Testing the European stream typology of the Water Framework Directive for macroinvertebrates

In the EU Water Framework Directive (WFD) a typological framework is defined for assessing the ecological quality of water bodies in the future. The conditions in the Directive impose a strong demand for `new' assessment systems. During the AQEM project an assessment system was developed for European streams using macroinvertebrates. The aim of this study was to test if the typology suggested in the WFD is useful for developing an assessment system for macroinvertebrates in streams. In total 889 streams of 29 stream types were sampled in eight countries all over the major geographical gradients in Europe. These stream types fit the WFD typological demands and fit to the major European geographic regions (ecoregions). The sites included gradients from reference conditions (for the definition see Nijboer et al., 2004) to sites with bad ecological quality. Despite standardisation there were large differences between the participating countries concerning the number of taxa, the number of specimens and the taxonomic resolution. The data, including macroinvertebrates and environmental variables were analysed by using Canonical Correspondence Analysis (CCA). The observed macroinvertebrate distribution largely supported the WFD typological criteria. This means that the major macroinvertebrate distribution patterns in European streams follow climatological and geomorphological conditions and are well distinguished in terms of stream types. Furthermore, it was shown that large scale factors affected the macroinvertebrate distribution even on a very fine scale. Most explanatory variables seemed to be scale independent. Even at a fine scale major factors concerning geology, geomorphology and hydrology added to the species occurrences. Within stream types morphology together with physico-chemistry best explained the macroinvertebrates distribution. In conclusion, the WFD typology is useful for an assessment system for streams using macroinvertebrates. The large scale factors were indeed the variables that explained most of the variation in species composition. But as these factors even strongly act at the scale of stream types, a further refinement is most probably necessary to disentangle typological actors from water quality ones.     

The AQEM/STAR taxalist – a pan-European macro-invertebrate ecological database and taxa inventory

The European list of aquatic macro-invertebrate taxa, and its associated ecological database, originated within the context of the AQEM project and have been extended during the STAR project. The AQEM/STAR taxalist is a product of co-operation between applied freshwater ecologists and scientists from different zoological fields, applied partners and the administration. The basic idea is that a sound understanding of benthic invertebrate ecology is a prerequisite for the implementation of a biological approach to aquatic ecosystem management in Europe. The database has been generated under the management of BOKU (University of Natural Resources and Applied Life Sciences, Vienna) and UDE (University of Duisburg-Essen) and provides an important means of standardisation and unification of ecological classifications in Europe. This paper outlines the aims for setting up the AQEM/STAR macro-invertebrate taxalist and autecological database and provides a current summary of the numbers of aquatic orders, families, species, and species occurrences in 14 European countries. The number of available and applicable assignments of taxa to each ecological parameter is summarised and examples are given for different parameters and taxonomic groups. Gaps in the autecological information are identified and discussed. Besides its ecological relevance, the operational character of this database is underlined by the fact that it provides the associated taxon codes for each of five different European assessment systems for nearly 10,000 European macro-invertebrate taxa.     

Distribution and ecology of Middle Atlantic Bight Oligochaeta

Distribution and ecology of continental shelf oligochaetes are poorly understood. From 1975 through 1977 baseline ecological studies were conducted in the Middle Atlantic Bight prior to oil exploration activities. A total of 38 species in three oligochaete families were collected. Of this total one family and 29 of the species were undescribed at the time of collection. The most abundant and widely distributed species were Limnodriloides monothecus, L. medioporus, Bathydrilus longus, Phallodrilus biprostatus, and Tubificoides diazi. No single species dominated the oligochaete fauna. All the oligochaete species responded to changes in physical parameters over the shelf similarly to the total macrofauna. Seasonal trends within the oligochaetes were not found. Species that preferred coarser sand sediments were all small interstitial forms. The finer sand and eurytopic species were all burrowers. Zoogeographically the Middle Atlantic Bight oligochaetes were predominantly temperate (16 species) with some southern species (8). Seven species were broadly distributed from the Caribbean to Massachusetts or around the world. Seven species were only known from the area studied.Contribution No. 1397 of the Virginia Institute of Marine Science.     

