Potential effects of a non-indigenous predator in its expanded range: assessing green crab, <Emphasis Type="Italic">Carcinus maenas</Emphasis>, prey preference in a productive coastal area of Atlantic Canada

The non-indigenous green crab (Carcinus maenas) is an important predator on bivalve wild beds in coastal areas worldwide. This study explored size-dependent green crab prey preference on American oysters (Crassostrea virginica), blue mussels (Mytilus edulis), and soft-shell clams (Mya arenaria) in a productive coastal system of Atlantic Canada. Using two sizes of prey and three different experimental manipulations, small, medium, and large green crabs were given a choice among these three bivalves, and their daily feeding rates were monitored over the course of 3 days. For both prey sizes, green crabs showed an early feeding preference for soft-shell clams and, only as they declined in numbers, a switch toward mussels and subsequently toward oysters. We found that such changes in the timing (order) of prey preference are related to prey differences in shell thickness, a fairly reliable indicator of prey shell strength.     

Assimilation efficiencies and turnover rates of trace elements in marine bivalves: a comparison of oysters, clams and mussels

Assimilation efficiencies (AEs) and physiological turnover-rate constants (k) of six trace elements (Ag, Am, Cd, Co, Se, Zn) in four marine bivalves (Crassostrea virginica Gmelin, Macoma balthica Linnaeus, Mercenaria mercenaria Linnaeus, and Mytilus edulis Linnaeus) were measured in radiotracer-depuration experiments. Egestion rates of unassimilated elements were highest during the first 24 h of depuration and declined thereafter. Significant egestion of unassimilated Co, however, continued for up to 5 d in Macoma balthica, Mercenaria mercenaria and Mytilus edulis. With the exception of the extremely low values for ^110 mAg, ¹⁰⁹Cd, and ⁶⁵Zn in C. virginica, physiological turnover-rate constants (k) showed no general pattern of variation among elements, bivalve species or food types, and were relatively invariant. Values from  ≤0.001 to 0.1 d⁻¹ were observed, but excluding those for Co, most values were  ≤0.04 d⁻¹. In all four species, the AEs of Ag, Am, and Co were generally lower than those of Cd, Se, and Zn. The AEs of Ag, Cd, Se, and Zn in these bivalves are directly related to the proportion of each element in the cytoplasmic fraction of ingested phytoplankton, indicating that >80% of elements in a prey alga's cytoplasm was assimilated. C. virginica, Macoma balthica, and Mercenaria mercenaria assimilated ∼36% of the Ag and Cd associated with the non-cytoplasmic (membrane/organelle) fraction of ingested cells in addition to the cytoplasmic fraction. The ratio of AE:k, which is proportional to the consumer–prey trace-element bioaccumulation factor (concentration in consumer:concentration in prey) was generally greater for Cd, Se, and Zn than for Ag, Am, and Co. This ratio was lowest in Mytilus edulis, suggesting that this bivalve, the most widely employed organism in global biomonitoring, is relatively inefficient at accumulating important elements such as Ag, Cd, and Zn from ingested phytoplankton. Received: 7 February 1997 / Accepted: 24 February 1997     

The role of marine aggregates in the ingestion of picoplankton-size particles by suspension-feeding molluscs

