Ecological assessment of soft bottom benthic communities in northern Spanish estuaries

The implementation of the Water Framework Directive requires the classification of the ecological quality status of benthic macroinvertebrates in costal and transitional waters. The uncertainty and complexity of this task have lead to the creation of specific Geographical Intercalibration Groups (GIGs) for the different ecoregions and types of water bodies established. In this framework, several metrics are under study in the United Kingdom, Spain and Denmark for North East Atlantic coastal waters (NEA-GIG). All of them include in their formulations the AMBI index. Nevertheless, few advances have been made in relation to transitional waters. This paper aims to test the suitability of those methodological procedures for their application in Northern Spanish estuaries. The results show evident divergences in classification of ecological status among methods, although the correlations of corresponding ecological quality ratios (EQRs) are good. Thus, we think that it may be necessary to modify the boundaries between each ecological status category and adjust the reference conditions for the variety of community-types existing in estuarine water bodies. However, some problems arise in the assessment of some naturally stressed communities. In this situation, we found an overall dominance of species tolerant to organic enrichment in all status categories and low range of variation of AMBI index. This aspect introduces some uncertainty in relation to the ability of this index to detect a deleterious effect in these estuarine communities naturally stressed. On the other hand, we found some sites that achieve good status according to the metrics used, even though the number of opportunistic species was high. The use of combined approaches that incorporate physicochemical condition of sediments would be a straightforward approach to reduce the risk of failing in the assignation of ecological status category.     

The use of a biotic index (Bentix) in assessing long-term effects of dumping coarse metalliferous waste on soft bottom benthic communities

The biotic index Bentix [Simboura, N., Zenetos, A., 2002. Benthic indicators to use in Ecological Quality classification of Mediterranean soft bottom marine ecosystems, including a new biotic index. Mediterr. Mar. Sci., 3 (2), 77–111] developed for the classification of ecological status of zoobenthic communities was tested in a Mediterranean coastal area (Aegean Sea, N. Evvoikos gulf, Greece) subject to coarse metalliferous waste discharge. The residues in the form of slag are the byproducts of the smelting procedure of a local ferronickel plant and are discharged in a permitted area 8 km from the shore. This practice adopted over the last 36 years has led to the formation of a thick underwater deposit of slag on the sea bottom. A decade of monitoring resulted to a long series of data on macrozoobenthos communities in the area. Application of the Bentix and other indices on the data has revealed the long-term impact of dumping on the benthic communities and the potential of the indices used to assess the community health. Among the indices tested, Bentix index succeeded in assessing the long-term trends of decline or recovery of the community health. The index proved efficient in detecting differences of the ecological status of the communities attributed to the transferring of the core of the dumping activity over the area studied, as validated by the geophysical data provided.     

Baseline study of soft bottom benthic assemblages in the Bay of Santander (Gulf of Biscay)

The Bay of Santander constitutes one of the most important estuarine areas in northern Spain (Gulf of Biscay) which has been significantly stressed by sewage discharges. However, an improvement in the environmental quality of the bay is expected once the new sewer system comes into operation. The evaluation of such an evolution needs to be contrasted with the current situation, established in specific baseline studies. For this purpose, a field study was carried out on the soft bottom assemblages of the estuary in spring of 1998. At each station, the benthic fauna abundance, grain size, organic matter, carbon, nitrogen, phosphorus and heavy metals concentrations were analysed. The community structure and the physical and chemical characteristics of the sediment reflected the higher impacts both near the points of discharges and in the low water exchange zones, whereas those areas showing higher tidal flows presented an acceptable quality. High heavy metal concentrations for Cd, Hg, Pb and Zn indicated contamination of industrial sources in several areas of the Bay. This information allows to establish a reference situation in order to improve the design of a specific monitoring program for this area.     