The role of oligochaetes in the management of waters

To better understand the role of an aquatic oligochaete in the management of water quality, it was determined if oligochaetes were representative of water quality, if indices based on oligochaetes were widely applicable to access water quality, and if oligochaete assemblages were related to environmental factors other than organic pollution. It was found that if only one monitoring sample a year was taken in December, about 73% of the more common species were collected. Seasonality did not seem to affect the taxa collected except for forms in low presence, e.g., Nais. Several indices we used were not applicable to regulated streams. Oligochaete ordination procedures revealed only a partial picture of the environmental conditions of regulated streams.     

Effects of invasive benthic macroinvertebrates on assessment methods of the EU Water Frame Work Directive

The assessment of the ecological status of aquatic ecosystems for the implementation of the European Water Framework Directive is based on different methods, many of which include the study of macroinvertebrate organisms. We examined whether and how invasive species affect the calculation of metrics used for fresh water assessment utilizing data from different substrate types in the streams of Central Germany. In these streams, the proportion of macroinvertebrate invasives increased from a very low fraction of the community up to 75% of species, or 96% of individuals, within 15 years. The values of the German Saprobic Index (GSI, reflecting the degree of organic load) did not differ significantly between different substrate types or within the examined stream reaches. However, saprobic valences of the species recorded did differ between substrates and between localities. This obvious contradiction is caused by an abundance of exotic species all having a medium GSI index between 2.2 and 2.3. Moreover, the minimum total number of organisms required for this method cannot be reached for a large majority of the sites dominated by invasives making it impossible to use the GSI. Regarding distribution of zonation types, the proportion of metarhithral species is low, and it is even lower on hard substrates than it is on soft substrates. This contradicts the general assumption that dwellers preferring metarhithral reaches (=lower-trout region) will show an affinity for habitats with higher oxygen concentrations and stronger flow velocities, both appearing more on hard than on soft substrates. This latter contradiction results from the classification of the most abundant invasive species Potamopyrgus antipodarum, Dikerogammarus villosus, and Chelicorophium curvispinum in the AQEM taxa reference list. The metrics ‘locomotion type’ and ‘feeding type’ were also affected by the presence of invasive species. Possible options for overcoming these assessment problems are discussed.     

Comparative body wall musculature and muscle fibre ultrastructure in branchiobdellidans (Annelida: Clitellata), and their phylogenetic significance

The body wall muscles in five species of branchiobdellidans are all arranged in the oligochaete pattern and the muscle fibres are obliquely striated. The structure of the circular muscle fibres do vary to some degree. The longitudinal muscle fibres in Ankyrodrilus legaeus, Branchiobdella kozarovi, and Xironogiton instabilis all are round circomyarian and thus double-obliquely striated. These species represent three of the four genera composing the family Branchiobdellidae. Although Bdellodrilus illuminatus and Cambarincola fallax, from the families Bdellodrilidae and Cambarincolidae, respectively, also possess a few round circomyarian fibres, most are polyplatymyarian comparable to single-obliquely striated fibres. A similar division of branchiobdellidan families is obtained based on the number of anterior nephridial pores. The muscular structure in the branchiobdellidans shows both similarities and differences with the leeches and the lumbriculid oligochaetes. One phylogenetic explanation for this is that the branchiobdellidans separated from the common clitellate ancestor before the oligochaetes and leeches became recognizable taxa.     

Overview and application of the AQEM assessment system

The main objective of the European Union (EU) funded project AQEM¹was to develop a framework of an assessment system for streams in Europe based on benthic macroinvertebrates that fulfils the requirements of the EU Water Framework Directive. Initial assessment methods for 28 European stream types and more generally applicable tools for stream biomonitoring in Europe were generated. The development of the system was based on a newly collected data set covering stream types in Austria, the Czech Republic, Germany, Greece, Italy, The Netherlands, Portugal and Sweden. Altogether, 901 benthic invertebrate samples were taken using a standardised multi-habitat sampling procedure and a large number of parameters describing the streams and their catchments was recorded for all sampling sites. From the stream and catchment characteristics measures of stress were derived. A large number of metrics was tested independently for each of the stream types, to identify the response of each metric to degradation of a site. This process resulted in up to 18 core metrics for the individual stream types, which were combined into a different multimetric index in each country. The multimetric AQEM assessment system is used to classify a stream stretch into an Ecological Quality Class ranging from 5 (high quality) to 1 (bad quality) and often provides information on the possible causes of degradation. AQEM provides a taxa list of 9557 European macroinvertebrate taxa with associated autecological information, a software package for performing all the calculations necessary for applying the multimetric AQEM assessment system and a manual describing all aspects of the application of the system from site selection to data interpretation.     

