Growth and shell morphology of three mytilidae (Bivalvia) species from the Sea of Japan

The growth and shell morphology of bivalve mollusks Crenomytilus grayanus, Mytilus coruscus, and Modiolus modiolus from the Sea of Japan are examined. The changes in body proportions and shell form in the ontogenesis of C. grayanus, M. coruscus, and M. modiolus are different even in the cases when the mussels develop in similar environmental conditions. Rapid growth shapes a well-streamlined and flat form of shell; slow growth leads to the formation of a massive and convex form. The parameters of the Bertalanffy growth curve for each species are calculated. The differences are discussed from the standpoint of functional morphology and spatial distribution patterns of mytilids in the coastal areas of the sea.     

A comparison of the reproductive cycles of Modiolus modiolus (L.), Cerastoderma (=Cardium) edule (L.), and Mytilus edulis L. in Strangford Lough,Northern Ireland

Summary This paper examines the reproductive cycles of three ecologically important marine bivalves-Modiolus modiolus (L.), Cerastoderma edule (L.), and Mytilus edulis L. in Strangford Lough, Northern Ireland over a period of almost five years.Whilst Modiolus does not appear to become sexually mature until it is several years old, Mytilus and Cerastoderma can reproduce in their first and second years of life respectively. In Cerastoderma and Modiolus sexual maturity is preceded by a period of rapid somatic growth.The subtidal Modiolus population remained in a more or less fully ripe condition virtually throughout the period of this investigation suggesting that this particular population lacked any marked cyclical reproductive activity. We interpret this as evidence of slow but almost continuous release of gametes throughout much of the year, a suggestion which is supported by recruitment data. A small intertidal population of Modiolus in Belfast Lough monitored over a period of two years exhibited a much more seasonal cycle. Here spawning occurred mainly during the autumn and winter. These data suggest that localised environmental factors are exceedingly important in controlling the annual reproductive cycle of this species. Cerastoderma from the mid-tidal sand flats ripened rapidly during the spring and spawned over a relatively restricted period in the summer. In Strangford Lough Mytilus occurs predominantly in the low-shore and while it spawns mainly in the spring and summer the annual cycle is considerably more protracted and variable than in Cerastoderma.Variations in the duration of the spawning periods in these bivalves can perhaps be explained in terms of both environmental stability and the immediate physical conditions experienced by these particular populations. The reproductive strategies exhibited by Cerastoderma, Modiolus, and Mytilus in Strangford Lough are considered in relation to population stability and to the different patterns of mortality which characterise these species in their respective local habitats.     

Electrophoretic investigation of systematic relationships in the marine mussels Modiolus modiolus L., Mytilus edulis L., and Mytilus galloprovincialis Lmk. (Mytilidae; Mollusca)

Genetic variation was assayed electrophoretically at 13–16 loci in Modiolus modiolus, Mytilus edulis, and Mytilus galloprovincialis. High genetic distance ( D ) values were observed between Modiolus modiolus and Mytilus edulis (1.516 ± 0.523) and between Modiolus modiolus and Mytilus galloprovincialis (1.564 ± 0.539), whereas the distance between Mytilus edulis and M. galloprovincialis (0.167 ± 0.118) was rather low. The systematic status ot Mytilus edulis and M. galloprovincialis is discussed in relation to these lindings and the genetic distance values are used to estimate divergence times which in turn are compared with paleontological estimates. The observations of high average heterozygosity in Modiolus modiolus, and high correlations of locus heterozygosity between taxa are discussed briefly.     

