Macroinfaunal structure and effects of thermal discharges in a mesohaline habitat of Chesapeake Bay,near a Nuclear Power Plant

The soft-bottom macroinfauna of the Chesapeake Bay near the Calvert Cliffs Nuclear Power Plant, Maryland, USA, was studied quarterly in 1977. A total of 42,900 organisms distributed among 55 taxa was taken in 180 Smith-McIntyre grabs. Two communities and an ecotone between these two communities were identified in association with three sediment types: sand, sand-clay, and clay. Reproductive activities of 8 numerically dominant species are described. The macroinfauna at Calvert Cliffs was largely seasonal; species richness and organism abundances were lowest in late summer. The polychaetes Scolecolepides viridis and Heteromastus filiformis and the bivalves Macoma balthica and Mya arenaria were dominant in March-June. In September-December, the polychaetes Neanthes succinea, Paraprionospio pinnata, and Glycinde solitaria and the bivalve Gemma gemma became dominant. There were clear indications of differences associated with sediment types in total number of species, total number of individuals, and abundances of the dominant species. The shallower and intermediate habitats had higher sand-clay ratios, lower organic carbon contents, and larger numbers of organisms than the deeper clay habitat. Within each depth, no apparent detrimental effects caused by the thermal discharges on total number of species, total number of individuals, abundances of the dominant species, species diversity or evenness were found. Increased abundance was observed at the discharge stations for a few species, especially for Mya arenaria and N. succinea.     

Accumulation of polychlorinated biphenyls in the cockle Cerastoderma edule and the tellin Macoma balthica

Two species of marine bivalve, Cerastoderma edule (Linnaeus) and Macoma balthica (Linnaeus), were exposed to particles contaminated with three commercial polychlorinated biphenyl (PCB) mixtures, Aroclors^® 1242, 1254 and 1260. The accumulation of PCB in bivalve tissue was monitored for periods of up to 40 days. In addition to measuring total PCB concentrations, which reached 60 ppm of Aroclor 1242 in M. balthica, the fate of individual PCB homologues of different chlorine content, ranging from 2 to 8 chlorine atoms per biphenyl molecule, was also determined. Selective accumulation of PCBs with 5 chlorine atoms per biphenyl molecule occurred, whilst isomers of very low and very high molecular weights were accumulated at much slower rates.     

Variable feeding behavior in three species ofMacoma (Bivalvia: Tellinacea) as a response to water flow and sediment transport

Many species of tellinacean bivalves mainly deposit-feed but are also known to vary their feeding behavior in response to predation and to the availability of suspended organic matter relative to that of sedimentary organic matter. This study showed that three species of the Pacific genusMacoma (Macoma nasuta Conrad, 1937,Macoma secta Conrad, 1837, andMacoma inquinata Deshayes, 1855, from the San Juan Islands, Washington, USA) varied their deposit-feeding behavior in response to water flow near the sediment-water interface and to sediment transport. In the summers of 1987, 1988 and 1989, water velocity was varied in a large racetrack flume, in order to change both water velocity and bottom sediment transport. In quiet water, the inhalant siphon was protruded far from the siphon hole. As water velocity increased, with little or no sediment transport, the deposit-feeding radius decreased. Qualitative observations suggested that this was related to the drag on the siphon. InM. secta, the distance of siphon protrusion was not related to body size. Under conditions of higher near-bottom water velocity, combined with bottom sediment transport, some individuals ceased to deposit-feed, while others sustained feeding by ingesting sediment within the siphon hole, at least 1 cm beneath the sediment-water interface. These results suggest that hydrodynamic conditions are a major determinant of feeding behavior, and previous explanations of variable feeding behavior as a response to predation may have to be adjusted to accommodate this expanded set of responses.     