Diversity and distribution of Tubificidae, Naididae, and Lumbriculidae (Annelida: Oligochaeta) in the Netherlands: an evaluation of twenty years of monitoring data

Data from 24 water management districts and the rivers Rhine and Meuse in the Netherlands were used to study geographical distribution, relative occurrence, and environmental requirements of 76 aquatic oligochaetes (families Tubificidae, Naididae, and Lumbriculidae) (Annelida, Clitellata). Approximately 50% of the 76 species that occur in the Netherlands are uncommon, rare, or very rare. The other half of the species are common, very common or abundant. The abundant species are: Stylaria lacustris, Ophidonais serpentina, Limnodrilus claparedeianus, Limnodrilus hoffmeisteri, and Lumbriculus variegatus. With the exception of several brackish water species (those restricted in distribution to water management districts close to the sea that are influenced by salt water influx) and specific running water species (restricted mainly to the eastern part of the Netherlands), most of the species occurred throughout the whole Netherlands. The species distribution was related to environmental variables using ordination. In general, species distribution was correlated with either large waters with high chloride and phosphorus concentrations and a high hydrogen ion concentration (as pH), or with small forested (running) waters in more natural (undeveloped) areas that occasionally become intermittent. Vegetation cover was positively correlated with several swimming species in the family Naididae. While the distribution of aquatic oligochaetes in some families occurring in the Netherlands is known to some extent, the occurrence and distribution of rare and small taxa, particularly those that are difficult to identify taxonomically, is virtually unknown. Some of the rare oligochaete taxa, especially those associated with unique habitats, have received only cursory attention. Also in the data studied, the observations of the more rare species were too few to draw conclusions. To improve our knowledge of oligochaete distribution in the Netherlands, additional research should focus on rare species associated with special habitats and water types (natural areas) and those taxa in poorly known families. The standardisation of sampling and processing methodologies, particularly the use of nets and sieves with fine-meshed screening, will ensure the collection of the smaller species of oligochaetes. Subsequently, oligochaetes should be identified to species level by experienced taxonomists trained in oligochaete identification. Finally, many aquatic oligochaete species are identifiable only when sexually mature and therefore the time of year in which samples are collected is critical to the accurate representation of true oligochaete diversity at any given site. For analysing the relation between species and environmental variables the best option is to use composite data from spring and autumn.     

The effect of organic pollution on the abundance and distribution of aquatic oligochaetes in an urban water basin, Taiwan

Aquatic oligochaetes are abundant in polluted areas and are, therefore, commonly used as bioindicators to study organic pollution in rivers and streams. In order to develop a species-level oligochaete biotic index to reflect the River Pollution Index (RPI) in the Taichung Water Basin in Taiwan, we conducted a systematic sampling scheme to collect aquatic oligochaetes from the sediment samples of watercourses in the Taichung Water Basin, Taiwan. We evaluated the relationships between aquatic oligochaetes and the sewage pollution using statistical methods. The distribution of aquatic oligochaetes in relation to environmental variables, such as water quality and sediment characteristics of the regional urban contaminated streams was expressed by Canonical Correspondence Analysis (CCA). We identified 17 species of aquatic oligochaetes (Annelida, Clitellata) including 3 species of Tubificidae, 13 species of Naididae, and 1 species of Enchytraeidae from the watercourses of an urban region in the Taichung Water Basin in Taiwan, during the summer and winter of 2005 and 2006. A positive correlation was found between the total abundance of aquatic oligochaetes and the RPI (r = 0.58, P < 0.05). However, only population density of the most abundant tubificid, Limnodrilus hoffmeisteri, increased with increasing RPI values and a significantly negative correlation was found between the population density of the naidid, Nais communis, and RPI values. The results of CCA indicated that certain naidids, such as Aulophorus furcatus and Allonais gwaliorensis also tolerated extremely polluted environments in upper stream or stony habitats, implying that tubificids should not be the sole representation of simple biotic indices but should also include pollution-tolerant naidids. We found that the community structure of aquatic oligochaetes was influenced by short-term variations in microhabitat rather than according to seasonal factors in our study region. The results proved that aquatic oligochaetes were sensitive enough to provide a supplement for the regional urban pollution assessment applications for biotic indicators at the species-level.     

