Oligochaeta in Spartina stems: the microdistribution of Enchytraeidae and Tubificidae in a salt marsh,Sapelo Island,USA

The distribution and abundance of Enchytraeidae and Tubificidae in and around Spartina alterniflora plants in a tidal salt marsh on Sapelo Island, Georgia, USA were studied using two different sampling techniques: wet funnel extraction and stem dissection. At least 80% of all worms inhabited leaf sheaths at the bases of S. alterniflora plants, and densities were low in sediment, root and surface debris samples. Oligochaete densities were dependent on the position within the marsh, the height on stems and the stage of sheath decay. Six predominant species were identified and included Marionina appendiculata, Marionina spartinae, Marionina waltersi, Marionina paludis, and Monopylephorus parvus. Individual species were distributed differently on stems and enchytraeids were more common than tubificids on standing-dead and further up S. alterniflora stems. Estimates of oligochaete densities in salt marsh habitats are increased dramatically when the numbers of worms on stems are considered. Possible advantages of the stem microhabitat are discussed in relation to the biology and ecology of oligochaetes.     

Phylogeny and origins of Enchytraeidae

Phylogenetic analyses of Achaetinae (Enchytraeidae: Oligochaeta) and other bisetate enchytraeids indicate that the achaetines include the earliest species of Enchytraeidae but that Achaetinae is not monophyletic. The earliest species of bisetate enchytraeid now extant are restricted to South America, Africa, and India. As this part of the enchytraeid lineage is ancestral to other enchytraeid taxa, it is suggested that Enchytraeidae may have arisen in South America or a contiguous Southern land mass.Less than 50% of the 11 genera of Enchytraeidae considered are supported by the results of these analyses as evolutionary or phylogenetic groups. Five are substantiated as monophyletic: Achaeta, Lumbricillus, Fridericia, Randidrilus, and Enchytronia. In addition to most of the achaetine genera, Marionina is shown to be in great need of revision. Its recognition is a continuing source of confusion to taxonomic resolution of Enchytraeidae.     

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.     

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.     

The occurrence of species of semi-aquatic Enchytraeidae (Oligochaeta) in Ireland

The Enchytraeidae are essentially terrestrial oligochaetes but many species have marked aquatic tendencies. Over two thirds of recorded Irish species were found in soils which were submerged or frequently flooded and 35% showed a distinct preference for these conditions. Relatively few species were living in soils subject to drought. Red blood was present in 28 species, all but one from soils with more than 55% water. Cognettia sphagnetorum and C. glandulosa developed red blood in very wet conditions. In a survey of Irish wetlands, samples were taken from bog, heath, marsh, fen, margins of lakes and rivers, and saltmarsh. The influence of various environmental parameters was determined using ordination techniques. Magnesium and pH were found to be the most important factors. A high level of magnesium distinguished coastal sites and pH 5.2 separated two clusters representing acid peat and marsh-fen-aquatic sites. Groups of indicator species characterized each of the three clusters. The ecological distribution of the indicator species is described, and their usefulness in classifying enchytraeid communities is discussed.     

The distribution of Oligochaeta on an exposed rocky shore in southeast Ireland

The distribution of oligochaetes was studied on a wave-exposed, granite shore at Carnsore Point, County Wexford. Habitats sampled were crevices and shallow surface cracks in rock, Lichina pygmaea turf, a Mytilus edulis bed, barnacles, Laurencia pennatifida turf, Corallina officinalis turf and pools. Lichina, mussels, barnacles greater than 8 mm in height, Laurencia and Corallina were sampled quantitatively.Three species of Tubificidae and nine species of Enchytraeidae were recorded including two new species described from this locality and five which are undescribed. Oligochaetes were present in all the habitats sampled but were rare in pools and numbers were low on barnacle-covered rock except where barnacles were more than 8 mm in height. The number of species increased with decreasing tide level and was highest in lower shore crevices. Highest densities were recorded from Corallina turf and in barnacles > 8 mm. Lumbricillus semifuscus was the most widespread species, occurring in crevices and mats at all tide levels; other species had more restricted distributions. Grania species were confined to Corallina turf. Only three species of oligochaete were recorded from cracks where they were almost the only fauna present. Species diversity and density were influenced by physical structure of the habitat, particularly the amount of retained sediment.The oligochaetes are members of a rich cryptofauna in habitats which provide them with organic matter and moisture and protect them from environmental extremes and wave damage and from predators during immersion. Reasons for the scarcity of oligochaete records from exposed rocky shores and the high proportion of new species in this study are discussed.     

