Tidal and circadian patterns of European eel during their spawning migration in the North Sea and the English Channel.

Technological advances in tracking methods enable the mapping of anguillid eel migration routes from continental habitats to their spawning sites in the ocean. However, the behaviour and orientation abilities of anguillids are still poorly understood, and have only rarely been studied on the continental shelf. Here we present the results of a study into the vertical and horizontal movement behaviour of 42 European eels (Anguilla anguilla L.) tagged with electronic tags that migrated through the North Sea and English Channel towards and into the Atlantic Ocean during their spawning migration. We used actograms, periodograms and linear mixed effects models to determine the periodicity and significance of the timing and pattern of vertical movement and activity. Overall, eels had a complex behavioural repertoire that included classical diel vertical migration (DVM), reverse DVM and vertical movement behaviours that synchronized with tidal patterns. All of the eels that were tracked showed one or more of these behaviours during their time at liberty, and many exhibited all of them. We also observed that the eels had a higher horizontal migration speed when the current in the favourable direction was stronger. This, together with the vertical movement synchronized with the tides, suggests the eels adopt selective tidal stream transport. Finally, tracked eels had a higher vertical movement range at night compared to daytime. We hypothesize that these behaviours are driven by bio-energetic efficient movement, navigation and predator avoidance.

Glass eel (Anguilla anguilla L. 1758) feeding behaviour during upstream migration in an artificial waterway.

The transition from marine to fresh water is a challenging task for juvenile eels. This critical step in the early eels' life is preceded by a metamorphosis from the oceanic larval to the continental glass eel stage, requiring major energy-demanding morphological, physiological and behavioural modifications during which time these animals do not feed. The success of the glass eels' inland migration after metamorphosis will largely depend on remaining energy levels, which can be supplemented only by resuming food uptake. Although it is crucial for their survival and the maintenance of the population, the feeding behaviour of glass eels is still an understudied aspect of the eels' complex life cycle. Many uncertainties about the phenology, diet, potential prey preferences and their relation with migration modus (migratory vs. sedentary) still remain. In this study, the authors analysed the stomach and gut contents of 458 European glass eels (Anguilla anguilla L. 1758) captured in a drainage canal connecting a small mesotidal estuary with an adjacent polder area during the spring migration seasons of 2016 and 2017. They demonstrated that although glass eels started feeding briefly upon arrival in the estuary, food uptake for early arrivals was restricted to a minority that sparsely feed on detritus and some worm-like benthic invertebrates. Along the season, food uptake intensified eventually engaging all glass eels and their dietary palette diversified including a wide array of planktonic and benthic organisms. Crustacean plankton (mainly cyclopoid copepods) was an important part of the glass eel diet, whereas benthic oligochaetes were less abundant as food source in spite of their high presence in the sediments. No clear differences in feeding behaviour could be observed between migratory and sedentary glass eels. This study showed that glass eels can use highly artificial and dynamic drainage canals as feeding ground during their critical marine/freshwater transition. This outcome is also a plea to improve the accessibility of alternative (unnatural) migration routes between the ocean and suitable freshwater growth habitats for the European eel.

Mapping silver eel migration routes in the North Sea.

Recent developments in tracking technology resulted in the mapping of various marine spawning migration routes of the European eel (Anguilla anguilla). However, migration routes in the North Sea have rarely been studied, despite many large European rivers and hence potential eel growing habitat discharge into the North Sea. In this study, we present the most comprehensive map to date with migration routes by silver European eels in the North Sea and document for the first time successful eel migration through the English Channel. Migration tracks were reconstructed for 42 eels tagged in Belgium and 12 in Germany. Additionally, some eels moved up north to exit the North Sea over the British Isles, confirming the existence of two different routes, even for eels exiting from a single river catchment. Furthermore, we observed a wide range in migration speeds (6.8-45.2 km day-1). We hypothesize that these are likely attributed to water currents, with eels migrating through the English Channel being significantly faster than eels migrating northward.

Fish species sensitivity classification for environmental impact assessment, conservation and restoration planning.

