Oncholaimus Campylocercoides Research Articles

Do functional traits and biochemical biomarkers of the nematode Oncholaimus campylocercoides De Coninck and Schuurmans Stekhoven, 1933 affected by fluoranthene and polystyrene microplastics? Results from a microcosm bioassay and molecular modeling

The current experiment measured the multifaceted effects of polystyrene and fluoranthene, acting alone or in a mixture, on the meiobenthic nematode species Oncholaimus campylocercoides. This Oncholaimid was first experimentally selected from an entire nematode assemblage taken from the Jeddah coasts (Saudi Arabia). Several discernible changes were found in morphometry and functional traits after exposure to single and combined treatments. An increase in the activity of the biochemical biomarkers catalase and glutathione S-transferase was also observed following the exposure of males and gravid females of O. campylocercoides to 37.5 ng fluoranthene·g−1 dry weight (DW) and 62.5 mg polystyrene·kg−1 DW paralleled by a higher vulnerability of females. Moreover, the reproduction and feeding of this species were impaired, starting from 37.5 ng fluoranthene·g−1 and 62.5 mg polystyrene·kg−1, respectively. These results have been confirmed by good binding affinities and molecular interactions of fluoranthene and polystyrene with both GLD-3 and SDP receptors.

Meiobenthic nematode Oncholaimus campylocercoides as a model in laboratory studies: selection, culture, and fluorescence microscopy after exposure to phenanthrene and chrysene.

Numerous studies have focused on the response of meiofauna after exposure to polycyclic aromatic hydrocarbons (PAHs), but none has been devoted to their uptake into nematode body compartments. The present study monitored PAH uptake by Oncholaimus campylocercoides which was selected after 40 days in the laboratory through original protocols from natural sediments collected in the Old Harbor of Bizerte, Tunisia. To achieve the mono-species level, a grain size magnification was applied by gradually adding a biosubstrate made from either the crushed shells of Mytilus galloprovincialis or minced leaves of Posidonia oceanica. After selection, O. campylocercoides was cultured and fed with earthworm powder (560 mg.l-1). Thereafter, it was exposed for 3 weeks to phenanthrene and chrysene (38, 116, and 348 ppb). Fluorescence microscopy revealed higher intensities of PAHs at the spicules, mouths, and pharynges compared with the other organs considered. Moreover, the buccal fluorescence showed a significant correlation with that measured in the biosubstrate made with shells of M. galloprovincialis.

Do presence of gray shrimp Crangon crangon larvae influence meiobenthic features? Assessment with a focus on traits of nematodes.

In this study, a microcosm experiment was conducted for 30days to assess the impact of the presence of juvenile gray shrimp Crangon crangon on meiofauna. The results suggested that juvenile shrimp had a significant negative impact on the abundance of nematodes and copepods, but no effect on polychaetes. Moreover, nematodes showed a significant decline in individual weight. The collected nematodes were taxonomically identified and assigned to five functional traits: shapes of the tail and amphid, life history, feeding types, and adult length. The nematode traits were affected by the number of shrimp introduced, and descriptors followed normal or inversed bell-shaped curves. When no shrimp were present, the nematofauna had a higher species richness compared with treatments of 4, 8, and 12 shrimp. Bell-shaped curve patterns were common in relation to the two phases of feeding for C. crangon. During the first phase, C. crangon consumed the nematode species Oncholaimus campylocercoides; thereafter, shrimp fed mostly on the nematode Anticoma eberthi and copepods.

Migratory behavior of free-living marine nematodes surrounded by sediments experimentally contaminated by mixtures of polycyclic aromatic hydrocarbons

An experiment was conducted using microcosms with connected sedimentary compartments to assess the effects of three polycyclic aromatic hydrocarbons (PAHs) on free-living marine nematodes from Bizerte Bay (northeastern Tunisia). Over 30 days, the nematofauna were exposed to four treatment sediments, including one with chrysene (150 ppb), chrysene (150 ppb) plus fluoranthene (75 ppb), chrysene (150 ppb) and phenanthrene (15 ppb), and an uncontaminated reference. Numerical and qualitative community simplifications occurred in contaminated sediments relative to the reference. The diversity of nematodes differed based on hydrocarbon combinations. Multivariate analyses revealed communities in contaminated compartments differed from those in the initial natural sediment and reference. Differences in sensitivities of nematodes to hydrocarbons and in migratory abilities occurred. Rhabditis sp., Calamicrolaimus honestus, and Oncholaimus campylocercoides were in all compartments and classified as tolerant to PAHs. Nematodes observed only in control compartments, including Parasphaerolaimus paradoxus, Encheliidae (sp.), Trichotheristus mirabilis, and Theristus pertenuis were considered sensitive.