Effects of Lunatia heros predation on the population dynamics of Mya arenaria and Macoma balthica in Maine,USA

The interaction between the naticid snail predator Lunatia heros and 2 iteroparous, infaunal, intertidal bivalves was investigated in Lubec, Maine, USA. The Mya arenaria population consists primarily of young, small individuals. M. arenaria survivorship is low when young (3.5% yr^-1 for the first 5 yr), then increases. M. arenaria can attain a length of 60 mm, but it is susceptible to L. heros attack only until it is 30 mm long. It delays reproduction until it is 20 mm long (3.8 yr) and diverts its resources instead into rapid early growth (4.9 mm yr^-1 for the first 5 yr). The Macoma balthica population has a larger proportion of older individuals than does that of M. arenaria. Survivorship is higher for M. balthica than for M. arenaria (76.3% yr^-1 for the first 5 yr). Unlike M. arenaria, M. balthica attains a final length of only 25 mm and all sizes are susceptible to L. heros attack. M. balthica grows slowly (2.7 mm yr^-1 for the first 5 yr) and diverts its resources into earlier reproduction at a length of 10 mm (2.9 yr). These contrasting life-history patterns and the possible relationship between bivalve resource allocation and refuges from predation are discussed.     

Cycles and rates of reproduction of six Baltic Sea bivalves of different zoogeographical origin

All bivalves investigated (Astarte borealis, Astarte elliptica, Macoma calcarea, Cyprina islandica, Macoma baltica, Cardium lamarcki) are dioecious. The mean egg-diameters were determined by a series of measurements for 6 bivalves and compared with data from other authors. M. calcarea does not have a direct non-pelagic larval development as previously assumed, but probably produces pelagic planktotrophic larvae. The arctic species exhibit no modification of their mode of development in the boreal area. The boreal forms spawn over a relatively short period in spring and summer. The arctic forms revealed, over long periods, ripe eggs and sperms. Apparently they begin to spawn in winter or early spring. This extension and shift of the spawning season represents adaptation of the Aretic species to the specific conditions of the Baltic Sea. The number of eggs laid by the most important bivalves were determined and correlated with the larval ecology and length of generation time.     

Human trampling as short-term disturbance on intertidal mudflats: effects on macrofauna biodiversity and population dynamics of bivalves

The effect of physical disturbance in the form of trampling on the benthic environment of an intertidal mudflat was investigated. Intense trampling was created as unintended side-effect by benthic ecologists during field experiments in spring and summer 2005, when a mid-shore area of 25 × 25 m was visited twice per month by on average five researchers for a period of 8 months. At the putatively-impacted location (I) (25 × 25 m) and two nearby control locations (Cs) (25 × 25 m each), three sites (4 × 4 m) were randomly selected and at each site, three plots (50 × 50 cm) were sampled after 18 and 40 days from the end of the disturbance. Multivariate and univariate asymmetrical analyses tested for changes in the macrofaunal assemblage, biomass of microphytobenthos and various sediment properties (grain-size, water content, NH₄ and NO₃ concentrations in the pore water) between the two control locations (Cs) and the putatively-impacted location (I). There were no detectable changes in the sediment properties and microphytobenthos biomass, but variability at small scale was observed. Microphytobenthos and NH₄ were correlated at I to the number of footprints, as estimated by the percentage cover of physical depressions. This indicated that trampling could have an impact at small scales, but more investigation is needed. Trampling, instead, clearly modified the abundance and population dynamics of the clam Macoma balthica (L.) and the cockle Cerastoderma edule (L.). There was a negative impact on adults of both species, probably because footsteps directly killed or buried the animals, provoking asphyxia. Conversely, trampling indirectly enhanced recruitment rate of M. balthica, while small-sized C. edule did not react to the trampling. It was likely that small animals could recover more quickly because trampling occurred during the growing season and there was a continuous supply of larvae and juveniles. In addition, trampling might have weakened negative adult-juvenile interactions between adult cockles and juvenile M. balthica, thus facilitating the recruitment. Our findings indicated that human trampling is a relevant source of disturbance for the conservation and management of mudflats. During the growing season recovery can be fast, but in the long-term it might lead towards the dominance of M. balthica to the cost of C. edule, thereby affecting ecosystem functioning.     