Meiofauna of the profundal zone of the northern part of Lake Ladoga as an indicator of pollution

Benthic meiofauna was sampled at 19 stations, mainly in the northern part of Lake Ladoga, from depths between 13 and 199 m and from types of environment ranging from sheltered areas near pollution sources to less polluted open areas. About 80 taxa were identified, of these 70 to the species level. The greatest numbers of species were oligochaetes (24 species) and harpacticoids (8 species). Certain quantitative ratios of meiofauna were shown to be correlated with environmental data. The species of the oligochaete families Lumbriculidae and Aeolosomatidae and the harpacticoids as a collective group, excluding Canthocamptus staphylinus, were most clearly confined to the less eutrophied environments. The oligochaete species Amphichaeta leydigii, Dero digitata and Tubifex tubifex, the resting stages of Cyclopinae, and Eucyclops serrulatus among the Eucyclopinae were most clearly concentrated in the eutrophicated or polluted environments. The groups of Naididae, Cladocera and Eucyclopinae did not have much value as indicators. Shannon diversity index correlated positively with the total phosphorus content of the water, and number of species decreased with increasing depth. In general, it seems that the pollution tolerance of the meiofauna in a very large lake such as Lake Ladoga is high, presumably due to the effective mixing of water masses.     

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.     

Cold tolerance and dehydration in Enchytraeidae from Svalbard

When cooled in contact with moisture, eight species of arctic Enchytraeidae from Svalbard were killed by freezing within minutes or hours at −3 and −5 °C; an exception was Enchytraeus kincaidi which survived for up to 2 days. When the temperature approached 0 °C the enchytraeids apparently tried to escape from the moist soil. The supercooling capacity of the enchytraeids was relatively low, with mean supercooling points of −5 to −8 °C. In contrast, specimens of several species were extracted from soil cores that had been frozen in their intact state at −15 °C for up to 71 days. Compared to freezing in a moist environment, higher survival rates were obtained during cooling at freezing temperatures in dry soil. Survival was recorded in species kept at −3 °C for up to 35 days, and in some species kept at −6 °C for up to 17 days. Slow warming greatly increased survival rates at −6 °C . The results strongly suggest that arctic enchytraeids avoid freezing by dehydration at subzero temperatures. In agreement with this, weight losses of up to ca. 42% of fresh weight were recorded in Mesenchytraeus spp. and of up to 55% in Enchytraeus kincaidi at water vapour pressures above ice at −3 to −6 °C. All specimens survived dehydration under these conditions. Accepted: 12 December 1997     

The macroinvertebrate fauna of the drainage channels of the Gwent Levels, South Wales

SUMMARY. From a survey of the fauna at sixty sites in permanent drainage channels of the Gwent Levels during the spring of 1976 and at some of these sites (thirty-three) during the autumn, speeies and site associations were identified using classification and ordination techniques. Sites in some groups had identifiahly similar environmental characteristics and grouping seemed particularly related to the vegetational stage in the hydrosere, water How and the incidence of salt contamination from the Severn Estuary. Characteristics which were common within particular species groups and whieh, in part, explained their distributions were: adaptation to low oxygen concentrations, food preferences and dispersal mechanisms. Whilst there was some temporal stability in site groups between the spring and autumn surveys, such associations were not idenlical. The oxygen status of channels appeared a major factor determining faunal distribution. Twelve sites, representing a range of channel types, were sampled six times in a 12-month period. Far more species were colleeled within the water column and on plants than in the benthos. Several oligochaetes, e.g. Aulodrilus pluriseta, Dero digitata, Limnodrilus claparedeanus, were not recorded at most sites during the summer probably because of the SUStained lack of oxygen at these sites. Three contiguous reaches of channel were maintained during the period March—October 1977 with different plant cover (no Lemnu; complete Lemna cover: control with partial Lemna cover). The oxygen status of the reaches was different, that with complete Lemna cover being continuously anaerobic near the channel bed for several months, the control reach which had partial Lemna cover being anaerobic for a shorter period and that with no Lemna being only anaerobic at night. Whilst the distribution of macroinvertebrate species was prineipally related to this oxygen status, the control reach whieh was strueturally the most complex, having both submerged and floating plants, contained about twiee as many species as the other reaches during the summer (colleeted in the water column and on plants): benthic samples in this and the reach with complete Lemna cover contained only between zero and two species during the period May- September.     