Effects of increasing levels of pharmaceutical penicillin G contamination on structure of free living nematode communities in experimental microcosms

A microcosm experiment was conducted to examine the effects of the pharmaceutical (penicillin G) on free living nematode communities of a Tunisian coastal zone (South-Western Mediterranean Sea). Sediments were contaminated with five penicillin G dose [D1 (3mgL−1), D2 (30mgL−1), D3 (300mgL−1), D4 (600mgL−1), D5 (700mgL−1)], and effects were examined after 30 days. Results showed significant differences between nematode assemblages from undisturbed controls and those from penicillin G treatments. Most univariate measures, including diversity (H′), species richness (d), equitability (J) and number of species (S) decreased significantly with increasing level of the antibiotic contamination. Results from multivariate analyses of the species abundance data demonstrated that responses of nematode species to the penicillin treatments were varied: Kraspedonema octogoniata and Paracomesoma dubium were eliminated at all the antibiotic doses tested and seemed to be intolerant species to penicillin G contamination; Oncholaimus campylocercoides although survived even the highest dose D5, showed definite reduction in its abundance and may be classified as “opportunistic” species at this dose, whereas, Nannolaimoides decoratus which showed a positive response with an increase in density even at highest concentration of contaminant, seems to be “penicillin G resistant” species.

Stress-induced selection of a single species from an entire meiobenthic nematode assemblage: is this possible using iron enrichment and does pre-exposure affect the ease of the process?

The present work proposes a new experimental design using iron enrichment to select a single species from an entire meiobenthic nematode community. The high diversity of nematodes makes it impossible to study their responses to stressors in terms of chemical biomarkers, as this kind of research demands a single species as a form of biological material. Two assemblages, one from a control site (Canal) and one from an iron-contaminated site (Fouledh) in Bizerte Lagoon (Tunisia), were used to validate the protocol and to study whether pre-exposure to iron could affect the ease of the selection process. Analyses of variance suggested that the abundance and species number of nematodes from the Canal and Fouledh sites decreased discernibly with exposure to iron. Multivariate analyses performed on nematode abundances revealed that Oncholaimus campylocercoides, Sabatieria granifer, Sabatieria punctata and Theristus flevensis were the most tolerant species, probably due to their deposit-feeding behaviour. Species with a restricted feeding spectrum showed a low tolerance potential. At the end of the experiment, O. campylocercoides became the unique component of the nematode community from both sites. The complete alteration of the community was achieved with more difficulty for nematodes from Fouledh than for those from Canal. The identity of the selected species was confirmed at both the morphological levels and by sequencing the nuclear small subunit ribosomal DNA (18S rDNA).

Responses of a free-living benthic marine nematode community to bioremediation of a PAH mixture.

The objectives of this study were (1) to assess the responses of benthic nematodes to a polycyclic aromatic hydrocarbon (PAH) contamination and (2) to test bioremediation techniques for their efficiency in PAH degradation and their effects on nematodes. Sediments with their natural nematofauna communities from Bizerte lagoon (Tunisia) were subjected to a PAH mixture (100 ppm) of phenanthrene, fluoranthene, and pyrene during 30 days. Nematode abundance and diversity significantly decreased, and the taxonomic structure was altered. Results from multivariate analyses of the species abundance data revealed that PAH treatments were significantly different from the control. Spirinia parasitifera became the dominant species (70 % relative abundance) and appeared to be an "opportunistic" species to PAH contamination while Oncholaimus campylocercoides and Neochromadora peocilosoma were strongly inhibited. Biostimulation (addition of mineral salt medium) and bioaugmentation (inoculation of a hydrocarbonoclastic bacterium) were used as bioremediation techniques. Bioremediation treatments enhanced degradation of all three PAHs, with up to 96 % degradation for phenanthrene resulting in a significant stimulation of nematode abundance relative to control microcosms. Nevertheless, these treatments, especially the biostimulation provoked a weak impact on the community structure and diversity index relative to the control microcosms suggesting their feasibility in biorestoration of contaminated sediments.