Suspension-feeding molluscs are important members of coastal communities and a large body of literature focuses on their feeding processes, including the efficiency of particle capture. Some molluscs, such as bivalves, capture individual picoplankton cells (0.2–2.0 μm) with a retention efficiency of less than 50%, leading to the assumption that such particles are not an important food resource. Picoplankton, however, are often concentrated in particle aggregates of much larger size. This study investigates the ability of suspension feeders to ingest picoplankton-size particles (0.2–2.0 μm) bound in marine aggregates. We fed clams (Mercenaria mercenaria), mussels (Mytilus edulis), oysters (Crassostrea virginica), scallops (Argopecten irradians) and slipper snails (Crepidula fornicata) 1.0- and 0.5-μm fluorescent particles (either polystyrene beads or bacteria) that were (1) dispersed in seawater, or (2) embedded within laboratory-made aggregates. Dispersed 10-μm beads were also delivered so that feeding activity could be determined. Ingested fluorescent particles were recovered in feces or isolated digestive glands and quantified. Results indicate that aggregates significantly enhance the ingestion of 1.0- and 0.5-μm beads by all species of bivalves, and enhance the ingestion of bacteria (greatest cell dimension ca. 0.6 μm) by all suspension feeders examined. Differences among species in their ability to ingest aggregates and picoplankton-size particles, however, were evident. Compared to mussels and clams, scallops and oysters ingested fewer aggregates with 1.0-μm beads or bacteria, and slipper snails ingested the most dispersed beads and bacteria. These differences may be a consequence of variations in gill structure and mechanisms of particle processing. Our data demonstrate that suspension feeders can ingest picoplankton-size particles that are embedded within aggregates, and suggest that such constituent particles may be an important food resource.     

Behavioral defenses of Crepidula fornicata against attack by Urosalpinx cinerea

Two behavioral defenses of the slipper shell Crepidula fornicata (L.) against attack by the oyster drill Urosalpinx cinerea (Say) are described: (1) Jabbing: when an oyster drill approaches, the slipper shell lifts the edge of its shell, extends the head and, with the radula, pricks or rasps the foot of the predator, which instantly retracts into its shell; sometimes, after repeated approaches, the oyster drill moves away. (2) Pivoting: when an oyster drill mounts C. fornicata, the latter slowly and repeatedly rotates its shell in a horizontal plane at a constant rate and through a constant arc. If this action brings the predator's shell to bear against an obstacle, motion ceases and C. fornicata maintains the pressure until the oyster drill is dislodged. These aggressive tactics are adaptive defensive measures for a nearly sessile prey otherwise highly vulnerable to oysterdrill attack.     

Genetic structure in native and non-native populations of the direct-developing gastropod Crepidula convexa

In many marine invertebrate species, larval development plays an important role in population connectivity and gene flow: species with direct benthic development generally show more genetic structure than those with planktonic development. We used nuclear markers (microsatellites) to determine population genetic structure of the direct-developing snail Crepidula convexa (Gastropoda: Calyptraeidae) in seven populations with 15–85 individuals each within its native range of the northwest Atlantic and compared it to Crepidula fornicata, a congener with planktonic development. Our results are consistent with general expectations and previous work in these species with other markers: C. convexa had greater population structure and even at a regional scale shows significant isolation-by-distance, in contrast to C. fornicata. We also genotyped a single population of C. convexa introduced to the northeastern Pacific to investigate the prediction of reduced genetic diversity following introduction (founder effect). We did not find a reduction in genetic diversity, suggesting that this non-native population may be characterized by multiple introductions. This pattern is consistent with many other introduced populations of marine invertebrates, including C. fornicata.     

Environmental stress,competition and dominance of Crassostrea virginica near Beaufort,North Carolina,USA

Observations and experiments were made at 2 intertidal areas near Beaufort, North Carolina, USA from May 1977 to July 1978 to determine why the oyster Crassostrea virginica dominated the community in areas protected from wave action but not in areas directly exposed to waves. Barnacles, oysters, the green alga Enteromorpha sp. and the mussel Brachiodontes exustus were the main occupiers of primary space at the mid and low intertidal levels of exposed areas. The intertidal community at the protected site consisted of a mid intertidal occupied by the barnacles Balanus amphitrite and Chthamalus fragilis, and the oyster C. virginica, and a low intertidal dominated by C. virginica. The exposed area was highly variable with high colonization and mortality for all species producing large seasonal changes in structure. The protected site remained constant throughout the year; there was no evidence of further colonization of either barnacles or oysters and mortality was very low. Recolonization experiments, selective removal of species, and growth and survival data demonstrated that C. virginica does not become dominant at exposed locations because (1) the constant wave shock at the ocean site reduces growth and increases mortality of young and adult oysters and (2) oysters are outcompeted by the mussel B. exustus. The monopolization of space by C. virginica at protected sites contrasts with studies north of Cape Hatteras where the abundance of predators produces a more diverse and heterogeneous community. Predation was unimportant in Beaufort because predators were absent at the exposed areas and the oyster drill Urosalpinx cinerea was restricted to the subtidal zone at the protected site. This absence of predators indicates a higher level of environmental stress near Beaufort compared to areas farther north.     

