal composition of species in the D. pulex species complex that have colonized periglacial habitats created by ice-sheet retreat in western Greenland. We analysed 61 populations from a young ( 150 years) of lakes and ponds. We identified 42 asexual clones that varied widely in spatial distribution. Beta-diversity was higher among older than among younger systems. Lineage sorting by the environment explained 14% of the variation in clonal composition whereas the pure effect of geographical distance was very small and statistically insignificant (R-adj(2) = 0.010, P = 0.085). Dispersal limitation did not seem important, even among young habitat patches. The observation of several tens of clones colonizing the area combined with environmentally driven clonal composition of populations illustrates that population assembly of asexual species in the Arctic is structured by environmental gradients reflecting differences in the ecology of clones.

The trophic interactions between viruses, bacteria and protists play a crucial role in structuring microbial communities and regulating nutrient and organic matter flux. Here, we show that the impact on viral density by heterotrophic flagellates is related to their feeding behaviour (feeding on sedimented particles - Thaumatomonas coloniensis, filter feeding of suspended particles - Salpingoeca sp., and actively searching raptorial feeding - Goniomonas truncata). Phage MS2 was co-incubated with flagellates and the natural bacterial and viral community originating from the same groundwater habitats where the flagellates were isolated. Three complementary assays, i.e. flow cytometry, qPCR and plaque assay, were used for enumeration of total viruses, total MS2 phages, and free and infectious MS2, respectively, to provide insights into the grazing mechanisms of the flagellates on viruses. Phage MS2 was actively removed by the suspension feeders T. coloniensis and Salpingoeca sp. in contrast with the actively raptoriale grazer G. truncata. The decline of viral titre was demonstrated to be caused by ingestion rather than random absorption by both qPCR and locating protein fluorescently labelled MS2 inside the flagellates. Further, we indicate that phages can be used as a minor carbon source for flagellates. Collectively, these data demonstrate that eliminating viruses can be an important function of protists in microbial food webs, carbon cycling and potentially water quality control.