More than 35 years of studies on marine nematodes from Tunisia: a checklist of species and their distribution.

This work provides an inventory of species of free-living nematodes from Tunisian waters, based on samples collected from September 1977 until March 2013. Sediment samples were taken from 8 ecosystems: the Lagoons of Bizerte, Ghar El Melh and Bou Ghrara, Northern and Southern Lakes of Tunis, Old Harbor of Bizerte, Bizerte Bay and Ichkeul Lake. A total of 31 families, 133 genera, and 249 species of marine nematodes are currently known from Tunisia. The Xyalidae (with 30 species), Cyatholaimidae (25), Chromadoridae and Linhomoeidae (each 21) and Comesomatidae (20) are the richest families. Among them, Daptonema (with 9 species), Pomponema and Sabatieria (with 7 each) are the most species-rich genera. Some species were recovered only from a specific type of ecosystem. In detail, 109 species were recovered from lagoons, 56 species from sandy beaches and 6 species from mudflats. The 78 remaining species were widely distributed in Tunisian waters and four of these (Terschellingia longicaudata, Oncholaimus campylocercoides, Sabatieria pulchra and Theristus flevensis) were present in most of the 8 ecosystems studied.

Impacts of bioremediation schemes for the mitigation of a low-dose anthracene contamination on free-living marine benthic nematodes

A microcosm experiment was used to examine (1) the effects of different bioremediation schemes on degradation of anthracene and the structure of free-living marine nematodes in a lightly contaminated (4.5μgg(-1)) sediment from Bizerte lagoon and (2) the responses of the nematode community upon an artificial spiking of a low dose anthracene (1μgg(-1)). For that purpose sediment microcosms were incubated in laboratory for 40days. Bioremediation techniques decreased the anthracene contamination, and interestingly, biodegradation were more efficient when anthracene was artificial supplied into the sediment suggesting that the addition of bioavailable anthracene stimulated the bacterial community to adjust towards a PAH-degrading community. Spiking with this low dose of anthracene provoked significant changes in the nematode community structure and abundance, with the elimination of specific species such as Mesacanthion diplechma, the decrease of the dominant species Oncholaimus campylocercoides and the increase in abundance of opportunistic species such as Spirinia parasitifera. This would suggest a low tolerance of the nematode community despite the presence of a weak anthracene contamination in the sediment that could have allow dominance of an anthracene tolerant nematode species. Anthracene toxicity was alleviated in biostimulation treatments, leading to a strong increase in nematode abundance, concomitantly with changes in the nematode community structure; Prochromadorella neapolitana became the most abundant species.

Impacts of permethrin contamination on nematode density and diversity: A microcosm study on benthic meiofauna from a Mediterranean coastal lagoon

A microcosm experiment was used to examine the response of nematode in terms of density and diversity at different levels of permethrin contamination. The sediments were contaminated with three permethrin concentrations [P1: low (5 mg kg−1), P2: medium (25 mg kg−1) and P3: high (250 mg kg−1)] and the effects were evaluated after 30 days. The results from univariate and multivariate analyses showed significant differences between nematode assemblages from uncontaminated control and those from permethrin treatments. All univariate indices changed significantly at all the levels of permethrin contamination. In fact, the total nematode abundance (I), Shannon-Weaner index (H′), species richness (d), evenness (J′) and number of species (S) decreased significantly in all the contaminated microcosms. In addition, the results from multivariate analyses of the species abundance data demonstrated that permethrin affects the responses of nematode species. These significant modifications in nematode community structures with response to permethrin contamination were the consequences of a different specific tolerance to this pesticide. Thus, Araeolaimus bioculatus, Calomicrolaimus honestus, Oncholaimus campylocercoides and Theristus pertenuis characterized by increased abundances in all treated replicates, appeared to be “permethrin-resistant” species. Daptonema trabeculosum was eliminated in all the doses tested and seemed to be a very sensitive species to permethrin contamination.