Availability of copper from phytoplankton and water for the bivalveMacoma balthica. I. Separation of uptake pathways using the radiotracer64Cu

To study the role of food in Cu accumulation by bivalves, algae spiked with Cu can be used. With spiked algae, however, redistribution of Cu between the disolved solved and the particulate phase hampers the assessment of the contribution of food. This occurred in efforts to label algae with the radiotracer⁶⁴Cu. A method was designed to overcome this problem of redistribution. By adding excess EDTA to seawater, the biological availability of dissolved Cu was minimized. The effectiveness of complexation by EDTA was controlled through adsorption onMacoma balthica shells and uptake inM. balthica tissue.Communication number 644 of the Centre for Estuarine and Coastal Ecology, Yerseke, The Netherlands     

Minimal predation upon meiofauna by endobenthic macrofauna in the Exe Estuary,south west England

Field manipulation experiments were performed in the Exe Estuary, south west England, in October 1988, to investigate the importance of the meiofaunamacroinfauna trophic link in benthic trophodynamics. Four hypothetically meiofauna-predacious endobenthic macrofauna species were selected for manipulation using the criteria of high abundance and different modes of feeding: Cerastoderma edule (filter-feeder), Nereis diversicolor (omnivorous scavenger), Ophelia bicornis (sand-ingester), Scrobicularia plana (deposit-feeder). Enclosures constructed from plastic tubing, 63 m nylon monofilament mesh and galvanized steel were deployed, containing adult members of these taxa at densities raised to approximately four times that of the surrounding sediment. The experiments ran for 12 tidal cycles. Differences in phyletic meiofaunal abundance between treatment and control enclosures at the termination of the experiment were assessed using both uni- and multivariate dataanalysis techniques. Only two univariate significant differences (p<0.05) existed for the N. diversicolor treatment and two for the S. plana treatment. Annelida, Turbellaria and copepod nauplii were the only meiofauna taxa affected. No univariate significant differences were recorded for either the C. edule or O. bicornis treatments. Multidimensional scaling ordination of the data revealed no consistent changes in community composition between treatments and controls. It is concluded that the experiments provide evidence of minimal predation by macroinfauna upon meiofauna.     

Inhibition of larval settlement to a soft bottom benthic community by drifting algal mats: An experimental test

The hypothesis that drifting red algal mats inhibit settlement of planktonic larvae was tested in a field experiment in 1986. Substratum free of algae (caged boxes) was compared with substratum covered with algae (natural substratum and open boxes). Whereas settling densities of 1500 to 5500 ind. m^-2 of the bivalves Macoma balthica, Cardium glaucum and Mya arenaria were observed in sediment without algal cover, no recruits of the same species were found beneath the algae during the period of peak settlement (June to July). The same difference was observed for the polychaete Nereis diversicolor, although in this case occasional individuals were found beneath the algae. The results demonstrate that algae mats may be efficient inhibitors of larval settlement to benthic soft-substratum communities. It is concluded that algae act as a larval filter.     

Metallothionein-like metalloproteins in the Baltic clam Macoma balthica : seasonal variations and induction upon metal exposure

Metallothionein-like proteins (MTLs) have been measured by differential pulse polarography every two months during a period of two and half years in the marine bivalve Macoma balthica from two locations of the Western Scheldt Estuary. The MTL concentrations (0.85 to 7.81 mg g⁻¹ dry wt) are comparable to most values found for other marine invertebrates. These concentrations vary significantly with the seasons (higher in winter, lower in summer), which is mainly due to body weight fluctuations. Strong correlations exist between MTL and metal levels. Upon short-term exposure to a mixture of Cd, Cu and Zn, there are on average significant MTL increases, but important seasonal variations have been observed: in winter, the clams are more sensitive to metals, uptake more Cd and Cu and induce more MTL than during the warmer months. Received: 16 May 1997 / Accepted: 6 June 1997     