Mitochondrial DNA analysis of native and selectively inbred Chesapeake Bay oysters,Crassostrea virginica

Mitochondrial DNA (mtDNA) genotypes were examined in 1989 in threeCrassostrea virginica (Gmelin) populations native to Chesapeake Bay, USA, and in one population selectively inbred for rapid growth for ten generations. We wished to determine whether this character would be useful as a genetic marker for distinguishing between the inbred line and the native oysters and to determine whether detectable genetic differences exist among present-day native populations ofC. virginica. Thirty mtDNA haplotypes were identified. The average percentage nucleotide difference between native haplotypes was 1.8%. Inbred oysters were characterized by mtDNA haplotypes distinctly different from ancestral native oysters, indicating a high degree of genetic differentiation between the two groups (average percentage mtDNA nucleotide sequence divergence,=21.8%). The common native haplotype was not present in the selectively inbred sample, and six of the seven haplotypes detected in the inbred oysters were not found in the survey of native oysters. Chi-square tests on haplotype frequencies indicated that the native populations were not significantly different from one another. However, the distribution and relatedness of haplotypes suggest that significant change in the oyster gene pool may have occurred over the past few decades.     

Scanning electron microscopy of metamorphosis in four species of barnacles (Cirripedia Thoracica Balanomorpha)

Metamorphosis and early juvenile development was followed in the laboratory in Balanus amphitrite and B. improvisus (family Balanidae) and in Semibalanus balanoides and Elminius modestus (family Archaeobalanidae) from cyprid settlement until 24 h after ecdysis. Stages of development were studied in vivo and with scanning electron microscopy. Events in metamorphosis and early juvenile development are very similar in all four species and can be interpreted in terms of a highly modified moult. Notably, there was no indication whatsoever of the amorphous decorticated settler previously described from metamorphosis of B. amphitrite. The shape of a juvenile barnacle with cirri and incipient shell plates is apparent immediately after the shedding of the cypris carapace, and rudimentary peduncle can be distinguished below the developing wall plates. A basal row of cuticular hairs encircles the peduncle in all species except S. balanoides. These hairs seem to serve a restraining function during early development, when the juvenile barnacle is only attached by the initially secreted cyprid cement. Similarly situated hairs are also present in very young juveniles of the lepadomorph Scalpellum scalpellum. In Semibalanus balanoides nothing indicates that the rostrum originates from the fusion of two latera as previously claimed, since this plate is single as soon as it can be distinguished.     

Mode of particle ingestion in five species of suspension-feeding bivalve molluscs