Oligochaetes in southern European groundwater: new records and an overview

This work deals with an overview of the faunal composition and presents a general ecological approach to the groundwater oligochaetes in southern Europe. The species richness is high and similar to that of the superficial waters in the same area. Among the 150 species that we found, about 29 are stygobiont, so the total number of stygobiont species in the area studied has been increased to 58. Ten of these species are tubificids with marine phyletic affinities. A list of the most frequently found species and the relative abundance of the different families and genera collected in the main environments investigated are presented. Although enchytraeids often dominated the fauna, lumbriculids and tubificids exhibit a high level of diversity and comprised the majority of stygobiont and stygophilous taxa found. The discovery of members of the family Parvidrilidae in southern European caves is the first outside of North America. This indicates that the family is widely distributed in the Palearctic region and typical of underground waters. Here, we update the previous evaluation of the knowledge about oligochaetes inhabiting groundwater: 96 species of stygobiont freshwater oligochaetes (sensu stricto) are now known in the world and 81 of them have been recorded from the Palearctic region.     

Species composition,distribution, and abundance of oligochaetes in the Kanawha River,West Virginia

Species composition, distribution, and abundance of oligochaetes in the Kanawha River, West Virginia, were investigated. Eighteen taxa of oligochaetes including 7 tubificids, 8 naidids, and representatives of the families Enchytraeidae, Lumbriculidae, and Aelosomatidae were collected. While most of the species collected are cosmopolitan, Pristina synclites and Stephensoniana sp. previously reported only from Asia and Africa were collected in the Kanawha. Other peculiarities of the Kanawha oligochaete fauna included the common occurrence of specimens of Limnodrilus cervix and Limnodrilus hoffmeisteri with atypical penis sheaths. The dominant and most widely distributed of the Kanawha River oligochaetes were immature tubificids lacking capilliform chaetae, Branchiura sowerbyi, Dero cf. obtusa, Limnodrilus hoffmeisteri, and Aulodrilus pigueti.Oligochaete densities ranged from 401–2403 individuals/m² in areas presumed to be nonpolluted, to 1–671 individual/m² at sites with suspected toxicity. The highest density of oligochaetes (2835 individuals/m²) occurred at a site of organic enrichment.     

Taxonomic composition of rotifers in Lake Peipsi

161 rotifer taxa 46 of which are new records for Estonia were identified in the north-western part of Lake Peipsi. The greatest number of taxa occurred in August (93), followed by September (79). In all, 19 families (Monogononta) are represented with 41 genera. The dominant families are Brachionidae (32 taxa) and Synchaetidae (16). The species of Anuraeopsis, Conochilus, Kellicottia, Keratella, Notholca, Polyarthra and Synchaeta are dominating. The taxonomic composition is analysed in the ecological aspect. Some species of zoogeographic interest, including a supposedly endemic Ploesoma peipsiense, are reported.     

Establishing reference conditions for European streams

The European Water Framework Directive stipulates that Member States have to assess the ecological status of a water body by comparing the present to the expected reference condition. In the AQEM project participating countries used the criteria from the Water Framework Directive to select reference sites for each stream type. The purpose of this study was to evaluate the suitability of these criteria and to validate the reference conditions chosen, by comparing the classification of sites before analysis with the final assessment of the sites using the assessment system which was developed within the AQEM project. Our study showed that not all criteria for reference conditions could be met for all stream types. This implies that `true' reference sites could not be selected for all stream types within the AQEM project. In our study, the differences between ecoregions, countries, and stream types were interpreted in terms of human impact. Validation of reference conditions showed differences between classification of sites as reference sites based on criteria to be used in the field and directly interpreting environmental or biological data and the result of the final assessment system. In some countries it was not possible to select reference conditions, because most of the a priori criteria were met. For example, reference conditions for Dutch stream types were established using historical data and predictions based on data from other geographical regions. Using data from adjacent countries appeared to be a feasible method for establishing reference conditions. Reference sites taken from other geographical areas represented a higher ecological quality than the Dutch sites of good ecological quality. However, metric results showed a large overlap between good and high ecological status. Historical information, on the other hand, was not found to be useful in metric calculations because of the confounding differences in sampling methods. One strong advantage of using historical information is that rare species that are locally extinct but still occur in the same stream type in other geographical areas can be added to the expected reference conditions. When and which methods can be used for establishing reference conditions is illustrated in a decision tree.     