The sterol constituents and Δ5,7-sterol content of some bivalve molluscs

• 1.1. The bivalve molluscs Cerastoderma edula, Chlamys opercularis, Ensis soliqua, Modiolus modiolus, Mya arenaria, Mytilus edulis and Pecten maximus contained mixtures of C₂₆-, C₂₇-, C₂₈- and C₂₉-sterols. Cholesterol, 24-methylcholesta-5,22-dien-3β-ol and 24-methylene-cholesterol were the major sterols.
• 2.2. The sterols of Cerastoderma edula, Mya arenaria and Mytilus edulis contained 6–16% of cholesta-5,24-dien-3β-ol.
• 3.3. All the molluscs contained Δ^5,7-sterols in amounts ranging from 2 to 21% of the total sterols.
• 4.4. Cholesta-5,7-dien-3β-ol and 24-methylcholesta-5,7,22-trien-3β-ol were identified in Mytilus edulis. 25-Norcholesta-5,7,22-trien-3β-ol was detected in Modiolus modiolus.

     

Anaerobic survival potential of four bivalves from different habitats. A comparative survey

A comparative survey of the anaerobic survival potential of four different bivalve species and the interference of associated bacteria has been carried out. Individuals from both subtidal and intertidal environments were considered by selecting the following species: Mytilus edulis (subtidal epifaunal), Spisula subtruncata (subtidal infaunal), Macoma balthica (intertidal infaunal) and Cerastoderma edule (intertidal infaunal). Anaerobiosis was simulated in the laboratory by subjecting individuals to the following conditions: nitrogen atmosphere, air atmosphere and anoxic seawater incubation. Moreover, the effect of the antibiotic CA (chloramphenicol) was investigated, either as a pre-treatment of individuals kept under normoxic conditions for a week or directly added to the anoxic incubation media. According to survival performances of the individuals, intertidal animals that use to cope with tidal fluctuations in the coastline (emersion processes) had an extraordinary greater capacity to survive aerial exposure as compared to both nitrogen gas and anoxic seawater incubations most likely due to their capacity to perform aerobiosis at certain rate from atmospheric oxygen availability. Specifically, Macoma balthica enlarged its survival potential up to 24.8 days (LT(50)) under air exposure at 12 degrees C as compared to other specific treatments used here (4.9 days). The latter pattern was also observed, although in a much lower magnitude, for the other intertidal species Cerastoderma edule that survived 3.7 and 4.6 days (LT(50)) under nitrogen atmosphere and anoxic seawater incubation, respectively as compared to 9.5 days for emersed individuals. In contrast to the subtidal species, aerial exposure of both intertidal species led to a much higher survival performances than incubation of individuals in anoxic media with the presence of antibiotic. Survival capacity of the subtidal species Mytilus edulis and Spisula subtruncata was statistically similar under air and nitrogen atmospheres and anoxic seawater incubation. Then, subtidal species have a limited ability to air breathing as a conclusion of a similar survival in atmospheric and anoxic seawater incubations. Remarkably, M. edulis represented the only exception when considering longer-term survival capacity compared to the LT(50) values. Indeed, differences in LT(90) values for M. edulis were statistically different, values decreasing significantly from 19.7-19.9 days (under both nitrogen and air atmospheres) to 16.7 days when individuals are incubated in anoxic seawater. This may be due to the adverse effects of anaerobic bacteria that spontaneously proliferate within the static seawater incubations. As well as for S. subtruncata, possible aerobic processes under aerial exposure of mussels seemed to be not significant for the enlargement of its survival potential, since results obtained for both air and nitrogen atmospheres are similar. Pre-treatment with the antibiotic chloramphenicol caused survival capacity to increase by a factor of approx. 2 (M. edulis) and 34-44% (S. subtruncata). In contrast to intertidal species, the direct addition of the antibiotic to the incubation media caused the highest survival performances in both subtidal species. Habitat differences and species-dependent variability must be considered as significant sources of variation when studying the anaerobic performance of individuals using the most common experimental anaerobic techniques to test survival potential.     