Ultrastructural study of neoplastic cells in Macoma balthica (Bivalvia) from the Gulf of Gdansk (Poland)

In the Baltic clam Macoma balthica from the Gulf of Gdansk, neoplasia is considered as a serious epizootic linked to bad environmental conditions and high levels of pollutants. Previous research was focused on the diagnosis, prevalence, seasonality and histopathological characteristic of the cancer. This study is focused on electron microscopy analyses describing cell ultrastructure abnormalities related to neoplasia. Examinations using the electron microscopy highlighted changes confined to anatomic ultrastructures, shapes and functions of neoplastic cells. The lobulated appearance of the nucleus, changes in cellular matrix and the occurrence of large granular cells with hyperchromatic nuclei, atypical Golgi structures and deterioration of rough endoplasmic reticulum manifested the disease. The presence of atypical mitochondria, free ribosomes and hypertrophic nuclei suggests the adaptation of neoplastic cells to increased mitotic activity, while the observed modification of cellular membranes may reflect functional changes connected to increased pinocytotic activity or intercellular transport. The cancer cells were found to appear in two types, abnormal round-shaped cells and spindle-shaped cells, both with increased frequencies of cell division. Round-shaped cells typical for disseminated neoplasia were observed in all affected bivalves, in a few cases co-occurring with abnormal spherical cells. Spindle-shaped cells containing some intracytoplasmic filaments, and with a tendency of the nuclei to be orientated as in a palisade were interpreted as myofibroblasts-like cells and were observed in five out of eleven clams diagnosed as neoplastic. This finding represents the first demonstration of phenotypic differences in the cell types co-occurring in animals diagnosed as neoplastic and by that suggests coexistence of two types of bivalve cancer, disseminated neoplasia and probable fibrosarcoma.     

Burrowing behaviour in the European lugworm Arenicola marina (Polychaeta: Arenicolidae)

A detailed analysis is presented of the mechanism of burrowing in Arenicola marina (L.). The following were recorded electronically, using further developments of the techniques introduced by Trueman: movement into the sand, internal pressure at various points in the body, external pressure in the surrounding sand. The results are interpreted in relation to direct visual observation of the burrowing process. The structural and functional specializations of the lugworm for burrowing and the factors concerned in the release and control of burrowing activity, are discussed.     

Ultrastructural investigation of the nuchal organs of Pygospio elegans (Polychaeta)

Summary The differentiation of the dorsal organs as well as the structure of the nuchal organs and their relation to the central nervous system in adult Pygospio elegans were studied by electron microscopy and compared to the nuchal organs of the larvae. The nuchal organs are represented by paired ciliary bands on the dorsal side of the first setiger, delimiting a median caruncle that is completely filled with epidermal and nervous tissue. They are composed of ciliated supporting cells and bipolar primary sensory cells constituting the nuchal ganglia, which are integrated into the brain. Microvillus-like processes of the ciliated cells give rise to a secondary covering layer over the sensory epithelium. The size of the nuchal organs is a sexually dimorphic feature.Dorsal organ formation is concomitant with the onset of sexual maturation in the male sex only. They appear as metameric ciliary bands on the dorsal side of the anterior body region and consist of ciliated cells accompanied by lateral accumulations of tubular gland cells. In the gametogenic segments they are structurally associated with the male genital pores and may be involved in reproduction. The results refute previous theories that dorsal organs are sensory and have a common origin to nuchal organs.Abbreviations ac anterior commissure of the brain - ace anterior circumesophageal connective - bb basal body - bl basal lamina - c cuticle - ca caruncle - cc ciliated cell - ci sensory cilium - co microvillar cover - d septate desmosome - db dorsal blood vessel - dn dorsal nerve cord - ea efferent axons - ec epidermal cell - eg elementary granules - g Golgi complex - i filamentous inclusion - lm longitudinal muscles - ly lysosome - mc motile cilia - mv microvillus - n neuron - ng nuchal ganglion - nn nuchal nerve - nu nucleus - oc olfactory chamber - pa palp - pc posterior commissure of the brain - pce posterior circumesophageal connective - rer rough endoplasmic reticulum - sI setiger I - sb sensory bulb - sc sensory cell - sd sensory dendrite - ser smooth endoplasmic reticulum - tf tonofilament bundle - v clear vesicles - za zonula adherens     

Macoma balthica in the White and Barents Seas: properties of a widespread marine hybrid swarm (Mollusca: Bivalvia)