In order to elucidate the mode of particle ingestion and the functional anatomy of the oesophagus in bivalves, a histological study was performed onMytilus edulis (Mytilidae),Crassostrea virginica (Ostreidae),Placopecten magellanicus, Chlamys varia, and juvenilePecten maximus (Pectinidae). Specimens were sampled from various sites in New Brunswick, Canada, and Brittany, France, from 1987 to 1989. The buccal, peribuccal, and oesophageal epithelia of all species contained a dense distribution of actively secreting mucocytes, although these were somewhat less abundant inCrassostrea virginica, which also has the shortest oesophagus. Mucocyte morphology, while constant within a family (Pectinidae), showed clear differences among families. Both acid and neutral mucopolysaccharides were secreted by the epithelial mucocytes of all species. Mucus and mucus-particle masses were observed in the peribuccal and buccal regions, as well as in the oesophageal lumina of all species, even in those specimens which had been maintained without feeding (Placopecten magellanicus) or held out of water for 48 h (C. virginica) prior to dissection and fixation. These results indicate that a basal level of mucus production and transport is continuous on the peribuccal, buccal, and oesophageal ciliated epithelia, regardless of the particle concentration in the external medium. Buccooesophageal glands, generally thought to be absent in the Bivalvia, were observed in one of the species examined (M. edulis). It is concluded that the mode of particle ingestion in these suspension-feeding bivalves is via ciliatransported mucus masses; the presence of buccooesophageal glands inM. edulis suggests a digestive role for the oesophagus in this species.     

Pathways of arsenic uptake and incorporation in estuarine phytoplankton and the filter-feeding invertebrates Eurytemora affinis,Balanus improvisus and Crassostrea virginica

Arsenic uptake from water and from phytoplankton was followed in the copepod Eurytemora affinis and the barnacle Balanus improvisus collected from the Patuxent River estuary, Chesapeake Bay, eastern coast of the USA in 1987, and in the oyster Crassostrea virginica obtained from a hatchery on the shore of Chesapeake Bay in 1987. Dissolved arsenic was readily taken up by phytoplankton and by shell material of B. improvisus and C. virginica; however, no dissolved arsenic was incorporated into the invertebrate tissues. When E. affinis, B. improvisus and C. virginica were fed phytoplankton containing elevated arsenic contents, significant arsenic incorporation occurred. Juvenile B. improvisus incorporated relatively more arsenic than adults of all three species. Compared to the 100 to 200% increase in arsenic content by phytoplankton exposed to dissolved arsenic, the 25 to 50% increase in these invertebrate species via trophic transfer is relatively small. Even though the trophic pathway for arsenic transfer is the major one for higher trophic levels within an ecosystem, the potential for direct arsenic impact to trophic levels other than phytoplankton appears to be minimal.     

Untersuchungen über die temperaturabhängigkeit des sauerstoffverbrauchs von Crepidula fornicata (Mollusca: Prosobranchia)

The oxygen consumption of the sessile gastropod Crepidula fornicata was measured during autumn 1962 as a function of soft body wet weight (0.04 to 5.2 g) and temperature (2° to 30° C). From the data obtained, the parameters of the allometric formula were derived for describing the relationship between respiration and body size. The parameters were then used for calculating the oxygen consumption of 3 different weight groups (0.1 g, 1.0 g, 10.0 g) at different temperatures. The resulting plot reveals different curves, demonstrating that the influence of temperature on the metabolism of C. fornicata is a function of body size. The ourves relating oxygen consumption to temperature may well be described by a modified Arrhenius-function proposed some years ago (Krüger, 1962). Biological temperature curves may be described by growth functions. Therefore, it is possible to apply the simple Walfordplot for identifying the “normal curves” of Krogh (1914).     

Environmental control of the breeding of three boreo-arctic cirripedes

Three species of barnacles with boreo-aretic distribution were shown to require maintenance for several weeks below a critical temperature before the breeding condition could be attained. The temperatures critical for Balanus balanoides (L.) Balanus balanus (L.) and Balanus crenatus (Bruguière) were found to be between 10° and 12°C, 10° and 14°C, and at about 17°C, respectively. Although the strong influence of continuous light and the weak influence of continued feeding in delaying the onset of breeding in B. balanoides were confirmed, there remained some outstanding anomalies between the breeding behaviour of this species under laboratory conditions and between the tide marks. It was found impossible to initiate breeding by the application of conditioning procedures significantly in advance of the time of the normal Autumn breeding season in B. balanoides. Breeding appears to be inhibited, independently of external conditions, for a set period after the preceding brood cycle. Evidence points to a similar, largely endogenous, control of breeding in B. balanus which also breeds once annually, but not in B. crenatus which breeds continuously so long as food and temperature levels permit.     