Species conservation, river rehabilitation, stock enhancement, environmental impact assessment and related planning tools require indicators to identify significant impacts but also mitigation success. Since river systems are shaped by disturbances from floods and droughts, typical riverine fish species should have evolved life history traits providing resilience against such disturbances. This study compiled and analyzed resilience traits of European lampreys and fish species to derive a novel sensitivity classification of species to mortality. We assembled life history traits like maximum length, migration type, mortality, fecundity, age at maturity, and generation time of 168 species and created a novel method to weigh and integrate all traits to generate a final sensitivity score from one (low sensitivity) to three (high sensitivity) for each species. Large-bodied, diadromous, rheophilic and lithophilic species such as sturgeons, sea trout, and Atlantic salmon usually appeared to have high sensitivity to additional adult fish mortality, whereas small-bodied, limnophilic and phytophilic species with fast generation cycles were of low sensitivity. The final scoring and classification of 168 European lampreys and fish species according to their sensitivity can be easily regionalized by selecting the most sensitive candidates according to the local species pool. This sensitivity classification has major implications for advancing impact assessment, allowing better targeting of species for conservation measures, benchmarking progress during rehabilitation and enhancing the objective evaluation of the success of restoration projects.

Urbanization drives cross-taxon declines in abundance and diversity at multiple spatial scales.

The increasing urbanization process is hypothesized to drastically alter (semi-)natural environments with a concomitant major decline in species abundance and diversity. Yet, studies on this effect of urbanization, and the spatial scale at which it acts, are at present inconclusive due to the large heterogeneity in taxonomic groups and spatial scales at which this relationship has been investigated among studies. Comprehensive studies analysing this relationship across multiple animal groups and at multiple spatial scales are rare, hampering the assessment of how biodiversity generally responds to urbanization. We studied aquatic (cladocerans), limno-terrestrial (bdelloid rotifers) and terrestrial (butterflies, ground beetles, ground- and web spiders, macro-moths, orthopterans and snails) invertebrate groups using a hierarchical spatial design, wherein three local-scale (200 m × 200 m) urbanization levels were repeatedly sampled across three landscape-scale (3 km × 3 km) urbanization levels. We tested for local and landscape urbanization effects on abundance and species richness of each group, whereby total richness was partitioned into the average richness of local communities and the richness due to variation among local communities. Abundances of the terrestrial active dispersers declined in response to local urbanization, with reductions up to 85% for butterflies, while passive dispersers did not show any clear trend. Species richness also declined with increasing levels of urbanization, but responses were highly heterogeneous among the different groups with respect to the richness component and the spatial scale at which urbanization impacts richness. Depending on the group, species richness declined due to biotic homogenization and/or local species loss. This resulted in an overall decrease in total richness across groups in urban areas. These results provide strong support to the general negative impact of urbanization on abundance and species richness within habitat patches and highlight the importance of considering multiple spatial scales and taxa to assess the impacts of urbanization on biodiversity.

A comparative hierarchical analysis of bacterioplankton and biofilm metacommunity structure in an interconnected pond system.

It is unknown whether bacterioplankton and biofilm communities are structured by the same ecological processes, and whether they influence each other through continuous dispersal (known as mass effects). Using a hierarchical sampling approach we compared the relative importance of ecological processes structuring the dominant fraction (relative abundance ≥0.1%) of bacterioplankton and biofilm communities from three microhabitats (open water, Nuphar and Phragmites sites) at within- and among-pond scale in a set of 14 interconnected shallow ponds. Our results demonstrate that while bacterioplankton and biofilm communities are highly distinct, a similar hierarchy of ecological processes is acting on them. For both community types, most variation in community composition was determined by pond identity and environmental variables, with no effect of space. The highest ß-diversity within each community type was observed among ponds, while microhabitat type (Nuphar, Phragmites, open water) significantly influenced biofilm communities but not bacterioplankton. Mass effects among bacterioplankton and biofilm communities were not detected, as suggested by the absence of within-site covariation of biofilm and bacterioplankton communities. Both biofilm and plankton communities were thus highly structured by environmental factors (i.e., species sorting), with among-lake variation being more important than within-lake variation, whereas dispersal limitation and mass effects were not observed.

The effects of zinc on the structure and functioning of a freshwater community: A microcosm experiment.