Persistence of polychlorinated biphenyls in marine bivalves

Bivalves (Cerastoderma edule and Macoma balthica) which had previously been exposed to Aroclors^® 1242, 1254 and 1260 were able to reduce their tissue burdens of chlorobiphenyls with 2 to 5 chlorine atoms in short-term static assay systems. Elimination rates decreased with increasing chlorination and removal of isomers with more than 5 chlorine atoms was not recorded. Position, in addition to the number, of chlorine atoms influenced the persistence of chlorobiphenyls. Isomers with most ortho-substituted chlorine atoms were least persistent. Experiments with single isomers indicated variation in the elimination of low chlorinated isomers between bivalve species. These biological and chemical influences on tissue residues, together with environmental parameters such as temperature and suspended solids, are considered in relation to the use of bivalves as bio-indicators of marine pollution.     

Transfer of brevetoxins to a tellinid bivalve by suspension- and deposit-feeding and its implications for clay mitigation of <Emphasis Type="Italic">Karenia brevis</Emphasis> blooms

Blooms of the brevetoxin-producing Karenia brevis in the Gulf of Mexico cause massive fish kills, food poisoning and adverse respiratory effects in humans. Sedimentation of toxic cells following inert clay application could reduce toxin incorporation by commercially important suspension-feeding bivalves and thus prevent direct public health impacts, but could potentially lead to brevetoxin (PbTx) accumulation by benthic deposit-feeders. The goal of this study was therefore to compare suspension- and deposit-feeding as pathways for brevetoxins. We investigated: (1) the effect of toxic K. brevis on both feeding modes using a facultative deposit-suspension feeding tellinid bivalve, the clam Macoma balthica, as a model species and (2) the relative effectiveness of brevetoxin transfer via suspension- and deposit-feeding over 24-h exposure. Sedimentation of K. brevis was achieved by treatment with 0.25 g phosphatic clay l⁻¹ and brevetoxin concentrations were measured by ELISA. Karenia brevis reduced both suspension- and deposit-feeding activity. This study demonstrates that brevetoxins can be rapidly accumulated by a surface deposit-feeding bivalve from sedimented K. brevis cells and that comparable toxin levels can be attained by both suspension- and deposit-feeding modes [1.2–1.6 μg PbTx (g tissue wet weight)⁻¹]. Deposit-feeding clams generally do not pose a direct threat to humans but may provide a pathway for brevetoxin food web transfer.     

Développement larvaire de deux Tellinacea,Scrobicularia plana (Semelidae) et Donax vittatus (Donacidae)

Scrobicularia plana Da Costa and Donax vittatus L. were reared in the laboratory through settlement. Fertilizable eggs were obtained by perfusing the ovary with 5% 0.1 M ammonium hydroxyde in sea water. S. plana eggs have a thick chorion, inside which the early larval stages develop; they hatch as straight-hinge larvae more than 60 h after fertilization. This brood protection is considered to be an adaptation to osmotic pressure changes and pollution in the environment. D. vittatus eggs have a very thin chorion and are unprotected. Further development is planktotrophic and very similar for the two species. Under laboratory conditions, the pediveliger stage is attained 3 weeks after fertilization and settlement occurs 1 week thereafter. S. plana spat stop growing until a suitable substratum is available. Meanwhile they undergo a byssus drifting important for postlarval dispersion. Post larvae resume growth as soon as a small quantity of fine sand is added to the rearing jar. The exhalant siphon is developed first, when the post larvae reach a length of 600 m; the inhalant siphon is formed later, at a length of approximately 900 m. Evolution from the larval hinge to the juvenile hinge stage occurs sooner in S. plana than in D. vittatus. Comparison of laboratory larval development with field development indicates that spawning occurs in June and August for S. plana in North Wales (UK).     

