Lysosomes and sulfide-oxidizing bodies in the bacteriocytes of Lucina pectinata, a cytochemical and microanalysis approach

Lucina pectinata is a large tropical clam living deeply burrowed in the black, reducing mud of mangrove swamps. It is known to possess hemoglobin in the cytoplasmic areas of its bacteriocytes, which harbor sulfide-oxidizing bacteria. The bacteriocytes also possess lysosome-like microbodies containing either membrane whorls or electron-dense granules in which free heme compounds have been identified. The cytochemical detection of acid phosphatase and arylsulphatase through EDX (energy-dispersive X-ray) microanalysis strongly suggests that the bacteriocytes of L. pectinata contain, in fact, two different types of microbodies. Some of these (devoid of dense granules) possess a variable amount of lysosomal enzymes and occasionally a limited quantity of iron, which may result from a recycling process of hemoglobin. Their main function seems to be the digestion of a limited proportion of symbiotic bacteria. They represent genuine secondary lysosomes with a functionally acidic pH. The second type of microbodies is characterized by dense granules containing sulfur and iron hemes but no lysosomal enzymes. Their sulfide-oxidizing activity was substantiated by benzyl viologen assay, with Na₂S as a substrate. These microbodies appear to be similar to the sulfide-oxidizing bodies (SOBs) described in the bacteriocytes of other bivalve species with symbiotic thioautotrophic bacteria; however, their sulfide-oxidizing activity appears to be non-enzymatic. They are discrete organelles, characterized by a functionally basic pH and pseudoperoxidasic activity, and have been termed SOBs. Therefore, the bacteriocytes of L. pectinata possess at the same time functional lysosomes and functional SOBs. Received: 17 August 2000 / Accepted: 20 December 2000     

Ultrastructure of the gill of the hydrothermal vent bivalveCalyptogena magnifica,with a discussion of its nutrition

Calyptogena magnifica Boss and Turner, 1980, a new Vesicomyidae found during the Galápagos expedition in hydrothermal vents of the East Pacific Rise, was collected in the same Rise at 21°N during the Oasis expedition (March 1982), and samples of the gill were fixed for ultrastructural observations. The large size and structure of the gill indicate that this is the organ mainly involved in the nutritional processes ofC. magnifica. Despite the classic structural appearance of the external cilia of its gill, and an obvious production of mucus,C. magnifica is not a filter-feeder, as it does not use filtering processes to provide its major source of nutrition. Negligible particulate transfer is evidenced by reduction of the ciliary groove, of the labial palps and of the digestive tube, as well as by the absence of mucous strings. Histological and ultrastructural observations endorse the hypothesis that endocellular chemoautotrophic bacteria play an important role in the nutrition of the clam. Except for a superficial zone of ciliated cells, most of the gill tissue is comprised of cells which appear to be bacteriocytes, and which are perfectly integrated into the gill tissue and contain abundant and normally reproducing bacteria. The differences observed in the structure of the bacteriocytes suggest a cyclic process of their colonization by bacteria, their possible resorption, and their replacement by new bacteria-infected cells. Energetic substrates (sulfides and organic molecules) are probably directly absorbed by the bacteriocytes through the microvilli of the epithelial cells. Abundant fingerprint-like mitochondria in ciliate cells attest to a particularly high metabolic activity, perhaps related to active biosynthesis.     

Phenotypic variations in the gills of the symbiont-containing bivalve Lucinoma aequizonata

The marine bivalve Lucinoma aequizonata (Lucinidae) maintains a population of sulfide-oxidizing chemoautotrophic bacteria in its gill tissue. These are housed in large numbers intracellularly in specialized host cells, termed bacteriocytes. In a natural population of L. aequizonata, striking variations of the gill colors occur, ranging from yellow to grey, brown and black. The aim of the present study was to investigate how this phenomenon relates to the physiology and numbers of the symbiont population. Our results show that in aquarium-maintained animals, black gills contained fewer numbers of bacteria as well as lower concentrations of sulfur and total protein. Nitrate respiration was stimulated by sulfide (but not by thiosulfate) 33-fold in homogenates of black gills and threefold in yellow gill homogenates. The total rates of sulfide-stimulated nitrate respiration were the same. Oxygen respiration could be measured in animals with yellow gills but not in animals with black gills. The cumulative data suggest that black-gilled clams maintained in the aquarium represent a starvation state. When collected from their natural habitat black gills contain the same number of bacteria as yellow gills. Also, no significant difference in glycogen concentrations of the host tissues was observed. Therefore, starvation is unlikely the cause of black gill color in a natural population. Alternative sources of nutrition to sulfur-based metabolism are discussed. Denaturing gradient gel electrophoresis (DGGE) performed on the different gill tissues, as well as on isolated symbionts, resulted in a single gill symbiont amplification product, the sequence of which is identical to published data. These findings provide molecular evidence that one dominant phylotype is present in the morphologically different gill tissues. Nevertheless, the presence of other phylotypes cannot formally be excluded. The implications of this study are that the gill of L. aequizonata is a highly dynamic organ which lends itself to more detailed studies regarding the molecular and cellular processes underlying nutrient transfer, regulation of bacterial numbers and host–symbiont communication. Received: 1 September 1999 / Accepted: 1 February 2000     