Macrozoobenthos on an intertidal mudflat in the Danish Wadden Sea: Comparisons of surveys made in the 1930s, 1940s and 1980s

To assess whether long-term faunal changes have occurred on intertidal flats in the Danish Wadden Sea, results of faunal surveys in the Skallingen area during the 1930s, 1940s and 1980s were compared. Since the earlier studies, two species have disappeared (Scrobicularia plana andPetricola pholadiformis) and two have invaded the area (Tharyx killariensis andEnsis americanus). This is, however, not a local event as species changes have occurred on a larger scale (Wadden Sea region). Comparison of abundance data did not provide evidence of changes from the 1930s until the 1980s. Spatio-temporal fluctuations in two dominant species, the mudsnailHydrobia ulvae and the cockleCerastoderma edule chosen for closer examination, could be explained by natural processes. Growth data on cockles from the 1930s and the 1980s matched perfectly. During both periods, cockles showed a much lower growth rate than generally reported from the Wadden Sea, while at the same time they occurred at high densities (>2000 ind·m⁻²). Intraspecific competition among cockles is suspected as being a major cause of the low growth rates. Estimates of secondary production and P/B-ratios of cockles were also in general agreement during the 1930s and the 1980s when corrected for differences in the age structure of the cockle populations. With the possible exception ofMytilus edulis, which according to some observations has extended its range along the low-water line, the present comparison failed to demonstrate long-term faunal changes caused by increased eutrophication. This results is expected to be representative for intertidal flats not exposed to direct impacts from terrestrial run-offs and waste-water discharges.     

Now an empty mudflat: past and present benthic abundances in the western Dutch Wadden Sea

The benthic fauna of two areas in the western Dutch Wadden Sea, Posthuiswad and Staart van Schieringhals, was described in 1930–1960 and again between 1996 and 2005. Here, we document the changes. Whereas both areas formerly had high densities of species that biogenically structured the intertidal mudflats such as mussels Mytilus edulis and cockles Cerastoderma edule, by 1996 they had shown a tenfold decrease in the densities of molluscs, with no recovery till 2005. Although the number of species of polychaetes and crustaceans may not have changed much, their relative abundance did. Nowadays, more polychaete species are common than before. We briefly discuss whether the changes in benthic community composition could be due to industrial fishery practices or eutrophication effects.     

Modelling shellfish growth with dynamic energy budget models: an application for cockles and mussels in the Oosterschelde (southwest Netherlands)

Dynamic energy budget models for growth of individual cockles (Cerastoderma edule) and mussels (Mytilus edulis) are adjusted and calibrated to the Oosterschelde by formulating and parametrizing their functional responses using an extensive set of field observations. The resulting model predictions fit the observations satisfactorily. Results indicate that food quality and the importance of detritus as a food source are site-specific as well as species-specific. Despite these differences in their calibrated parameter values, both species show a very similar functional response. Compared with other systems, however, the functional responses of mussels in the present study are clearly higher than those of mussels in other systems. This may be explained by the absence of intra-specific competition in the measurement set-up that was used, and therefore supports the idea that the generally small functional response of M. edulis is caused by intra-specific competition.     

Interspecific comparison of the mechanical properties of mussel byssus

Byssally tethered mussels are found in a variety of habitats, including rocky intertidal, salt marsh, subtidal, and hydrothermal vents. One key to the survival of mussels in these communities is a secure attachment, achieved by the production of byssal threads. Although many studies have detailed the unique biomechanical properties of byssal threads, only a few prevalent species have been examined. This study assesses the variation in the mechanical properties of byssus in a broad range of mussel species from diverse environments, including intertidal and subtidal Mytilus edulis, Modiolus modiolus, Geukensia demissa, Bathymodiolus thermophilus, and Dreissena polymorpha. A tensometer was used to measure quasi-static and dynamic mechanical properties of individual threads, and several aspects of morphology were quantified. The results indicate that thread mechanical properties vary among mussel species, and several novel properties were observed. For example, of the species examined, D. polymorpha threads were the strongest, stiffest, least resilient, and fastest to recover after partial deformation. Threads of M. modiolus were characterized by the presence of two distinct yield regions prior to tensile failure. This comparative study not only provides insight into the ecological limitations and evolution of mussels, but also suggests new models for the design of novel biomimetic polymers.     