An electron-microscope study of periostracum formation in some marine bivalves. I. The origin of the periostracum

Electron microscopy has revealed the presence of specialised basal cells at the base of the periostracal groove in the bivalves Mytilus edulis (L), Nucula sulcata Bronn., Cardium edule (L) and Macoma balthica (L). A common basal cell type is observed which gives rise to a pellicle in all cases except N. sulcata. The pellicle is the first part of the periostracum to be elaborated, and it is suggestd that it may be an indication of specialisation in bivalves. In N. sulcata and C. edule, an additional cell type is present. In N. sulcata, this second basal cell type appears to be directly involved in periostracum formation, whereas in C. edule an indirect role is proposed.     

Single-step nested multiplex PCR to differentiate between various bivalve larvae

A nested multiplex polymerase chain reaction (PCR) assay has been developed that allows the discrimination between six bivalve larvae common to Danish coastal waters (Cerastoderma edule, Macoma balthica, Mytilus edulis, Spisula subtruncata, Ensis americanus and members of the order Myoida). This assay involves the simultaneous use of a pair of general universally targeted 18S rRNA gene primers, five specific 18S rRNA gene targeted oligonucleotide primers internal (nested) to the universal primer pair and one species-specific primer that is not nested (Mya). The specificity of each primer was evaluated in silico, empirically, and verified further by sequencing of amplification products from single larvae collected from plankton samples. Identification of individually isolated bivalve larvae from plankton samples was based on the size of the PCR product produced by the specific primers after visualisation by agarose gel electrophoresis. Preliminary studies indicated that this method was suitable for use with freshly collected and preserved larvae, and is therefore suitable for field application.Communicated by M. Kühl, Helsingør     

Passing the gut of juvenile flounder, Platichthys flesus: differential survival of zoobenthic prey species

We studied the resistance of benthic prey organisms (Ostracoda, Hydrobia spp., Macoma balthica) to digestion by juvenile flounder, Platichthys flesus (L.), in a series of aquarium experiments. Results showed that some food species are able to survive the gut passage (Ostracoda 53 to 75%, Hydrobia spp. 46 to 92%). This has ecological implications, as increasing eutrophication of the Baltic Sea favours prey species that are nondigestible for the juvenile flounder. Thus, seemingly small changes in food-web structure may have unpredictable effects for the predatory fish. Received: 8 March 1997 / Accepted: 14 March 1997     

Natural variability in distribution of an intertidal population of Macoma balthica subject to potential oil pollution at Port Valdez,Alaska

Natural variability in the abundance of an intertidal population of the lamellibranch Macoma balthica (Linnaeus, 1758) was measured during 1971 and 1972 in a study area near the proposed oil storage and tankship loading facility at the southern terminus of the Trans-Alaska pipeline in Port Valdez, Alaska. M. balthica were divided for analysis into a large and a small size category. Small temporal changes in population densities throughout the entire study area were detected for both size categories over several of the 7 sampling times of the 2-year period. Large and persistent differences in density were found among elevation contour intervals for either size category; however, variations in the density profiles on elevation occurred among sampling times. Large M. balthica became more equitably distributed and the small category less equitably distributed among elevation contours over the 2-year period. Densities of both size categories were more stable at the higher elevations of the study site. Large M. balthica were more homogeneously distributed along a given elevation contour interval than the small category. Mobility and time available to redistribute at a horizontal location would explain the more homogeneous distribution of large M. balthica if competition for food resources exists.     