A main molecular subdivision in the circumpolar Macoma balthica complex has been described between Atlantic and Pacific taxa. In NE Europe, the clams of the White and Barents Seas, however, show deviant genetic structures. Using allozyme and mitochondrial DNA data, we explore the hypothesis that these deviations result from hybridization between an Atlantic (M. b. rubra) and an invading Pacific (M. b. balthica) lineage. A practically pure Atlantic Macoma extends from France north to the Varanger Peninsula (NE Norway), whereas populations farther east have genetic compositions intermediate between true Atlantic and true Pacific. Admixture estimates range from 32 to 90% Pacific contribution, with a notable deviation in a nearly pure Atlantic outpost in the Mezen Bay (NE White Sea). The pattern of variation is not one of a simple collinear mixing however. Different characters exhibit different degrees of introgression, and the relative introgression varies regionally. Yet, there are practically no interlocus genotypic disequilibria between the diverged loci, which brings out the White Sea-Barents Sea M. balthica as the best-documented marine animal hybrid swarms so far, arisen through amalgamation of genomes previously isolated since pre-Pleistocene times. On top of the main admixture pattern, strong geographical structuring is also seen in characters unrelated to the principal systematic distinction. The persistence of the regional patterns indicates restricted gene flow at the present time, despite the high dispersal potential of the species. The causes of this structuring could be in a complex history of colonization events and features of local hydrography enhancing isolation and divergence of populations.     

Sublittoral soft bottom communities and diversity of Mejillones Bay in northern Chile (Humboldt Current upwelling system)

The macrozoobenthos of Mejillones Bay (23°S; Humboldt Current) was quantitatively investigated over a 7-year period from austral summer 1995/1996 to winter 2002. About 78 van Veen grab samples taken at six stations (5, 10, 20 m depth) provided the basis for the analysis of the distribution of 60 species and 28 families of benthic invertebrates, as well as of their abundance and biomass. Mean abundance (2,119 individuals m^?2) was in the same order compared to a previous investigation; mean biomass (966 g formalin wet mass m^?2), however, exceeded prior estimations mainly due to the dominance of the bivalve Aulacomya ater. About 43% of the taxa inhabited the complete depth range. Mean taxonomic Shannon diversity (H′, Log e) was 1.54 ± 0.58 with a maximum at 20 m (1.95 ± 0.33); evenness increased with depth. The fauna was numerically dominated by carnivorous gastropods, polychaetes and crustaceans (48%). About 15% of the species were suspensivorous, 13% sedimentivorous, 11% detritivorous, 7% omnivorous and 6% herbivorous. Cluster analyses showed a significant difference between the shallow and the deeper stations. Gammarid amphipods and the polychaete family Nephtyidae characterized the 5-m-zone, the molluscs Aulacomya ater, Mitrella unifasciata and gammarids the intermediate zone, while the gastropod Nassarius gayi and the polychaete family Nereidae were most prominent at the deeper stations. The communities of the three depth zones did not appear to be limited by hypoxia during non-El Niño conditions. Therefore, no typical change in community structure occurred during El Niño 1997–1998, in contrast to what was observed for deeper faunal assemblages and hypoxic bays elsewhere in the coastal Humboldt Current system.     

The control of burrowing and the migratory behaviour of Donax denticulatus (Bivalvia: Tellinacea)

Donax denticulatus burrows in a similar manner to other infaunal bivalves, but also reacts to stimulation by activity and so proved to be suitable material for experiments on the control of locomotion. Tactile responses of the foot initiate burrowing and pedal stretch receptors control the duration of each digging cycle. Prevention of adduction of the valves, removal of ligament or cardinal hinge teeth, had little effect on burrowing and it is suggested that part of the digging cycle is programmed within the nervous system without peripheral feedback. Experiments on migratory behaviour are discussed and indicate that no intrinsic mechanism is required for its control beyond the adaptation of behavioural responses, common to most bivalves, to different physical conditions.     

Free amino acids in the clam Macoma balthica L. (Bivalvia,Mollusca) from brackish waters of the southern Baltic Sea

Fourteen acidic and neutral free amino acids (FAA) were investigated in soft tissue of Macoma balthica from different depth zones of the Gulf of Gdansk (Baltic Sea) over a full seasonal cycle. The dry weight of the bivalves and physico-chemical parameters of overlying bottom water and surface sediments were measured simultaneously at each site. In the brackish waters of the Baltic, the main pool of FAA is composed of Ala, Gln, Arg, Gly and Orn which represent approximately 80% of the total. Compared to the full saline environments, the composition of FAA in the clams from the Baltic differs substantially. The differences can be attributed to the lower salinity of the Baltic. In the Baltic, Gly appears to play a most important role in regulating intracellular osmolarity in the clams, a function performed primarily by Tau in Atlantic and North Sea populations. Spatio-temporal variations of the FAA are affected by biotic and environmental parameters; their respective influence differs with the amino acids. The concentration of Arg depends on its uptake from the external medium. However, its level might be temporarily modified by stress-induced metabolic transformation (e.g. hydrolysis to Orn) caused by changes in the ambient environment. The concentration of Ala increases with depth, probably because of physiological adaptations of the animal to diminishing oxygen concentration through anaerobic glucose catabolism. Biosynthesis of Ala, similarly to Gln, in the shallower zone is generally related to the physiological state of an organism. The concentration of Gly is most likely regulated by internal mechanisms driven by gonadal development and reproduction.     