Electrophoretic identification and genetic analysis of bivalve larvae

Taxonomic identification and genetic analysis of larval marine invertebrates have been vexing problems. We describe a polyacrylamide mini-gel electrophoresis technique for resolving proteins from individual larval bivalves (shell length 250 to 350 m) and apply this technique to three species of laboratory-cultured larval oysters [Ostrea edulis L., 1758, Crassostrea gigas (Thunberg, 1793) and c. virginica (Gmelin, 1791)] reared during summer 1989. Electrophoretic patterns of proteins clearly discriminate among the three species and allow genetic analysis of a polymorphic allozyme locus (Pgi) in field-collected larvae and juveniles of C. virginica. This technique provides an economical tool for largescale taxonomic, ecologic, and genetic studies of meroplanktonic stages of various species.     

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.     

Adjustment of the components of energy balance in the gastropod Crepidula fornicata in response to thermal acclimation

Synchronous measurements were made of the routine rate of oxygen consumption and the clearance rate of Phaeodactylum tricornutum at different exposure temperatures by specimens of the suspension-feeding gastropod Crepidula fornicata which had been acclimated to temperatures between 10° and 25°C. The results show that the cost of activity (l O₂ consumed h^-1/ml seawater cleared h^-1) increases dramatically in individuals exposed to short-term increases of temperature up to 30°C, especially in limpets acclimated to 10°C. The process of thermal acclimation, however, results in two compensatory adjustments in energy expenditure and uptake which profoundly affect the energetics of water transport. Firstly, the routine oxygen consumption shows lateral translation of the rate-temperature curve which results in the maintenance of a relatively uniform energy expenditure despite an increase in acclimation temperature from 10° to 25°C. Secondly, because of the form of the rate-temperature curve for filtration by C. fornicata, lateral translation in response to warm acclimation results in an increase in the maximal clearance rate. Lateral translation of the rate-temperature curves for feeding rates and for oxygen consumption in response to thermal acclimation may thus be linked to maintain a balance between energy gain and expenditure. In this way, the greatly increased cost of activity which would occur with increase of temperature in the absence of acclimation is evaded. The minimal maintenance energy requirement, and hence the greatest scope for growth and reproduction, is then adjusted to coincide with temperatures prevailing in the environment.     

Seasonal changes in the free glycerol level in the body parts of the adult barnacle Balanus balanoides

Seasonal changes in the free glycerol level in body parts of the adult barnacle Balanus balanoides have been followed during the 1969 season. Glycerol levels were highest in winter when the barnacles were most cold-tolerant and lowest in summer when they were least cold-tolerant. It is suggested that glycerol, and possibly other solutes, may be concentrated in the blood of B. balanoides, thus promoting supercooling and reducing the amount of ice formed at sub-zero temperatures.     

The transport of oyster larvae in an estuary

Vertical and horizontal distributions of 3 larval stages of the oyster Crassostrea virginica were measured concurrently with phytoplankton species compositions, phytoplankton size distributions and physical hydrographic parameters in tributaries of the Chesapeake Bay (USA) during the oyster spawning seasons of 1980 and 1981. The superposition of the biological distributions upon the physical hydrographic data provide instantaneous distributions of the entire system which are consistent with the upstream transport of oyster larvae. Oyster larvae distributions in the Choptank River and its Broad Creek and Tred Avon River tributaries can be described in terms of three contiguous regions: (1) a common spawning region, (2) an intermediate, upstream transport region and (3) a seed bed region where major spat set occurs. The phytoplankton species compositions and abundances in the size fraction less than 10 m in the tributary system during the transport were sufficient to supply optimum growth requirements of developing larvae. The transport proposed can explain the 30 yr record of consistently higher spat set success in one tributary, Broad Creek, relative to an adjoining tributary, the Tred Avon River. This may be a general mechanism whereby oysters maintain reproductive success and emigrate to seed bed regions in the Chesapeake Bay.Contribution No. 1144 of the McCollum-Pratt Institute and Department of Biology     