A major problem with risk assessment of chemicals is the extrapolation of laboratory single-species toxicity tests, which oversimplify the actual field situation by ignoring species interactions, to natural communities. The authors tested if the bioavailability-normalized 5% hazardous concentration (HC5) estimated from chronic planktonic single-species toxicity data (HC5plankton ) for zinc (Zn) is protective for a plankton community and investigated the direct and indirect effects of Zn (at HC5 and HC50) on a freshwater community's structure and function. Microcosms were exposed to 3 different Zn concentrations (background, HC5plankton = 75 µg Zn/L and HC50plankton = 300 µg Zn/L) for 5 wk. The planktonic groups revealed a consistent no-observed-effect concentration for the community of 75 µg Zn/L, similar to or higher than the HC5plankton , thus suggesting its protectiveness in the present study. At 300 µg Zn/L a significant reduction in cladocerans resulted in increases of rotifer, ciliate, and phytoplankton abundance. In addition, the phytoplankton community shifted in dominance from grazing-resistant to edible species. Contrary to the species sensitivity distribution (SSD) prediction, which identified phytoplankton as the most sensitive group, only the total chlorophyll and the abundance of 2 phytoplankton species were adversely affected at 300 µg Zn/L. Thus, although the HC5 estimated from the bioavailability-normalized SSD was overall protective for the plankton community, the SSD was not able to correctly predict the species sensitivity ranking within their community context at the HC50. Environ Toxicol Chem 2016;35:2698-2712. © 2016 SETAC.

The common bloom-forming cyanobacterium Microcystis is prone to a wide array of microbial antagonists.

Many degraded waterbodies around the world are subject to strong proliferations of cyanobacteria - notorious for their toxicity, high biomass build-up and negative impacts on aquatic food webs - the presence of which puts serious limits on the human use of affected water bodies. Cyanobacterial blooms are largely regarded as trophic dead ends since they are a relatively poor food source for zooplankton. As a consequence, their population dynamics are generally attributed to changes in abiotic conditions (bottom-up control). Blooms however generally contain a vast and diverse community of micro-organisms of which some have shown devastating effects on cyanobacterial biomass. For Microcystis, one of the most common bloom-forming cyanobacteria worldwide, a high number of micro-organisms (about 120 taxa) including viruses, bacteria, microfungi, different groups of heterotrophic protists, other cyanobacteria and several eukaryotic microalgal groups are currently known to negatively affect its growth by infection and predation or by the production of allelopathic compounds. Although many of these specifically target Microcystis, sharp declines of Microcystis biomass in nature are only rarely assigned to these antagonistic microbiota. The commonly found strain specificity of their interactions may largely preclude strong antagonistic effects on Microcystis population levels but may however induce compositional shifts that can change ecological properties such as bloom toxicity. These highly specific interactions may form the basis of a continuous arms race (co-evolution) between Microcystis and its antagonists which potentially limits the possibilities for (micro)biological bloom control.

The Impact of Conservation Management on the Community Composition of Multiple Organism Groups in Eutrophic Interconnected Man-Made Ponds.

Ponds throughout the world are subjected to a variety of management measures for purposes of biodiversity conservation. Current conservation efforts typically comprise a combination of multiple measures that directly and indirectly impact a wide range of organism groups. Knowledge of the relative impact of individual measures on different taxonomic groups is important for the development of effective conservation programs. We conducted a field study of 28 man-made ponds, representing four management types differing in the frequency of periodic pond drainage and the intensity of fish stock management. We disentangled the relative importance of direct and indirect effects of pond management measures on the community composition of phytoplankton, zooplankton, aquatic macro-invertebrates, submerged and emergent vascular plants. With the exception of phytoplankton, pond management had strong effects on the community composition of all investigated biota. Whether management affected communities directly or indirectly through its impact on fish communities or local environmental conditions in the pond varied between organism groups. Overall, the impact of pond drainage regime and fish community characteristics on the community composition of target organism groups were more important than local environmental conditions. The majority of taxa were negatively associated with fish density, whereas multiple emergent plant species and several taxa of aquatic macro-invertebrates were positively affected by increased drainage frequency. The effects of fish community and drainage tended to be largely independent. The present study indicates that pond drainage is an important element for biodiversity conservation in eutrophicated shallow and interconnected man-made ponds.

Expansion of the 'Reticulosphere': Diversity of Novel Branching and Network-forming Amoebae Helps to Define Variosea (Amoebozoa).