Ultrastructure of the body wall of three species of Grania (Annelida: Clitellata: Enchytraeidae)

De Wit P., Erséus C. and Gustavsson L.M. 2011. Ultrastructure of the body wall of three species of Grania (Annelida: Clitellata: Enchytraeidae). —Acta Zoologica (Stockholm) 92 : 1–11. The body wall of three species of Grania, including the cuticle, epidermis and the musculature, are studied using TEM. The cuticle is similar to previously studied enchytraeids, with an orthogonal grid pattern of collagen fibers. This pattern is also seen in Crassiclitellata, which has been suggested as the sister taxon of Enchytraeidae. Variation of epicuticular and fiber zone patterns seen in Naididae (formerly Tubificidae and Naididae) seem to be lacking in Enchytraeidae. The fiber thickness, however, varies between Grania species and may be a phylogenetically informative character. The epidermis consists of supporting cells, secretory cells and sensory cells. Basal cells, typical for Crassiclitellata, were not observed. The clitellum of Grania seems to consist of two types of gland cells, which develop from regular epidermal tissue. It is possible that more cell types exist in different regions of the clitellum, however. The body wall musculature is arranged somewhat differently from that of closely related taxa; this refers to the reduction of circular and outer, triangular longitudinal muscle fibers, while the inner, ribbon‐shaped longitudinal muscle fibers are well‐developed. A search was conducted for the cause of the peculiar green coloration of Grania galbina De Wit and Erséus 2007, and it was concluded that neither cyanobacteria nor epidermal pigment granules were present in the fixed material.     

Oligochaete response to changes in water flow in the Dokka Delta,Lake Randsfjorden (Norway), caused by hydroelectric power development

The results reported here are part of a joint study on possible ecological effects in the delta area of Lake Randsfjorden, Southern Norway, caused by a reduction in water flow as a result of hydroelectric power development.From 1988 to 1990 the mean daily water flow decreased sharply and the total load of suspended matter was greatly reduced. During the same period there was an increase in oligochaete abundance from 6,096 ind. m^–2 in 1988 to 100,051 ind. m^–2 in 1989 and 154,345 ind. m^–2 in 1990. Piguetiella blanci was the dominant species in 1988 making up 48.5% of the total community. In 1989, Limnodrilus hoffmeisteri, Nais variabilis/Nais communis and enchytraeids were the most abundant oligochaetes, making up 29.3%, 26.1% and 28.2%, respectively of the total number. Reduced water flow from 1988 to 1989 and 1990 caused reduced disturbance of the bottom sediments and is considered the main reason for the change in species composition and abundances.     

Seasonal variation in the diel vertical distribution of the migratory alga Cryptomonas marssonii (Cryptophyceae) in a small,highly humic lake

The diel vertical distribution patterns of a migratory alga Cryptomonas marssonii in a small, steeply stratified humic lake were investigated during a summer season (10 diurnal experiments between May and September) using a close-interval Blakar-type sampler. The results indicate that the cells were phototactic; they were typically concentrated at the surface or subsurface during daylight, while in darkness the highest densities were recorded in deeper water, usually near the upper limit of anoxia. During a dense blue-green bloom in August the cells of C. marssonii were also concentrated by day into the same water layer, where oxygen was depleted. However, the cells seemed to avoid totally anoxic water. Because the vertical distribution pattern of C. marssonii had special diurnal and seasonal characteristics, care is needed when designing a sampling programme for a phytoplankton population dominated by this species.     