Laboratory Study on Individual and Combined Effects of Cobalt- and Zinc-Spiked Sediment on Meiobenthic Nematodes

Free-living nematodes are the most abundant taxa among the meiobenthos and the predominant prey for bottom-feeding fishes. They are able to accumulate toxicants from sediments which explain their use in this study as possible tools in nutritional quality assessment of fishes. Nematodes from sediments of Ghar El Melh lagoon (Tunisia) were subjected to cobalt and/or zinc enrichment in a microcosm experiment for 30days. Three levels (low, medium, and high) of each treatment were used. Nematode abundance and diversity significantly decreased, and the taxonomic structure was altered. Results from multivariate analyses of the species abundance data revealed that all treatments were significantly different from the control. Both univariate and multivariate analyses of the data showed that the differential response occurred in all treatments, but the assemblages from microcosms contaminated with zinc alone were much more negatively affected compared with those exposed to cobalt alone. The presence of cobalt simultaneously with zinc seems to reduce its impact on nematode species composition. Such a result is suggestive of antagonistic interactions between these two metals. The responses of nematode species to the cobalt and zinc treatments were varied. Oncholaimellus mediterraneus, Oncholaimus campylocercoides, and Neochromadora trichophora were significantly affected with cobalt contamination but, they were not eliminated. Exposed to zinc, Hypodontolaimus colesi was eliminated and seemed to be an intolerant species versus this metal. Some of these species, "cobalt-sensitive" or "zinc-sensitive", were also affected by the metal combination even at low dose: O. mediterraneus, N. trichophora, and H. colesi. Differential sensitivity to cobalt and/or zinc may result in a subsequent competitive release of more tolerant species. A list of this kind of species was established to be used as a possible preventive tool versus contaminated fish. This was most evidently the case in Marylynnia stekhoveni and O. campylocercoides which increased at all doses of cobalt and zinc, respectively. These two resistant species have also the opportunity to dominate the nematode assemblage when the studied metals were added together. The level of health risk is probably higher for humans assimilating additional amount of cobalt and zinc, especially heavy smokers and/or patients using some medications including salts of these metals.

Experimental validation of the “relative volume of the pharyngeal lumen (RVPL)” of free-living nematodes as a biomonitoring index using sediment-associated metals and/or Diesel Fuel in microcosms

Since 1981, many numerical indices have been developed for meiobenthic nematodes but later discarded. To overcome problems and drawbacks in this type of indices, the “relative volume of the pharyngeal lumen (RVPL)” was first introduced in biological assessments in 2006 and it focused on the assumption that pollution is commonly expressed in terms of reduced food intake. This index, evaluated by using a morphometric approach, expresses the sucking potential of the pharynx. A series of microcosm experiments was performed in order to test the “relative volume of the pharyngeal lumen (RVPL)” as a biomonitoring index in the cases of 8 free-living nematode species (the deposit-feeders, Daptonema normandicum, Desmodora longiseta, and Leptonemella aphanothecae; the epistrate feeder, Xyala striata; the facultative predators, Oncholaimus campylocercoides and Oncholaimellus mediterraneus; and the predators Bathylaimus capacosus and Mesacanthion hirsutum). Each species was exposed to various concentrations of toxicants (nickel, copper, chromium and Diesel Fuel) used separately or in combinations. Our results showed that RVPL increased with increasing amount of toxicants, visibly caused by a systematic decrease in body volume with toxicants. Responses of all examined species suggested an additive toxicological action. Predators ( M. hirsutum and B. capacosus) showed the highest potential of adaptation because of their higher means of RVPL. Observations during this experiment highlight the importance of pharyngeal pumping and cuticular structure to tolerating elevated catabolic water loss under stress conditions. Thus the ability to osmoregulate is an important factor in overcoming food sedimentary toxicants. Based on all these findings, RVPL was found to be an accurate and precise index of monitoring the effect of metals and Diesel Fuel in marine ecosystems.

Examination of the bioindicator potential of Oncholaimus campylocercoides (Oncholaimidae, Nematoda) from Bizerte bay (Tunisia)

We have tested the use of the free-living nematode Oncholaimus campylocercoides as a bioindicator of petroleum pollution. This was analysed using a polluted-reference sampling strategy for one year in Bizerte bay (Tunisia). We noted a decrease in body dimensions together with many biometric anomalies when organisms are exposed to petroleum compounds. The three first moults and the egg-production phase of specimens of O. campylocercoides from the polluted site TRIC (Tunisian Refining Industries Company) took longer. The egg volume was negatively affected by pollution and the development from egg to juveniles J1 in one side and from juveniles J4 to males from TRIC beach on another side seems to be faster. Our results demonstrated that biometry, life cycle and fecundity are useful indices to biomonitor hydrocarbon pollution in marine ecosystems. Juveniles of the second and third stage and gravid females of O. campylocercoides seem to be the most appropriate bioindicators.