A comparison of variability in size dimensions of intertidal and subtidal mussels (Mytilus edulis) (Mollusca: Bivahia)

Variability in size dimensions of bivalve species has previously been claimed (1) to decrease with size and (2) to be greater for individuals in an intertidal habitat than in a subtidal habitat. Results from this study showed that the decrease in variability with size was an artifact of the techniques previously used, and that variability was the same for all sizes of Mytilus edulis. The previous study utilized an interspecific comparison to examine the effects of habitat on variability, which is considered unacceptable because differences in variability are likely to be a consequence of species differences rather than the effects of different habitats. Results presented here from the intraspecific comparison of M. edulis showed that variability was the same for mussels from an intertidal habitat and from a subtidal habitat.     

Divergence of unique DNA sequences of bivalve mollusks from Mytilinae subfamily (Bivalvia mytilidae)

The DNA-DNA hybridization method was used to determine the divergence degree of unique sequences of five bivalvia species belonging to the Mytilinae subfamily. The matrix of delta Tm values for heteroduplexes of unique sequences was found which made it possible to define three phylogenetic branches within the subfamily. Under non-stringent hybridization conditions (55 degrees C, 0.5M PB) the divergence between species of any two branches was about 14% of nucleotide substitutions. The hybridization of [3H] unique sequences of Mytilinae with DNA fragments of Modiolus modiolus, a representative of the closest relative Modiolinae subfamily, showed that the divergence rate of unique sequences in two phylogenetic lines of the Mytilus genusis higher than in the line of the Crenomytilus genus I0.35-0.23% and 0.1% of nucleotide substitutions per one million years, respectively). According to the matrix of delta Tm values for five species of Mytilinae and Modiolus modiolus, a phylogenetic tree was built reflecting the differences between the divergence rates in different branches.     

Mytilus galloprovincialis (Mollusca: Bivalvia) in a warm-temperate South African estuarine embayment

The distribution, population structure, condition and gonad indices, and the attachment strengths of the invasive Mediterranean mussel Mytilus galloprovincialis are described from the Knysna Estuary, South Africa. Mussels were found in the rocky intertidal at the mouth of the estuary and on all man-made hard substrata as far as 12?km from the mouth in 2013. However, mussel beds were found only in the lower reach of the estuary, where mussel densities were as high as 120 per 0.01?m². When compared to mussels from the embayment regions of the estuary, those at the estuary mouth, where wave action was high, were generally smaller, had significantly lower condition and gonad indices, and greater attachment strength. This suggests that mussels within the sheltered regions of the estuary invest more energy in growth and reproduction, and that this is one reason for their success as an invasive species in the Knysna estuarine embayment.     

Mollusc diversity in an Arca zebra (Mollusca: Bivalvia) community, Chacopata, Sucre, Venezuela

The diversity of a subtidal epifaunal mollusk community was studied from September, 1990 to September, 1991 in Chacopata, Sucre State, Venezuela. There were 40 species (24 bivalves and 16 gastropods). The diversity indexes (H' = 2.087, J' = 0.392, 1/D = 0.528) were low when compared with other tropical zones. Monthly diversity reached its maximum in September, 1990 (1.63 bits/ind.) and July, 1991 (1.60 bits/ind.); minimum diversity occurred in June, 1991 (0.52 bits/ind.). A Log series model applied to species abundance data showed a straight line with a diversity index alpha of 5.56. Of 40 species identified, the turkeywing Arca zebra was dominant (69% in number of individuals and 72% of biomass) followed by Pinctada imbricata, Modiolus squamosus, Chama macerophyla and Anadara notabilis. The predatory snails Phyllonotus pomum, Chicoreus brevifrons and Murex recurvirostris seemed to have trophic relationships with A. zebra. The total mean biomass in wet weight (469.20 +/- 263 g m-2, shell included) was high which indicates that A. zebra, a species with a rapid growth rate, occupies a central role in the assemblage as an efficient filter feeder that converts planktonic food into available biomass, supporting one of the most important fisheries in Venezuela.     