Elemental composition of the hydrothermal vent clam Calyptogena magnifica from the East Pacific Rise

Specimens of the hydrothermal vent clam Calyptogena magnifica were collected at a depth of 2 600 m from 21° N on the East Pacific Rise in spring, 1982. The elemental composition was determined for the total soft tissues, individual organs, and the shell. The soft parts contained high concentrations of a number of trace metals. Iron, copper, and zinc, at 760, 148 and 2 152 g g^-1 dry weight, respectively, were present at the highest concentrations. Silver and antimony exhibited the greatest enrichments when their concentrations in C. magnifica were compared with those measured in a shallow-water marine mussel, Mytilus edulis. In contrast to the soft parts, the shell was not a site of metal accumulation or deposition. Within the soft parts; gills, kidney, pericardium, and mantle were involved in the concentration of various metals; the degree of concentration varied depending on the particular metal under consideration. C. magnifica possesses metal body burdens which are ordinarily considered to be very high and potentially toxic in other species.     

Spawning and development of three coral-associated Lithophaga species in the Red Sea

Lithophaga date mussels from three species (L. lessepsiana, L. simplex and L. purpurea) were removed from their stony coral hosts in the Red sea at Eilat, Israel. Spawning, observed in the laboratory on several occasions during 1987–1988, appeared to be closely tied to lunar periods, occurring primarily during the last quarter and the new moon. Embryonic and larval development was typical of that described for other mytilids and, except for pigmentation differences, which could be discerned during embryogenesis, the developmental stages of the three species were indistinguishable. Development to the pediveliger stage took 3 to 4 wk in standard culture conditions, but raising the temperature to 27.5 °C increased the growth rate of larvae of L. lessepsiana by as much as three-fold, so that the pediveliger stage was attained in 16 d. Larvae resulting from spawning by L. simplex adults removed from the coral Astreopora myriophthalma grew significantly faster in culture than larvac from adults removed from the coral Goniastrea pectinata (comparison of slopes, p<0.05). The latter individuals showed a 6-d growth plateau at the early umbone stage. Metamorphically competent larvae were capable of delaying metamorphosis for up to 4mo, which would allow an extended period for dispersion and would increase the chance of finding a suitable substratum in the natural environment.     

Deep distributions of oceanic cirripede larvae in the Sargasso Sea and surrounding North Atlantic Ocean

From deep ( 1000 m), multi-depth zooplankton samples taken in the Azores frontal region from November 1980 to June 1981, high numbers of the two larval forms ofLepas pectinata, the commonest lepadid cirripede in this area, were sampled. The nauplii and cyprid larvae are large and long-lived. Nauplii were sampled where they feed, in the upper 150 m, closely associated with the fluorescence maximum. The cyprids were mainly sampled between 300 and 400 m, with a sharp cut-off in numbers just below the distribution peak. This deep distribution is intriguing, as the cyprids, which were not migrating diurnally, are non-feeding. To complete their life cycle they must settle on floating debris at the surface, where very few were sampled. They are negatively buoyant and there is no apparent physical reason for this deep distribution. There was no obvious pattern of changes in individual length or length/dry weight distribution with depth, the population appeared homogeneous. There was also no accumulation of other zooplankton at the same depth, or evidence from gut contents of large carnivores that the distribution pattern was caused by selective predation. A close association exists betweenL. pectinata andSargassum spp. weed, the cyprids preferentially settling on small fragments of weathered weed, the supply of which may be enhanced after winter. During winter, the harsh environment at the surface of the northern Sargasso Sea may discourage cyprid settlement. The deep distribution may be an ontogenetic migration conferring some survival advantage and saving energy due to the lower temperature and oxygen levels at depth. If it is a behavioural mechanism, duration of the cyprid stage will be extended until a more clement time of year, allowing them to settle over a longer period. The cyprids of other species were also sampled at depth, so the phenomenon appears to be widespread. If subsequent research confirms these observations as an ontogenetic migration, our current understanding of the fouling behaviour of oceanic lepadid cirripedes would be considerably modified.     