Inorganic carbon fluxes at the water–sediment interface in five littoral systems in Spain (southern Europe)

The assemblage of littoral oligochaetes in six crater lakes in Central Mexico, was studied throughout a yearly cycle. To establish species composition, richness, density and biomass, 14 localities were sampled in the lakes. A total of eight species belonging to the families Naididae (five species), Tubificidae (two species), and Enchytraeidae (one species) were found. The dominant species, Limnodrilus hoffmeisteri, contributed with up to 99% in both abundance and biomass. Sediment organic matter is the most important environmental variable explaining the differences in density and biomass. Seasonal (dry and rainy seasons) changes were not significant for density and biomass. Higher density and lower biomass values characterized these lakes in contrast to other tropical and subtropical lakes worldwide. The small size of the dominant species L. hoffmeisteri was recorded in all lakes and explained the low biomass recorded in the area of study. The correlation between L. hoffmeisteri and four other species (Dero (Dero) nivea, D. (D.) digitata, Nais variabilis and Tubifex tubifex) was negative. The naidid species were positively correlated (>0.5) to each other.     

The significance of Cognettia sphagnetorum(Enchytraeidae) on nitrogen availability and plant growth in wood ash-treated humus soil

A laboratory microcosm experiment was established to study whether the role of Cognettia sphagnetorum (Enchytraeidae) in affecting Scots pine (Pinus sylvestris) seedling growth is influenced by wood ash-amendment, i.e., neutralisation of the raw humus soil. Coniferous forest soil, inoculated with soil microbes and nematodes, was either treated with wood ash or left as ash-free control. Wood ash (corresponding to an amount of 5000 kg ha^–1) was either spread on the soil surface or mixed into the soil. Enchytraeid and pine seedling biomass, abundance of nematodes, and water leachable NH₄ ⁺-N and NO₃ ^–-N were measured 26 and 51 weeks after initiation of the experiment and root length and N concentration of needles were measured 51 weeks after initiation of the experiment. Wood ash when mixed into the soil, reduced the biomass of C. sphagnetorum. Nematodes were unaffected by the treatments. In the ash-free soils C. sphagnetorum had little influence on pine growth, but it did decrease root length and root to shoot ratio. In the absence of enchytraeids wood ash decreased pine biomass production and root length. However, the presence of enchytraeids in the ash-treated soil compensated the ash-induced negative effects on the pine performance. Enchytraeids increased and wood ash decreased water leachable NH₄ ⁺-N in the presence but not in the absence of enchytraeids, while water leachable NO₃ ^–-N was not affected by the treatments. It was concluded that C. sphagnetorum can be important in ensuring nutrient cycling and plant growth in situations when an ecosystem encounters disturbances.     

The distribution and ecology of littoral Oligochaeta and Aphanoneura (Annelida) of the Natural and Historical Reserve of Isla Martín García,Río de la Plata River,Argentina

The objective of this paper is to provide data on the distribution and ecology of oligochaetes and aphanoneurans occurring in the Río de la Plata, Argentina, including details regarding, their densities, faunal characteristics, species richness, diversity and equitability. Between 1995 and 1997, a total of five stations on the Río de la Plata were sampled on eight different occasions. The taxocenosis of Oligochaeta and Aphanoneura included 30 species belonging in six families: Naididae, Tubificidae, Narapidae, Enchytraeidae, Lumbricidae and Aeolosomatidae. The density varied between 44495 ind m^–2 and 57 ind m^–2. Species richness varied between 1 and 17, the diversity ranged from 0.07 to 2.55, and equitability varied between 0.08 and 0.64. According to dominance and frequency, the dominant, constant and widely distributed species were: Nais variabilis, Stephensoniana trivandrana, Amphichaeta leydigi, Chaetogaster diastrophus, Limnodrilus hoffmeisteri, Aulodrilus pigueti, Narapa bonettoi and the species of Enchytraeidae and Aeolosomatidae.     