Secondary production of an intertidal mussel (Mytilus edulis L.) population in the Eastern Scheldt (S.W. Netherlands)

We monitored an intertidal mussel (Mytilus edulis L.) population between June 1981 and June 1982 in the Eastern Scheldt estuary (S.W. Netherlands). Density and biomass of the population remained relatively constant over the study period. The shell length growth was described by a Gompertz growth curve. The parameters of this equation were estimated from a log-log-modified Ford-Walford plot of the growth-ring data. The slope of the relationship between animal weight and shell length is season-dependent, mainly due to the spawning cycle in larger mussels.Secondary production is estimated with the growth rate method. In the calculated growth rates the change in slope of the length-weight relationship is incorporated, as well as differences in length growth rates between summer and winter. Secondary production amounts to 156 g AFDW m^–2a ^–1 (expressed per m² of mussel bank). P:B is 0.50 a^–1. The mussel productivity is probably a limiting factor for the density of overwintering Oystercatchers (Haematopus ostralegus).     

Macroparasite community in molluscs of a tidal basin in the Wadden Sea

We provide a quantitative inventory of macroparasites in intertidal molluscs from a tidal basin in the Wadden Sea (eastern North Sea). Gastropods and bivalves contained a species rich macroparasite community consisting of trematodes (26 species), turbellarians (1), nematodes (1), copepods (2) and polychaetes (1) in 3,800 host individuals from 10 host species. Highest parasite burdens were observed in the gastropods Hydrobia ulvae and Littorina littorea and in the bivalves Cerastoderma edule and Mytilus edulis. In contrast, only one parasite species and no trematodes were found in Crepidula fornicata. The parasite community in the molluscs was similar to other Western European localities but some parasite species showed obvious differences, related to the large-scale distribution of intermediate and final hosts. Parasitism seems to be a common phenomenon in molluscs of the Wadden Sea and hence the detrimental effects observed in experiments can be expected to frequently happen in the field.     

Adaptations of the Ribbed Mussel, Modiolus demissus (Dillvvyn), to the Intertidal Habitat

SYNOPSIS. This is a review of the behavioral, biochemical-physiological,and morphological adaptations of Modiolus demissus (Dillwyn)to the high intertidal habitat. The ribbed mussel, Modiolus,has fully exploited the behavioral adaptations within the limitsimposed by the bivalve body plan. Air-gaping is undoubtedlya significant behavioral adaptation which permits aerial respirationand penetration of the high intertidal zone. Modiolus fullyutilizes physiological-biochemical adaptations also. The musselis both eurythermal and euryhaline: the thermal range is atleast 56 °C and the salinity range is at least 70%, bothimpressive ranges for an organism which conforms to these environmentalparameters. The two primary adaptations made by Modiolus arean extensive tolerance to dehydration and a very high enzymethermostability. These two adaptations are resjxmsible for Modiolus'tolerance toward desiccation, variation in salinity, thermalstress, and possibly anaerobic conditions. There is no obviousmorphological adaptation. Thus, the physiological adaptations,coupled with air-gaping, are responsible for the vertical penetrationof Modiolus into the intertidal zone and its horizontal migrationover an extensive geographic range.     

Water loss from aerially exposed mussels

The valve activity and water loss from aerially exposed Mytilus edulis L. and Modiolus modiolus (L.) have been investigated in a series of laboratory experiments. Mytilus is capable of maintaining long periods of complete, or almost complete, valve closure when exposed in air, and this allows the retention of water in the mantle cavity and protects the tissues from evaporative water loss. Over periods of three days or more the amount of water lost from emersed Mytilus was found to be less than that retained in the mantle cavity at the beginning of exposure, suggesting that during normal periods of exposure the tissues are never directly subject to water loss. In contrast, Modiolus shows periods of gaping during which water loss is rapid due to drainage from the mantle cavity and evaporation from the tissues. The exposure of the tissues to air that results from gaping, makes the water loss susceptible to environmental influences of which wind was found to be the factor which caused the greatest increase in water loss and, as a consequence, an important factor in causing death through dehydration. The different abilities of Mytilus edulis and Modiolus modiolus to control water loss may be related to their intertidal distributions.     