Integrated behaviour in the feeding phase of Caenorhabditis elegans (Nematoda)

Caenorhabditis elegans spends most of its life feeding. When feeding, it moves only short distances but intestinal contents are moved to and fro and thus mixed. The nematodes also defaecate and lay eggs during the feeding phase in bacterial culture. Defaecation is one of a series of events which causes the forward and backward movement of intestinal contents. These rhythmic movements, once initiated, appear to be controlled by a pacemaker and are unrelated to feeding rates. Oviposition is associated with movements of the genital tract and it can occur at any posture and does not influence the rate of other activities. After food-deprivation, C. elegans shows no measureable hunger response on being returned to bacteria. Experiments with excised parts of worms showed that isolated anterior ends will feed both in and out of bacteria and move forward and backward. Isolated posterior halves without a pharynx or circumpharyngeal commissure will lay eggs and defaecate.     

The benthic communities of the saline lakes Abijata and Shala (Ethiopia)

Lake Abijata lies in a shallow depression (maximum depth 8–9.5 m); the water is green with phytoplankton and it supports large fish and bird communities. Lake Shala lies in a deep caldera (maximum depth reputedly 260 m); phytoplankton is sparse and fish and bird communities scanty.Lakes Abijata and Shala, sampled in January, 1985, had conductivities of 14 000 and 21 000 microSiemens cm^-1 at 25 °C respectively, mainly due to high sodium, carbonate and chloride ions. Calcium concentrations are very low.The benthic fauna was studied with an Ekman grab to a depth of 8.5 m in Abijata and 15.5 m in Shala and was found to be dense in both lakes but varying greatly in composition at different depths. In Abijata the benthos consisted mainly of Ostracoda and Chironomidae, and in Shala mainly of Tubificidae, Ostracoda and Chironomidae. There were very few Nematoda. No true halophilic species were found but the community consisted of euryhaline forms found also in non-saline waters. Predatory invertebrates were absent and many of the dominant species, notably of the Chironomidae, were different from those of non-saline lakes nearby.     

Long-term changes in the benthic communities of the Pomeranian Bay (Southern Baltic Sea)

Long-term changes in the macrofauna of the Pomeranian Bay were studied by comparing survey data from the 1950s, 1980s, and 1990s. The study area has undergone significant eutrophication during the period of investigation. Biomass of filter-feeding bivalves increased significantly. Spatial distribution patterns of several species have changed. Strong decreases in species richness were caused by oxygen depletion at stations deeper than 15 m.Saduria entomon, Monoporeia affinis, andPontoporeia femorata vanished entirely between 1981 and 1993. Although a causal relationship between simultaneous increases of nutrient levels and macrobenthic biomass cannot be verified, eutrophication is proposed to be the major process affecing changes in macrofauna assemblages. In addition, changes in hydrography and climate increased frequency and severity of oxygen depletion events in the Pomeranian Bay since the mid 1980s.     

Ecological aspects of syllids (Annelida: Polychaeta: Syllidae) on Thalassia testudinum beds in Venezuela

The spatial and temporal variations of syllids associated with the tropical Thalassia testudinum were studied at four seagrass beds in the Morrocoy National Park. The epifaunal syllids were collected at quarterly intervals throughout one year using a modified suction sampler. Possible relationships between sampling sites, months and species were evaluated by Principal Components Analysis. The spatial-temporal variability of the predominant species was tested by means of a 2-factor ANOVA with the sampling sites and months as factors. A total of 1138 individual syllids were collected, belonging to 41 species and 12 genera, of which Branchiosyllis, Exogone, Odontosyllis, Sphaerosyllis and Syllis showed the highest diversity. Spatial variations were defined by the species of Branchiosyllis (B. exilis, B. lorenae, B. oculata) whereas temporal variability was defined by certain species of Syllis(S. beneliahui, S. broomensis and S. prolifera), the latter also being the most species rich genus present. Of the 41 species found, only these six showed consistent presence–absence and abundance patterns. Species richness and abundance were significantly higher in March at all sampling sites. Collectively, the syllid fauna of the Morrocoy seagrass beds was richer than recorded from other similar habitats. These findings are discussed in relation to the physical and biotic factors that may affect the variability of syllid populations in these seagrass habitats.     