Countering morphological ambiguities: development of a PCR assay to assist the identification of <Emphasis Type="Italic">Tubifex tubifex</Emphasis> oligochaetes

The freshwater oligochaete, Tubifex tubifex, is a common resident of organic-rich sediments worldwide. Although it is a familiar species to fish enthusiasts, toxicologists and parasitologists, T. tubifex often confounds definitive identification due to the similarity that immature specimens bear to several other common oligochaetes, and given the degree of plasticity of key morphological characters due to environmental conditions and/or age of specimen. To solve this identity crisis, we used a polymerase chain reaction based molecular approach and developed a T. tubifex specific assay with primers that amplify a 192 bp fragment of the internal transcribed spacer region 1 ribosomal DNA. We tested these primers on four T. tubifex mitochondrial genotypes, and on other oligochaete species from nine genera. The primers amplified all specimens identified morphologically asT. tubifex. They did not amplify any other species, including morphologically similar worms possessing hair chaetae (Dero digitata, Ilyodrilus templetoni, Tubifex ignotus or Rhyacodrilus spp.) or other oligochaetes often found with T. tubifex (Lumbriculus variegates, Limnodrilus hoffmeisteri, Stylodrilus heringianus or Trichodrilus sp.). This technique should remove the uncertainties all too often associated with identification of T. tubifex.     

The fossil record of extant elasmobranchs

Sharks and their relatives (Elasmobranchii) are highly threatened with extinction due to various anthropogenic pressures. The abundant fossil record of fossil taxa has allowed the tracing of the evolutionary history of modern elasmobranchs to at least 250 MYA; nonetheless, exactly how far back the fossil record of living taxa goes has never been collectively surveyed. In this study, the authors assess the representation and extent of the fossil record of elasmobranchs currently living in our oceans by collecting their oldest records and quantifying first appearance dates at different taxonomic levels (i.e., orders, families, genera and species), ecological traits (e.g., body size, habitat and feeding mechanism) and extinction risks (i.e., threatened, not threatened and data deficient). The results of this study confirm the robust representation of higher taxonomic ranks, with all orders, most of the families and over half of the extant genera having a fossil record. Further, they reveal that 10% of the current global species diversity is represented in the geological past. Sharks are better represented and extend deeper in time than rays and skates. While the fossil record of extant genera (e.g., the six gill sharks, Hexanchus) goes as far back as c. 190 MYA, the fossil record of extant species (e.g., the sand shark, Carcharias taurus Rafinesque 1810) extends c. 66 MYA. Although no significant differences were found in the extent of the fossil record between ecological traits, it was found that the currently threatened species have a significantly older fossil record than the not threatened species. This study demonstrate that the fossil record of extant elasmobranchs extends deep into the geologic time, especially in the case of threatened sharks. As such, the elasmobranch geological history has great potential to advance the understanding of how species currently facing extinction have responded to different stressors in the past, thereby providing a deep-time perspective to conservation.     

Are patterns in the taxonomic,biological and ecological traits of water beetles congruent in Mediterranean ecosystems?

1. Coleoptera species show considerable diversity in life histories and ecological strategies, which makes possible their wide distribution in freshwater habitats, including highly stressed ones such as saline or temporary waterbodies. Explaining how particular combinations of traits allow species to occupy distinctive habitats is a central question in ecology. 2. A total of 212 sites, sampled over a wide range of inland aquatic habitats in the south‐eastern Iberian Peninsula, yielded 272 species belonging to 68 genera and 11 families. The affinities of genera for 11 biological and 11 ecological traits, gathered from literature and the authors’ own expertise, were used to assess the degree of congruence between taxonomic, biological and ecological traits. 3. Taxonomic richness was significantly related to the number of both biological and ecological trait categories, with the richest families also showing the highest functional and ecological diversity. A fuzzy correspondence analysis performed on the abundance‐weighed array of biological traits separated genera according to categories of diet, feeding habits, respiration, reproduction and locomotion. A similar analysis of ecological traits revealed that preferences related to longitudinal distribution (headwater to mouth), local habitat and current velocity best discriminated genera. At the family level, there was a distinctive functional grouping of genera based on biological traits. Only Elmidae showed noticeable homogeneity across genera for both biological and ecological traits. 4. Co‐inertia analysis demonstrated a significant match between biological and ecological traits (Rv‐correlation = 0.35, P < 0.001). Elmidae genera displayed the highest concordance, whereas Hydraenidae demonstrated the lowest. 5. These results indicate that the predominance of habitat filtering processes in headwater streams yields biological trait conservatism (as shown by Elmidae genera), as well as trait convergence for some specific traits (for instance, respiration) among certain Dytiscidae genera and other typical rheophilic taxa, whereas other biotic factors, such as competition among species, appear more prominent in less stressed habitats. Further knowledge of traits, especially regarding physiological capabilities, is needed to better understand water beetle life history strategies.     