Assessment of the state of the Japanese scallopMizuhopecten yessoensis in Alekseeva Bight (Peter the Great Bay, Sea of Japan) based on morphological and biochemical parameters

The results of an integrated examination of the state of the scallopMizuhopecten yessoensis in Alekseeva Bight (Peter the Great Bay, Sea of Japan) are presented. In mollusks of different ages, shell height was measured; in animals of commercial size (over 100 mm), some size and weight characteristics (annual increment of shell and adductor muscle and soft tissue weight) were determined. The morphology of the digestive gland and gills was studied. In the adductor muscle and digestive gland, the concentration of heavy metals (HMs) (Hg, Cu, Zn, Mn, Pb, Cd, Ni, Co, and Cr) was determined. In the digestive gland, metallothionein and reduced glutathione concentration was also determined, as was the activity of glutathione-dependent enzymes (glutathione peroxidase and glutathione reductase). In scallops collected outside Alekseeva Bight, the linear growth rate and adductor muscle weight were on average 1.3 and 1.7 times greater, respectively, than in those collected in the bight. In scallop organs, numerous histomorphological alterations were revealed: digestive cell vacuolization and hemocyte infiltration of the digestive gland, hyperplasia and vacuolization of the respiratory epithelium, and connective tissue hypertrophy in gill filaments. The biochemical parameters of scallops from Alekseeva Bight differed substantially from those of mollusks collected outside the bight. We conclude that one of the factors negatively affecting the state of theM. yessoensis population in Alekseeva Bight is the contamination of the bight with HMs, especially mercury. This is consistent with the results of chemical analysis of bottom sediments and tissues of two mytilid species,Modiolus kurilensis andCrenomytilus grayanus, specimens of which were collected in the bight together with the scallops [3].     

The intertidal zoning of cockles (<Emphasis Type="Italic">Cerastoderma edule</Emphasis>) in the Wadden Sea,or why cockle fishery disturbed areas of relatively high biodiversity

Recently, Kraan et al. (ICES J Mar Sci 64:1735–1742, 2007) reported that areas in the Wadden Sea selected by mechanical cockle dredgers were of greater zoobenthic diversity than areas that remained undredged. The present paper attempts to explain this observation, focussing on the elevation-related zoning of the macrobenthic fauna on tidal flats. Dense beds of cockles Cerastoderma edule occur in a specific intermediate intertidal-height zone where more species are present (and reach their maximal abundance) than in either lower or higher zones. This coincidence of preferred intertidal zones in many species, including cockles, may cause the higher species richness (and higher total macrozoobenthic biomass) of the fisher-chosen dredging areas as compared to other nearby non-dredged tidal-flat areas.     

Reproduction and fecundity of the musselmytilus edulis at helgoland (North sea)

The reproductive cycle of aMytilus edulis population in the lower tidal zone at the island of Helgoland from Spring 1980 to Summer 1981 is described. In both years the spawning period extended from the end of April until the end of June at water temperatures between 6° and 14°C. The gametes were built up again in autumn and most individuals were mature in February. Sex ratio did not differ significantly from 1:1. Fecundity, egg diameter and gamete weight of this and another population in the subtidal zone were assessed and size frequency distribution of shell lengths was established. Based on data of body weight prior to spawning, the following tendencies could be outlined: animals of the intertidal population had significantly smaller eggs than those from the subtidal area. Although the soft body biomass per unit area of the subtidal population was nearly double that of the tidal, their annual egg output was nearly the same (2.3 × 10⁹ eggs m^?2). This was achieved by a smaller size at the onset of sexual maturation (18 mm instead of 30 mm shell length) and a higher egg output at comparable shell lengths in the intertidal. In both populations, large animals contributed most to gamete production, although they did not necessarily dominate in biomass.