Amoebae able to form cytoplasmic networks or displaying a multiply branching morphology remain very poorly studied. We sequenced the small-subunit ribosomal RNA gene of 15 new amoeboid isolates, 14 of which are branching or network-forming amoebae (BNFA). Phylogenetic analyses showed that these isolates all group within the poorly-known and weakly-defined class Variosea (Amoebozoa). They are resolved into six lineages corresponding to distinct new morphotypes; we describe them as new genera Angulamoeba (type species Angulamoeba microcystivorans n. gen., n. sp.; and A. fungorum n. sp.), Arboramoeba (type species Arboramoeba reticulata n. gen., n. sp.), Darbyshirella (type species Darbyshirella terrestris n. gen., n. sp.), Dictyamoeba (type species Dictyamoeba vorax n. gen., n. sp.), Heliamoeba (type species Heliamoeba mirabilis n. gen., n. sp.), and Ischnamoeba (type species Ischnamoeba montana n. gen., n. sp.). We also isolated and sequenced four additional variosean strains, one belonging to Flamella, one related to Telaepolella tubasferens, and two members of the cavosteliid protosteloid lineage. We identified a further 104 putative variosean environmental clone sequences in GenBank, comprising up to 14 lineages that may prove to represent additional novel morphotypes. We show that BNFA are phylogenetically widespread in Variosea and morphologically very variable, both within and between lineages.

How to maximally support local and regional biodiversity in applied conservation? Insights from pond management.

Biodiversity and nature values in anthropogenic landscapes often depend on land use practices and management. Evaluations of the association between management and biodiversity remain, however, comparatively scarce, especially in aquatic systems. Furthermore, studies also tend to focus on a limited set of organism groups at the local scale, whereas a multi-group approach at the landscape scale is to be preferred. This study aims to investigate the effect of pond management on the diversity of multiple aquatic organism groups (e.g. phytoplankton, zooplankton, several groups of macro-invertebrates, submerged and emergent macrophytes) at local and regional spatial scales. For this purpose, we performed a field study of 39 shallow man-made ponds representing five different management types. Our results indicate that fish stock management and periodic pond drainage are crucial drivers of pond biodiversity. Furthermore, this study provides insight in how the management of eutrophied ponds can contribute to aquatic biodiversity. A combination of regular draining of ponds with efforts to keep ponds free of fish seems to be highly beneficial for the biodiversity of many groups of aquatic organisms at local and regional scales. Regular draining combined with a stocking of fish at low biomass is also preferable to infrequent draining and lack of fish stock control. These insights are essential for the development of conservation programs that aim long-term maintenance of regional biodiversity in pond areas across Europe.

Planktonic ciliate community structure in shallow lakes of lowland Western Europe.

Temperate shallow meso- to eutrophic lakes can exist in one of two alternative states with contrasting foodwebs, referred to as the clear-water and the turbid state. We describe the planktonic ciliate communities of such lakes based on a survey of 66 northwestern European lakes. Ciliates were enumerated and identified to species level according to the quantitative protargol staining technique. Ciliate biomass was on average twice as high in the turbid than in the clear-water lakes. The ciliate communities were dominated by oligotrichs and protostomatids, and no differences in functional composition or α-diversity could be detected between turbid and clear-water lakes, although ß-diversity tended to be higher in the latter. At the species level, however, community structure strongly differed between turbid and clear-water lakes, and several indicator species could be identified for the different lake categories. Variation partitioning showed that nutrient status did not explain ciliate community structure independent of the alternative states, while lake area was identified as an additional structuring factor for the ciliate communities. These results stress the importance of the ecosystem structure in shaping ciliate communities in temperate shallow lakes and suggest that nutrient status has little direct effect on ciliate community structure in such lakes.

Strong effects of amoebae grazing on the biomass and genetic structure of a Microcystis bloom (Cyanobacteria).

Despite its importance for bloom toxicity, the factors determining the population structure of cyanobacterial blooms are poorly understood. Here, we report the results of a two-year field survey of the population dynamics of Microcystis blooms in a small hypertrophic urban pond. Microscopic enumeration of Microcystis and its predators and parasites was combined with pigment and microcystin analysis and denaturing gradient gel electrophoresis of the ITS rDNA region to assess population dynamics and structure. Two main Microcystis morpho- and ITS types were revealed, corresponding to M. aeruginosa and M. viridis. In both years, high population densities of naked amoebae grazing on Microcystis coincided with rapid decreases in Microcystis biomass. In one year, there was a shift from heavily infested M. aeruginosa to the less-infested M. viridis, allowing the bloom to rapidly recover. The preference of amoebae for M. aeruginosa was confirmed by grazing experiments, in which several amoeba strains were capable of grazing down a strain of M. aeruginosa, but not of M. viridis. Zooplankton and chytrid parasites appeared to be of minor importance for these strong and fast reductions in Microcystis biomass. These findings demonstrate a strong impact of small protozoan grazers on the biomass and genetic structure of Microcystis blooms.