Oncholaimids (Nematoda, Oncholaimidae) from Tenerife (Canary Islands)

Four species of the family Oncholaimidae: Oncholaimellus calvadosicus De Man, 1890, Oncholaimus campylocercoides De Coninck & Stekhoven, 1933, O. skawensis Ditlevsen, 1921 and Viscosia glabra (Bastian, 1865) were recorded for the first time from the Canary Islands. Samples were collected in the intertidal and shallow subtidal sandy soft-bottoms from Los Abrigos and Los Cristianos, two localities situated on the eastern and western coast of Tenerife, respectively. Meristic and autoecological data of the studied species are presented.

Effects of Increasing Levels of Nickel Contamination on Structure of Offshore Nematode Communities in Experimental Microcosms

A microcosm experiment was used to examine the effects of nickel on offshore nematode communities of a Tunisian coastal zone (Southwestern Mediterranean Sea). Sediments were contaminated with three nickel concentrations [low (250 ppm), medium (550 ppm) and high (900 ppm)], and effects were examined after 30 days. Results showed significant differences between nematode assemblages from undisturbed controls and those from nickel treatments. Most univariates measures, including diversity and species richness, decreased significantly with increasing level of Ni contamination. Results from multivariate analyses of the species abundance data demonstrated that responses of nematode species to the nickel treatments were varied: Leptonemella aphanothecae was eliminated at all the nickel doses tested and seemed to be intolerant species to nickel contamination; Daptonema normandicum, Neochromadora trichophora and Odontophora armata which significantly increased at 550 ppm nickel concentration appeared to be "opportunistic" species at this dose whereas Oncholaimus campylocercoides and Bathylaimus capacosus which increased at all doses tested (250, 550 and 900 ppm) seemed to be "nickel-resistant" species.

Individual and combined effects of lead and zinc on a free-living marine nematode community: Results from microcosm experiments

A microcosm experiment was carried out to study the influence of lead and zinc, individually and in mixtures, on a free-living nematode community of a Tunisian lagoon. Three levels (low, medium and high) of each heavy metal were used, separately and in combinations, for sediment contamination and effects were examined after two months. Results from the multiple comparison tests showed significant differences between nematode assemblages from undisturbed controls and those from lead and/or zinc treatments. Most univariate measures, including diversity and species richness, decreased significantly in the treated microcosms. Results from multivariate analyses of the species abundance data revealed significant effects of both metals, separately or in mixtures, at all the doses tested on nematode assemblages. All treatments were significantly different from the control and from each other. Both univariate and multivariate analyses of the data showed that the differential response occurred in all treatments but the communities from microcosms contaminated with lead and zinc separately were much more strongly affected. This result is suggestive of antagonistic interactions between Pb and Zn. The responses of nematode species to the lead and zinc treatments were varied: Calomicrolaimus honestus was eliminated at all lead doses tested and seemed to be intolerant species to Pb contamination whereas Oncholaimus campylocercoides increased significantly at low and medium lead contamination, was eliminated in Pb-highly amended sediment treatments and increased at all zinc doses used. This species could be categorized as “opportunistic” at low and medium lead doses and “zinc-resistant” species. Hypodontolaimus colesi was eliminated in all replicates treated with Zn even at low concentration and appeared to be a very sensitive species to zinc contamination.

Individual and combined effects of cadmium and diesel on a nematode community in a laboratory microcosm experiment