Oligochaetes (Naididae, Tubificidae, Opistocystidae, Enchytraeidae, Sparganophilidae and Alluroididae) of Guyana

About 50 species in more than 20 genera of the microdrile oligochaete families Tubificidae, Naididae, Opistocystidae, and Enchytraeidae and the freshwater megadrile families Sparganophilidae and Alluroididae were identified in recent collections made in Guyana. Only seven species in these families were previously recorded from Guyana. The aquatic oligochaete fauna has similar components to those of the southeastern United States, other locations in South America, and across lower latitudes in the northern hemisphere. A high diversity of species is found in the naidids especially in the genera Pristina and Pristinella, especially considering the small number of locations that have been sampled. The collections include approximately 12 new species of rhyacodrilines (Tubificidae), Dero, Pristina, Pristinella (Naididae), Brinkhurstia (Alluroididae), Hemienchytraeus and Aspidodrilus (Enchytraeidae). This is the first record of Aspidodrilus from outside of Africa and the first new record since 1952.     

Macroecology of Aquatic Insects: A Quantitative Analysis of Taxonomic Richness and Composition in the Andes Mountains of Northern Ecuador

To determine the spatial dynamics of Neotropical lotic insect species, specimens were collected from 41 streams on the eastern and western flanks of the Andes Mountains in Ecuador. We examined the manner in which taxonomic richness and composition differed with elevation, latitude, and versant. Statistical analyses were limited to 5 families (Ephemeroptera: Baetidae, Leptohyphidae, Oligoneuriidae; Heteroptera: Naucoridae; Coleoptera: Elmidae), comprising 32 genera and 85 species, for which identifications or morphospecies assignments were reliable. Assessment of taxonomic diversity was based on the richness of genera and species at each site. In addition, each site was characterized by species richness within each of 4 families with more than 10 species. The effects of versant and transect on composition and structure were family‐specific. Mean site differences between versants in elmid richness at generic and specific levels depended on transect. Only baetid richness was affected by versant and transect in a consistent manner. Variation among sites in composition based on all genera and species was captured using multidimensional scaling (MDS). Subsequent MANOVAs based on scores from MDS axes revealed that differences between versants were much stronger in the southern transect, although transects differed from each other at specific and generic levels. A Jaccard's similarity matrix was computed for each family to reflect the spatial organization of taxonomic composition. Mesogeographic patterns of species composition for each of the four families were correlated (Mantel analysis) at both the regional level and at the level of the entire study area. At the regional level, the only pair of families to exhibit correlated patterns of species composition was elmids and naucorids in the southwestern region. The pattern of species composition for each family was correlated with the patterns for one or more other families at the level of the entire study area. Thus, spatial dynamics of species composition was similar for the families examined, suggesting that the Andes exert a consistent influence on species distributions within families, regardless of ordinal affiliation. At a local scale, however, the way in which taxonomic composition changed with latitude and versant was family‐specific. Mayflies, the most vagile of the taxa studied, had the highest percentage of species overlap between versants. Of three genera of Naucoridae collected, species of Ambrysus, of probable Mexican origin, were found only on the eastern versant, corroborating other evidence that the genus is recent in South America. Moreover, dispersion by Ambrysus across the Andes Mountains may not have occurred, as it has for Cryphocricos and Limnocoris, which are of probable South American origin.     

Lacustrine profundal meiobenthos as an environmental indicator

Organic loading and eutrophy is indicated at profundal depths by large numbers of resting copepodid stages of cyclopoid copepods, by the occurrence of the naidid oligochaete species, Amphichaeta leydigii and Specaria josinae, and the harpacticoid species Canthocamptus staphylinus, and by a low meiobenthos/macrobenthos biomass ratio. An oligotrophic environment is indicated by the occurrence of the aeolosomatid oligochaetes Aeolosoma quaternarium, A. hemprichi and Rheomorpha neiswestnovae, the naidid oligochaete Chaetogaster langi and the harpacticoid species Attheyella crassa and a high meiobenthos/macrobenthos biomass ratio.