The “Tsesis” oil spill: Acute and long-term impact on the benthos

The “Tsesis” oil spill in October 1977 resulted in the release of over 1 000 tons of medium grade fuel oil in an archipelago in the brackish Baltic Sea. Considerable oil quantities reached the benthos by sedimentation. Within 16 d benthic amphipods of the genus Pontoporeia, as well as the polychaete Harmothoe sarsi Kinberg, showed reduction to less than 5% of pre-spill biomasses at the most impacted station. The clam Macoma balthica (L.) was more resistant, and showed little or no mortality, but was heavily contaminated by oil (about 2 000 μg g^-1 dry wt total hydrocarbons). The meiofauna was strongly affected, with ostracods, harpacticoids, Turbellaria and kinorhynchs showing clear reductions in abundance, while nematodes, as a group, were more resistant. In the winter following the spill gravid Pontoporeia affinis Lindström females showed a statistically significant increase in the frequency of abnormal or undifferentiated eggs. Food-chain transfer of oil to flounder [Platichthys flesus (L.)] was indicated. Not until the second summer after the spill were the first signs of recovery noted at the most heavily impacted station: Amphipods, H. sarsi and harpacticoids increased and the oil concentrations in M. balthica decreased (to about 1 000 μg g^-1). In the area where amphipods had been virtually eliminated, there was an unusually heavy recruitment of M. balthica, reaching 4 000 juveniles, of 1.5–2 mm length, per square metre, probably from settling in summer 1978. Three years after the spill Pontoporeia spp. biomass was still depressed in the most affected area, while H. sarsi showed normal biomass, and M. balthica abundance was inflated. Oil concentrations in M. balthica (about 250 μg g^-1) and flounder were only slightly elevated and the oil could no longer be confidently ascribed to “Tsesis” origin, even using GC/MS-analysis. Recovery was thus underway, but the long lifespan of M. balthica implies that the disturbed community composition may persist for many years at this station. Full recovery is likely to require more than 5 yr and may take a decade or more. An effort to evaluate the accumulated monetary loss to fishery from the accident indicates that direct costs of shoreline cleanup and vessel damage were considerably greater.     

Effect of salinity fluctuation on the osmotic pressure and Na+, Ca2+ and Mg2+ ion concentrations in the hemolymph of bivalve molluscs

Specimens of Chlamys opercularis, Modiolus modiolus, Mytilus edulis, Crassostrea gigas, Scrobicularia plana and Mya arenaria were exposed to both gradual (sinusoidal) and abrupt (square-wave) salinity fluctuations and measurements made of osmotic, Na⁺, Mg²⁺ and Ca²⁺ concentrations in the hemolymph and where applicable in the mantle fluid. In both sinusoidal and square-wave regimes fluctuating between 100 and 50% seawater (100%=ca. 32 S), the hemolymph Na⁺, Mg²⁺, Ca²⁺ and osmotic concentrations followed the concentrations of the external medium in Chlamys opercularis. The hemolymph and mantle fluid osmotic Na⁺, Mg²⁺ and Ca²⁺ concentrations of Modiolus modiolus, Mytilus edulis, Crassostrea gigas and S. plana followed those of the external medium as long as the molluscs' shell valves remained open. There were no changes in the ionic or osmotic concentrations of the hemolymph or mantle fluid of any of these species during periods of shell-valve closure. The hemolymph osmotic, Na⁺ and Mg²⁺ concentrations of wedged-open Modiolus modiolus, Mytilus edulis, C. gigas and S. plana followed those of the external medium. Hemolymph Ca²⁺ concentrations showed a damped response in C. gigas and Mytilus edulis. The hemolymph osmotic, Na⁺, Ca²⁺ and Mg²⁺ concentrations of Mya arenaria fluctuated in a similar manner to the external medium, but were damped. Wedged-open Mytilus edulis exposed to fluctuating salinity and supplied with a constant supply of 10 mM Ca²⁺ showed greater changes in hemolymph ionic and osmotic concentrations than M. edulis exposed to the same salinity fluctuation without a constant Ca²⁺ supply. Chlamys opercularis and Modiolus modiolus survived in a 50% seawater minimum sinusoidal salinity fluctuation for 10 days; wedged-open M. modiolus survived only 3 days. Burrowing had no effect on the osmotic, Na⁺, Mg²⁺ or Ca²⁺ concentrations of the hemolymph of Mya arenaria or S. plana exposed to fluctuating salinities. All of the species studied were shown to be osmoconformers.