A microcosm experiment was carried out to study the influence of cadmium and diesel, individually and in a mixture, on a free living nematode community of a Tunisian lagoon. Sediments were contaminated with cadmium that ranged in concentration from 0.54 to 1.40 mg Cd kg −1 (dry weight ( dw)), by diesel at 0.25 mg kg −1 ( dw), by a cadmium–diesel mixture (Cd at 1.40 mg kg −1+Diesel at 0.25 mg kg −1) and effects were examined after 90 days. Univariate analysis showed that all univariate indices did not change significantly neither at all the levels of cadmium contamination nor at 0.25 mg kg −1 ( dw) diesel concentration. But, at the cadmium–diesel mixture, significant differences were recorded between nematode assemblages from uncontaminated sediment control microcosm and those from cadmium–diesel mixture amended sediment treatments. Total nematode abundance ( I), mean individual weight ( bi), Shannon–Weaver index H′, species richness ( d), evenness ( J′) and number of species ( S) decreased significantly in microcosms contaminated with both cadmium and diesel. Results from multivariate analyses of the species abundance data demonstrated that responses of nematode species to the cadmium–diesel treatments were varied: Marylynnia stekhoveni, Calomicrolaimus honestus and Oncholaimellus mediterraneus were significantly affected at the cadmium–diesel contamination but they were not eliminated. These species could be categorized as “cadmium–diesel sensitive”. Enoploides sp. and Oncholaimus campylocercoides, characterized by increased abundances in cadmium–diesel amended sediment, seemed to be “cadmium–diesel resistant” species. All these species, “cadmium–diesel sensitive” or “cadmium–diesel resistant”, were not affected by either cadmium or diesel alone.

Sulphide tolerance of the marine nematode Oncholaimus campylocercoides-a result of internal sulphur formation?

The free-living, marine nematode Oncholaimus campylocercoides occurs in high abundance (up to 600 ind. 10 cm -2 ) at the fringe area of shallow-water hydrothermal vents off the Greek island of Milos in the Aegean Sea. It was found to have a sulphide tolerance (LT 50 ) of 4.5 d at 500 pM sulphide concentration and of 4 d at 1 mM sulphide. Light- and electron-microscopical inspections showed that the non-symbiotic O. campylocercoides, when exposed to sulphidic conditions, develops oily to viscous inclusions in the epidermis consisting of elemental sulphur in the form of S 8 -rings and polysulphur chains. The longer the exposure to sulphidic conditions, the more sulphur was formed, which disappeared after re-introduction of the nematodes in normoxic conditions for 12 h. Based on these results and on tolerance experiments with hydrogen sulphide, we suggest a model of sulphide metabolism in O. campylocercoides which could relate to its occurrence in sulphidic, hydrothermal sediments.

Benthic fauna of a shallow-water gaseohydrothermal vent area in the Aegean Sea (Milos, Greece)

The benthic community of shallow-water hydrothermal vents (10 m water depth) in a sandy bay on the south coast of Milos, Greece was studied. The macro- and meiofauna, as well as environmental factors such as temperature, salinity, sulfide concentration, grain size composition, ATP concentration, organic carbon, chlorophyll a and phaeopigments were evaluated. Samples were taken along transects from a seagrass meadow into a hydrothermally active area. Hydrothermally active spots were distinguished from the surrounding sediment by their black sediment coloration and an overlying white bacterial mat. For comparison, a control transect from a seagrass meadow into bare sand in a non-vent area in the same bay was studied. Overall, we found decreasing faunal diversity from the seagrass bed towards the hydrothermally active area. Along this gradient, four different zones could be distinguished. (a) The seagrass area had a relatively low diversity compared to the control stations, but much higher diversity than in the areas directly influenced by hydrothermal activity. (b) The border zone between the seagrass and the hydrothermal area, with moderate sulfide concentrations, pH values, seawater salinity and temperatures, was inhabited by an impoverished community dominated by the polychaete Capitella capitata, the nematodes Oncholaimus campylocercoides and Sabatieria sp., and an as yet undescribed chromadorid nematode (Chromadorina sp.). (c) The transition zone at the border of the hydrothermal area proper, with temperatures and sulfide concentrations higher than in the border zone, was characterized by high salinity (60‰) and low pH values. Along with the gastropod Cyclope neritea, single specimens of C. capitata and O. campylocercoides occurred. (d) The hydrothermal area proper, characterized by high sulfide concentrations, had high sediment temperatures and high salinities. The pH was as low as 6. Here, the gastropod Cyclope neritea was the only species occurring in considerable numbers. The gaseohydrothermal activity seems to largely exclude the surrounding fauna in a graded zonal pattern. The increasing hydrothermal stress entrains a change of the ambient biota towards a depauperate consortium of tolerant, opportunistic species, e.g. Capitella capitata (Polychaeta) and Oncholaimus campylocercoides (Nematoda). Although the hydrothermal area proper was characterized by a thick bacterial mat, the benthic community was characterized not by bacteria feeders but by scavenging species.