Coral preference behaviour by planktotrophic larvae of Spirobranchus giganteus corniculatus (Serpulidae: Polychaeta)

Planktonic larvae of the serpulid polychaete Spirobranchus giganteus, an obligate associate of live coral, were tested for preferences for materials diffusing from natural substrates. Choices offered were Acropora prolifera, a very abundant coral on the Heron Island reef flat; Palauastraea ramosa, a less abundant coral, dead coral rubble and a glass tube as a control. Larval life is approximately 12 days. The larvae tested were 1–4 days old. Adults of S. giganteus occur commonly on A. prolifera and much less frequently on P. ramosa. Experiments were designed to prevent contact between larvae and substrate. Larvae preferred A. prolifera over P. ramosa, rubble and the control. There was no preference expressed between the control and P. ramosa or the control and rubble. A preference by young S. giganteus larvae for a substance diffusing from coral, acting together with a known positive phototaxis, may be adaptive in that it may help to maintain larvae in surface waters over the reef and in the vicinity of a specific coral until they are old enough to settle.     

Spring benthic macroinvertebrate communities of selected streams in the High Caucasus (Azerbaijan SSR)

Investigations on invertebrate fauna were carried out at fifteen sites in some chosen streams of the Caucasus (Azerbaijan SSR) in March 1970. Seventy-five taxa of invertebrates were found in the investigated streams. Chironomidae constituted the most numerous group at all sites, Ephemeroptera, Plecoptera, and Simuliidae being other important components of the fauna. On the basis of percentage structure of dominance in the examined streams four types of faunistic communities were distinguished. The most common one was the type characteristic of high mountain streams and rivers with Diamesa sp. (gr. latitarsis) predominant. Nevertheless, in springtime this community was characterized by a greater number of taxa and a greater abundance of specimens as compared with the summer period. It may be thus assumed that the spring period in high mountain streams and rivers is more conducive to the development of the invertebrate fauna.     

Ultrastructural aspects of Ellipsomyxa mugilis (Myxozoa: Ceratomyxidae) spores and developmental stages in Nereis diversicolor (Polychaeta: Nereidae)

The ultrastructure of the spores and developmental stages of Ellipsomyxa mugilis in Nereis diversicolor were studied by transmission electron microscopy. The ultrastructure features and the developmental stages show many similarities with the general pattern described for other actinospores. However, several new features are definitely worth noting. For example, tetranucleated cells precede the formation of the initial pansporocyst, which preserves the 2 original enveloping cells until the end of sporogony. In the initial stages of sporogony, the future sporoplasm cell acquires the first secondary cell by an engulfment process. In the final stage of sporogony, spores are formed by a sporoplasm with 2 secondary cells and 1 somatic nucleus, and the polar capsule has a polar filament with a helicoidal arrangement possessing 7-8 coils.     

Ecophysiological aspects of cardiac activity in the subtropical mussel Perna viridis (L.) (Bivalvia: Mytilidae)

Cardiac activity in the subtropical mytilid mussel Perna viridis was monitored to assess the influence of various endogenous and exogenous parameters. Temperature had a positive, significant, effect on heart rate; the Q₁₀ ranged from 2.5 to 2.6. Salinity, over a wide range (15-35‰), had no significant influence on heart rate reflecting the species' euryhalinity. Reduced oxygen tensions, either induced by emersion or seawater hypoxia, elicited rapid, significant, bradycardia. Positive inotropisms (indicative of greater heart output) were evident during seawater hypoxia and may represent an energetically advantageous mechanism of reducing heart rate while increasing stroke volume to maintain haemolymph circulation to organs associated with digestion and basal metabolism. Malnourishment for 14 days did not affect heart rate. The heart rate of larger mussels was slower than those of smaller individuals. Gender-based differences in P. viridis' heart rate were not present. No diurnal (24 h) variation in cardiac activity was evident while mussels were exposed to relatively stable, immersed conditions. Heart rate plasticity often reflected the ambient environmental conditions and such a flexible physiological response may partly explain the species' dominance in transitional, polluted harbours.