Diversity and habitat specificity of free-living protozoa in commercial poultry houses.

Despite stringent biosecurity measures, infections by bacterial food pathogens such as Campylobacter are a recurrent problem in industrial poultry houses. As the main transmission route remains unclear, persistence of these infections has been linked to bacterial survival and possibly multiplication within protozoan vectors. To date, however, virtually no information is available on the diversity and occurrence of free-living protozoa in these environments. Using a combination of microscopic analyses of enrichment cultures and molecular methods (denaturing gradient gel electrophoresis [DGGE]) on natural samples, we show that, despite strict hygiene management, free-living protozoa are common and widespread throughout a 6-week rearing period in both water and dry samples from commercial poultry houses. Protozoan communities were highly diverse (over 90 morphotaxa and 22 unique phylotypes from sequenced bands) and included several facultative pathogens and known bacterial vectors. Water samples were consistently more diverse than dry ones and harbored different communities, mainly dominated by flagellates. The morphology-based and molecular methods yielded markedly different results: amoebic and, to a lesser degree, ciliate diversity was seriously underestimated in the DGGE analyses, while some flagellate groups were not found in the microscopic analyses. Some recommendations for improving biosecurity measures in commercial poultry houses are suggested.

Covariation between zooplankton community composition and cyanobacterial community dynamics in Lake Blaarmeersen (Belgium).

The cyanobacterial community composition in the mesotrophic Lake Blaarmeersen was determined by denaturing gradient gel electrophoresis (DGGE) of PCR-amplified 16S rRNA gene fragments during two consecutive years to assess the importance of different classes of explanatory variables (bottom-up and top-down factors, physical variables and phytoplankton) in cyanobacterial community dynamics. The most dominant cyanobacteria in Lake Blaarmeersen were Synechococcus (three genotypes), Limnothrix redekei and Anabaena/Aphanizomenon. Analyses of Similarity revealed that the cyanobacterial community in Lake Blaarmeersen differed significantly between the growing season and the winter season as well as between the epilimnion and hypolimnion during the stratified periods. Mantel tests revealed significant correlations between the DGGE data and bottom-up factors, physical variables, the phytoplankton community composition and, interestingly, the zooplankton community composition. In general, the zooplankton community composition (especially the cladoceran community) was more important in structuring the cyanobacterial community than the total zooplankton biomass. This study shows that grazing zooplankton communities can have a relatively strong impact on the cyanobacterial community dynamics and that this impact can be equally important as bottom-up processes regulated by nutrient concentrations and/or physical variables.

Rapid screening for freshwater bacterial groups by using reverse line blot hybridization.

The identification of phylogenetic clusters of bacteria that are common in freshwater has provided a basis for probe design to target important freshwater groups. We present a set of 16S ribosomal RNA gene-based oligonucleotide probes specific for 15 of these freshwater clusters. The probes were applied in reverse line blot hybridization, a simple method that enables the rapid screening of PCR products from many samples against an array of probes. The optimized assay was made stringent to discriminate at approximately the single-mismatch level. This made 10 of the probes highly specific, with at least two mismatches to the closest noncluster member in the global database. Screening of PCR products from bacterioplankton of 81 diverse lakes from Belgium, The Netherlands, Denmark, Sweden, and Norway showed that the respective probes were reactive against 5 to 100% of the lake samples. Positive reactivity of six highly specific probes showed that bacteria from actinobacterial clusters ACK-M1 and Sta2-30 and from verrucomicrobial cluster CLO-14 occurred in at least 90% of the investigated lakes. Furthermore, bacteria from alpha-proteobacterial cluster LD12 (closely related to the marine SAR11 cluster), beta-proteobacterial cluster LD28 and cyanobacterial cluster Synechococcus 6b occurred in more than 70% of the lakes. Reverse line blot hybridization is a new tool in microbial ecology that will facilitate research on distribution and habitat specificity of target species at relatively low costs.