The gill symbiont of the hydrothermal vent mussel Bathymodiolus thermophilus is a psychrophilic,chemoautotrophic, sulfur bacterium

Certain hydrothermal vent invertebrates, e.g. Riftia pachyptila and Calyptogena magnifica, are clearly established as harboring dense populations of chemoautotrophic sulfur bacteria in specialized tissues. By contrast, the physiological characteristics of the abundant intracellular gill symbiont of the vent mussel Bathymodiolus thermophilus have been questioned. The low activities of enzymes diagnostic for CO₂ fixation (Calvin cycle) and for sulfur-driven energy generation, as measured by other investigators, have been attributed to bacterial contamination of the gill surface. Based on research at the Galápagos Rift hydrothermal vents in 1988 and subsequent laboratory experiments, the current study confirms that the B. thermophilus symbiont is a psychrophile for which thiosulfate and sulfide stimulate CO₂ fixation. It strongly indicates that the symbiont is a chemoautotroph by establishing the following: (1) Sulfide and thiosulfate can stimulate CO₂ fixation by partially purified symbionts by up to 43-fold and 120-fold, respectively; (2) the ribulose-1,5-bisphosphate carboxylase/oxygenase activity of the symbiont is sufficient to account for its sulfide- or thiosulfate-stimulated CO₂ incorporation; (3) the symbiont's molar growth yield on thiosulfate, as judged by CO₂ incorporation, is indistinguishable from that of free-living chemoautotrophs. Due to the high protein-degrading activity of B. thermophilus gill lysate, it is also suggested that host lysis of symbionts plays a more important role in the nutrition of the vent mussel than in R. pachyptila or C. magnifica, for which no comparable protein-degrading activity was found.     

Hematological studies on some Nile fishes,Tilapia nilotica,Lates niloticus,and Labeo niloticus

Hemoglobin content of 3 Nile fishes — Tilapia nilotica Linnaeus, Lates niloticus Linnaeus and Labeo niloticus Forskal— were examined. Clear differences in hemoglobin and hematocrit values were observed, and seasonal variation in the hemoglobin content of Lates niloticus and Labeo niloticus were noticed. There is a correlation between hematocrit value and hemoglobin content of the 3 species examined. The chloride content is higher in the species with lower hemoglobin content.     

Analysis of fungal fruiting patterns at the Dawyck Botanic Garden

Since 1994 data on fungal fruiting have been gathered in the Dawyck Botanic Garden outside Edinburgh, including that for eight plots within a 7.5 ha wood, which has been left unmanaged in order to investigate relationships amongst fungal succession, habitat characteristics and local weather patterns. The climatic data are provided by a small meteorological station situated on site. To handle nearly 4000 entries of data so far collected a computer database was constructed. This assists querying of the information so that changes in fungal communities and patterns of succession can be observed, and the relevant data extracted for further analysis. To aid in this analysis, and to give the user full control over editing, updating and querying the database, a special computer program was compiled.Analysis of the database records revealed that the currently known Dawyck fungal community comprises over 620 different taxa, and the species list increases every year. Although some species of mycorrhizal fungi showed high specificity towards their preferred host either at species (e.g. Lactarius blennius with beech, Lactarius quietus with oak, Russula fellea and Russula mairei with beech), or at genus level (e.g. Suillus spp. with Pinaceae), some other common fungi (e.g. of the genus Amanita) were found to have very wide host ranges. In addition, unusual mycorrhizal associations have also been recorded. A lime tree (Tilia), which is supposedly primarily endomycorrhizal, has been found regularly associated with Russula cyanoxantha, Russula parazurea, Russula pectinata and Inocybe fastigiata. Many common Scottish fungi were found to be associated with introduced exotic trees, which is illustrated on the example of Pseudotsuga menziesii (Douglas Fir).The software created utilises the Microsoft Office suite of programs. Microsoft Access was used to construct the database, and Microsoft Excel was used for statistical investigations. The software was programmed using Microsoft Visual Basic, and provided a means of extracting the necessary data stored in Access and controlling the formulae used in Excel for their analysis. The recorded seasonal patterns of fungal fruiting was successfully described by a simulation mathematical model comprising differential and algebraic equations. This model may now be used as a submodel within more complex models describing terrestrial ecosystem processes. Examples of model simulations and sensitivity analysis are given and implications for future research discussed.     