Organics in wheat and rye straw pulp bleaching and combined mill effluents (II) ecotoxicological screening

Potential environmental effects of the discharge of industrial wastewater from manufacturing of bleached straw pulp have been studied. Bleached neutral sulphite straw pulping results in discharges of both black cooking and spent bleaching liquors. Time proportional samples were taken from the combined bleaching effluent and the combined mill effluent, (i) Mutagenic activity of the two samples in an Ames’ test, and (ii) acute toxicity of the combined mill effluent sample towards the photosynthetic activity of natural marine phytoplankton from the receiving waters were determined. Also, acute toxicity was determined of the slowly biodegradable or persistent organics remaining after aerobic stabilization of the sample towards (ii) photosynthetic activity of natural phytoplankton, (iii) mussels (Mytilus edulis), (iv) eels (Anguilla anguilla) and, (v) crustacean (Nitocra spinipes); and furthermore, (vi) reproduction test of the crustacean and (vii) growth inhibition test of the marine diatom (Phaeodactylum tricornutum) were performed. Mutagenicity emission factors (MEF's) and toxicity emission factors (TEF's) were calculated to classify the effluents. Combined mill effluent sample showed a mutagenic activity per t₉₀ of one fourth the activity of a kraft pulp bleaching effluent, and the inhibition of the photosynthesis of natural phytoplankton was significant up to 2600 times dilution. After aerobic stabilization, inhibition was observed up to 400 times dilution. Other effects of the combined mill effluent sample were only significant below 40 times dilution. The bleaching effluent showed a mutagenic activity per t₉₀ comparable to kraft bleaching effluents. Other effects were only significant below 20 times dilution.     

Responses of cirripede larvae to light. I. Experiments with white light

The phototactic response of the nauplius larva of Balanus balanoides, B. crenatus and Elminius modestus shows darkadaptation; the response of the cyprid of B. balanoides shows both phototaxis and low photokinesis. The phototactic responses and the orientation of the cyprid to white light at settlement require an intensity of illumination slightly above 10^-5 lux. The ability to select a shaded position by cyprids of R. balanoides requires a higher intensity of 10^-2 to 10^-4 lux; hence a different mechanism may be involved. Barnacle larvae are sufficiently sensitive to be able to respond to light beneath the sea surface, even on cloudy, moonless nights.     

Combined effects of temperature and salinity on fed and starved larvae of the mediterranean mussel Mytilus galloprovincialis and the Japanese oyster Crassostrea gigas

The combined effects of temperature, salinity and nutrition on survival and growth of larvae of the Mediterranean mussel Mytilus galloprovincialis and the Japanese oyster Crassostrea gigas were studied over a period of 7 d in the laboratory. Ripe adults, collected in spring and summer 1987 from natural populations in the Bay of Arcachon, France, were induced to spawn. Larvae of both species were cultured at four temperatures (15°, 20°, 25° and 30°C), four salinities (20, 25, 30 and 35S) per temperature, and two levels of nutrition (fed and unfed) per temperature/salinity combination. The fed larvae received a mixed algal diet of 50 cells each of Isochrysis galbana and Chaetoceros calcitrans forma pumilum per microlitre. In both bivalve species, larvae survived over a wide range of temperature and salinity, with the exception of mussel larvae, which died at 30°C. Statistical analysis indicated that nutrition had the greatest effect on larval development, explaining 64 to 75% of the variance in growth of M. galloprovincialis and 54 to 70% in growth of Crassostrea gigas. Unfed mussel larvae displayed little growth. Compared with temperature, the effect of salinity was very slight. M. galloprovincialis larvae exhibited best growth at 20°C and 35S and C. gigas at 30°C and 30S.