Marine Oligochaeta from the Koster Area, West Coast of Sweden, with Descriptions of Two New Enchytraeid Species

A total of 28 species of marine Oligochaeta (belonging to the families Naididae, Tubificidae and Enchytraeidae) are reported from eulittoral and sublittoral sediment samples taken in the Koster area on the Swedish West Coast. Lumbricillus algensis sp.n. and Grania ovitheca sp.n. are described and their morphological and taxonomical relations to other enchytraeids are discussed. L. algensis is characterized mainly by its spermathecae, which project into segment VI, and by its very large penial bulbs. G. ovitheca possesses no dorsal setae, the ventral setae being present posterior to the clitellum only, and its spermathecae consist of a narrow duct and a large egg-shaped to oval ampulla, filling most of the coelom of segment V. Four sublittoral species, Limnodriloides barnardi (Tubificidae), Lumbricillus semifuscus, Marionina sublitoralis and Grania roscoffensis (all Enchytraeidae) are reported as new to the Swedish fauna.     

Molecular Barcoding of Aquatic Oligochaetes: Implications for Biomonitoring

Aquatic oligochaetes are well recognized bioindicators of quality of sediments and water in watercourses and lakes. However, the difficult taxonomic determination based on morphological features compromises their more common use in eco-diagnostic analyses. To overcome this limitation, we investigated molecular barcodes as identification tool for broad range of taxa of aquatic oligochaetes. We report 185 COI and 52 ITS2 rDNA sequences for specimens collected in Switzerland and belonging to the families Naididae, Lumbriculidae, Enchytraeidae and Lumbricidae. Phylogenetic analyses allowed distinguishing 41 lineages separated by more than 10 % divergence in COI sequences. The lineage distinction was confirmed by Automatic Barcode Gap Discovery (ABGD) method and by ITS2 data. Our results showed that morphological identification underestimates the oligochaete diversity. Only 26 of the lineages could be assigned to morphospecies, of which seven were sequenced for the first time. Several cryptic species were detected within common morphospecies. Many juvenile specimens that could not be assigned morphologically have found their home after genetic analysis. Our study showed that COI barcodes performed very well as species identifiers in aquatic oligochaetes. Their easy amplification and good taxonomic resolution might help promoting aquatic oligochaetes as bioindicators for next generation environmental DNA biomonitoring of aquatic ecosystems.     

A taxonomic and faunistic survey of the marine Tubificidae and Enchytraeidae (Oligochaeta) of Italy. Introduction and preliminary results

On the basis of material collected along the Ligurian, Tyrrhenian, Ionic and Adriatic coasts of Italy, a tentative list of 26 species (18 tubificids, 8 enchytraeids) of marine Oligochaeta is presented. Most of the species are new to science, and it can therefore be concluded that there is a very high diversity of oligochaetes in the Mediterranean Sea.     

Superficial and hyporheic oligochaete communities as indicators of pollution and water exchange in the River Moselle, France

Benthic oligochaetes were sampled on three occasions (June, August and October 1992) in the upper (0–10 cm) and hyporheic (35–45 cm depths) sediments at five sites of the River Moselle, from upstream of the town of Epinal to Velle-sur-Moselle. The first site (upstream from Epinal) is considered as unpolluted and the four remaining sites are polluted by industrial effluents. The most polluted stations were generally dominated by the pollution tolerant taxon Limnodrilus. Numbers of individuals of this taxon decreased at the less polluted last site in recovery zone, and were also scarce in the first unpolluted site. It is noteworthy that these tendencies were observed in both superficial and hyporheic substrates and to the greatest degree in hyporheic ones. At the unpolluted site, the hyporheic habitat is dominated by the groundwater species Propappus volki, Pristina spp., Pristinella spp. At the less polluted site (last site), the deep sediments are dominated by groundwater species and the Tubificidae without hair setae decrease from June to October. As a result of water exchange between superficial and subterranean waters, superficial substrates of the first and the last stations tend to be colonised by a high proportion of hyporheic species that suggests that flow is primarily from subterranean to superficial waters. The contrary is the case at other polluted stations which are characterised by the invasion of hyporheic substrates by the pollution tolerant superficial taxa Limnodrilus. This suggests that water flows from the river to the deeper groundwater. These two stations are located near drinking water plants which utilise groundwater, thus increasing the vulnerability of groundwater to surface contaminants.