A survey of translocation in laminariales (Phaeophyceae)

A survey of translocation of photoassimilates in 13 genera of Laminariales is presented. All showed long-distance transport of ¹⁴C-labeled products from mature source tissue to meristematic sinks (haptera and intercalary growing regions). In plants with several laminae forming one frond, older laminae may provide assimilates for the growth of younger ones, and in Macrocystis spp., where fronds of different ages and developmental stage arise from a common holdfast, mature fronds initiate and support new fronds. Translocation velocities vary from species to species but are in the range of 55 to 570 mm/h. The results strongly support the hypothesis that Laminariales in general have an effective translocation system, on which their thallus growth depends.     

Coexistence of nine anemonefish species: differential host and habitat utilization, size and recruitment

The region of Madang, Papua New Guinea, has the highest reported species diversity of both anemonefishes (nine species) and their host anemones (ten species). To determine which factors may allow so many anemonefish species to coexist at this location, we studied their patterns of distribution, abundance, and recruitment. Population surveys at three replicate reef sites within four zones situated at varying distances from the mainland (nearshore, mid-lagoon, outer barrier, and offshore) indicated that each species of host anemone and anemonefish lived within a particular range of zones. Each species of anemonefish lived primarily with one species of host. Anemonefish species that lived with the same host species usually had different distribution patterns among zones (e.g., Amphiprion percula occupied Heteractis magnifica in nearshore zones, while A. perideraion occupied H. magnifica in offshore zones). Monitoring of natural populations showed that there were few changes (losses or recruitment) in the number or species of fishes associated with each individual anemone over periods ranging from 3 to 9 months. Recruitment was monitored on anemones with and without residents (resident fishes were removed) within each of three zones (nearshore, mid-lagoon, outer barrier). Significantly more anemonefishes recruited to anemones without resident fishes than to anemones with resident fishes. Each anemonefish species recruited to particular host species and zones. The distribution and abundance of the recruits of each fish species among zones were positively correlated with the distribution and abundance of resident fishes in the benthic habitat. This suggests that the spatial patterns of recruitment among zones strongly determined the distribution and abundance patterns of the benthic populations, and they were not the result of post-recruitment mortality or movement. Coexistence of the nine anemonefish species on the limited anemone resource was considered possible because of niche differentiation (i.e., differences in host and habitat utilization among zones), and the ability of two small species (i.e., Amphiprion sandaracinos and A. leucokranos) to cohabit individual anemones with other anemonefish species. Received: 29 July 1999 / Accepted: 1 September 2000     

Respiratory pigments of Yoldia eightsi, an Antarctic bivalve

Yoldia eightsi, an infaunal protobranch bivalve inhabiting the soft sediments of the Antarctic Sea, contains hemocyanin in its hemolymph and an intracellular hemoglobin in its gills. The concentration of hemocyanin in the hemolymph of Y. eightsi is very low, but the presence of this protein was documented by transmission electron microscopy after negative staining. The structure of Y. eightsi hemocyanin appears to be remarkably similar to that found in Gastropoda. The intracellular gill hemoglobin can be recovered after osmotic shock of the tissue and is isolated as a monomeric form, the molecular weight of which is 15.7 ± 0.4 kDa, as detected by FPLC gel-filtration experiments and SDS-PAGE. Isoelectric focusing experiments suggest the presence of a single native component with pI 6.6. Aggregation products accumulate during exposure to air and are resistant to SDS and β-mercaptoethanol treatment. This process can be avoided by maintaining the sample under a CO atmosphere. The oxygen dissociation curves, determined at pH 7.0 and 7 °C, are slightly sigmoidal, giving a Hill constant value of 1.35, probably related to the formation of aggregation products. The affinity for oxygen is moderately high, half saturation occurring at 3.15 mmHg. Received: 6 May 1997 / Accepted: 22 December 1997     

Vertical distribution of early and competent larvae of Concholepas concholepas in two systems of Chilean inland seas

The transport of marine invertebrate larvae is strongly influenced by their distribution in the water column, which could be affected by the biological features of the larvae and environmental variables. Larvae can modify their swimming behavior throughout their planktonic cycle, thereby changing the observed distributional patterns. This ability, coupled with oceanographic features, could induce landward or seaward transport. We studied the vertical distribution of C. concholepas larval stages in two differently stratified systems in Chilean inland seas; Refugio Channel (a strongly stratified channel, where previously has been described as a frontal system) and Guaitecas (a gently stratified system). Combinations of 12–24 h larval collection experiments were done simultaneously with fixed temperature and salinity profiles; meteorological data were also obtained. The results suggest that both salinity and day period influence the C. concholepas distributional patterns in the water column. Early veliger and competent larvae are concentrated in different parts of the water column, probably related to their transport capacities. The upper layer of the water column at the Refugio site, unlike the Guaitecas site, showed a stratified regime, which could affect larval density and larval length between the two sites. Finally, our results suggest that Refugio may be a sink habitat for C. concholepas.     

Methanotrophic symbiont location and fate of carbon incorporated from methane in a hydrocarbon seep mussel

Seep Mytilid Ia (SMIa), an undescribed mussel found at hydrocarbon seeps in the Gulf of Mexico, harbors intracellular methanotrophic symbionts. Two techniques were used to address the hypothesis that host digestion of symbionts is a significant mechanism of carbon transfer from symbiont to host in the SMIa association: lysosomal enzyme cytochemistry and ¹⁴C tissue autoradiography. Acid phosphatase activity was consistently localized in the Golgi apparatus and associated vesicles of gill cells, but was detected around bacteria in only three of approximately 50 bacteriocytes examined. These results indicate that the cellular equipment necessary for lysosomal digestion of symbionts is present in host bacteriocytes, but that acid phosphatase activity in symbiont vacuoles is rare at a given point in time. Tissue autoradiography was conducted with mussels collected in September 1992 to document carbon fixation by symbionts and follow the time course of transfer to host tissues. No asymbiotic host cell type showed a significant increase in relative grain density until at least 1 d after the end of incubation with ¹⁴C-methane. The ratio of label in the basal portion of bacteriocytes to total bacteriocyte label did not show a significant increase until 10 d after the end of the incubation period, indicating a slow increase of labeled carbon in the putative residual bodies, containing the remnants of lysosomal digestion. These results are consistent with the hypothesis that host digestion of symbionts is one route of nutrient acquisition in SMIa. Intracellular methanotrophic bacteria were found outside of the gill in SMIa juveniles, in mantle and foot epithelial tissues previously believed to be symbiont-free. These extra-gill symbionts and their host cells are morphologically similar to their gill counterparts and, like the gill symbionts, actively fix carbon from methane. Received: 29 March 1997 / Accepted: 12 May 1997     

A comparative study of the blood of four Tilapia species (Pisces)

The blood of 4 species of Tilapia has been examined. Mean erythrocyte count and size, hemoglobin content, and hematocrit value varied markedly among the species examined. A tendency towards a reciprocal relationship between the number of erythrocytes and their size was observed. In general, their is a correlation between the erythrocyte counts, hemoglobin content, and hematocrit value among the species examined.     

Life history of the nematode Theristus anoxybioticus from sublittoral muddy sediment at methane seepages in the northern Kattegat,Denmark

Field and laboratory studies from 1989 to 1994 of the nematode Theristus (Penzancia) anoxybioticus Jensen, 1995 inhabiting muddy sediment at methane seepages in the northern Kattegat, Denmark, have revealed a series of biological features of adults and juveniles not observed before in free-living marine nematodes. Reproductive adults were present in April and May only and inhabited the uppermost cm of sediment, while juveniles inhabited deep anoxic and sulphidic sediment layers during the remaining months of the year. This points to a downwards migration of newly hatched juveniles and an upwards migration during adulthood. Gut content analyses showed that adults feed mainly on benthic diatoms by swallowing whole cells while juveniles probably feed on bacteria; this differentiation in food clearly relates to the main bulk of potential food items in their respective microhabitats. The generation time is estimated to be one year, with a juvenile life span of about nine months, which is four to eight times longer than that found in related aerobic species of similar body size. Adults survived in oxic water but 50% died within 15 h in hypoxic water (5 to 6% atm sat), and all died within 12 h in anoxic water. Juveniles on the other hand died within 0.5 h in oxic water but survived anoxic water for at least 15 d. These results suggest that juveniles of T. anoxybioticus are adapted to an anaerobic life style.     

Feeding biology of the brackish-water oncholaimid nematode Adoncholaimus thalassophygas

The brackish-water inhabiting nematode Adoncholaimus thalassophygas (De Man, 1876) may be classified as an omnivore and predator according to the gut contents of the adults. A series of ¹⁴C experiments was conducted to investigate the feeding potential of labelled microorganisms and dissolved ¹⁴C glucose. The observed consistent failure to incorporate ¹⁴C from labelled microorganisms, and its consistent success with dissolved ¹⁴C glucose suggests that dissolved organic matter may be more important than microorganisms as a food source for his nematode. This corresponds also with direct observations of living worms which showed very little uptake of particulate food other than that originating from predation. Defecation and mean gulping rates were low and, therefore, ¹⁴C glucose uptake from solution can hardly be explained by the assumption that large amounts of water are swallowed and passing through the gut. By means of mucus secretion, A. thalassophygas forms compact agglutinations of detrital sediment which are sites of enhanced microbial activity. We suggest that hatched juveniles feed primarily on dissolved organic matter released by the intensive microbial activity within this habitat. Adults retain the ability to utilize dissolved organic matter, and supplement their diet by scavenging on carcasses and occasional predation. Thus, the biology of oncholaimid nematodes may be characterized by an intimate linkage to microbial metabolism, although they do not appear to feed directly upon microorganisms.     

Conidium ontogeny in marine hyphomycetous fungi: Asteromyces cruciatus and Zalerion maritimum

Developmental and ultrastructural aspects of conidium and conidiogenous cell formation in two species of marine hyphomycetes, Asteromyces cruciatus and Zalerion maritimum, are described, and a discussion of their accomodation in the experimental classification of Fungi Imperfecti, which is based on developmental characters, is presented. The blastic, basipetal and whorled pattern of conidiogenesis in A. cruciatus is compared to diagrammatic interpretations of conidium and conidiogenous cell formation in other hyphomycetous fungi, including Tritirachium album, Nodulisporium hinnuleum, Botrytis cinerea and Cladobotryum variospermum. Helicoid conidium formation in z. maritimum is also described, and it is suggested that the characters of this distinctive mode of blastic ontogeny define a separate category in the experimental classification which would include a large number of developmentally-related species.     

Beziehungen zwischen Körperbau und Lebensweise bei Blenniidae (Pisces) aus dem Roten Meer. III. Morphologie des Auges

The morphology of the eyes of 3 salariin Blenniidae have been investigated and compared: Salarias fasciatus (inhabiting the sublittoral), Istiblennius edentulus (eulittoral), and Alticus kirkii (supralittoral). An effective protection against desiccation in A. kirkii is offered by the very thick cornea conjunctiva. Extension of the visual field in this species is achieved by protrusion of the eyes from the head, by recession of the dermal pigment, and by a relatively large lense which allows better perception of marginal rays. The retina of I. edentulus is adapted for amphibious vision by the existence of many prominent swellings and folds, and of a central depression into which the lens can be with-drawn. Thereby, the stronger rays of light retraction present on land are focussed on the swellings, since the retina of the depression is under-developed. The cornea conjunctiva and propria of A. kirkii are separated, and thereby constitute an additional eye chamber. Presumably this cavity possesses a lower refractive index than the cornea or water, and thereby enables sharp vision in air, since the light-concentrating effect of the curved cornea is thus diminished. Additionally, the shape of the cornea propria can be varied, since it is centrally connected with the lens; by this means the eye chamber can be extended, enabling displacement of the tocal point of the light rays to correspond with the momentary residence. The adaptations described here for I. edentulus and for A. kirkii have so far not been reported in other amphibious vertebrates.

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Spatial distribution of the infaunal benthic communities of the Ría de Muros,north-west Spain

Benthic samples were collected at 13 stations in the Ría de Muros, north-west Spain. There are three major communities: (1) in the southern and outer part of the ría, an Amphiura filiformis—Thyone fusus community, inhabiting sandy sediments with low organic content; (2) in the middle part of the ría, and Amphiura chiajei—Maldane glebifex community, inhabiting muddy sediments that are not anoxic (sulphide not detected); (3) in Muros Bay and the inner part of the ría, a Spiochaetopterus costarum community, inhabiting anoxic sediments with high organic content. Biomass is very high (up to 46.2 g ash-free dry wt) in the first two communities. In the Spiochaetopterus costarum community, biomass and diversity are much lower, probably because of oxygen deficiency.