Genetic heterogeneity among New Zealand species of hydrothermal vent mussels (Mytilidae: Bathymodiolus)

Molecular systematic studies provide evidence for three new species of Bathymodiolus-like hydrothermal vent mussels (Bivalvia: Mytilidae) from relatively shallow waters (depth less than 750 m) associated with the Kermadec Arc off northern New Zealand. Mitochondrial COI sequences from the three putative new species differed substantially from those of other known bathymodiolin species from the Pacific and Indian Oceans. Population genetic analysis of one of these species (Bathymodiolus new species NZ-1) revealed heterogeneity in allozyme gene frequencies between samples collected from two seamounts about 50 km apart. Factors that might contribute to genetic differentiation between neighbouring seamounts are discussed.Communicated by M.S. Johnson, Crawley     

Ecological studies on intertidal New Zealand Sphaeromatidae (Isopoda: Flabellifera)

Ten sympatric rocky shore species and 3 brackish-water species of Sphaeromatidae were studied. Rocky shores were classified in terms of substrate and exposure to wave action. Although most of the rocky-shore species occur on all shore classes, vertical and horizontal distribution in each species, as well as densities, vary with substrate and wave action. A close relationship between distribution and breeding exists in the rocky-shore species; maximum differences in the vertical distribution of juveniles and adults occur, together with the highest reproductive capacities, in the species exposed to the greatest wave action. Tolerances to temperature, salinity and desiccation are wider in the species of brackish water and less exposed rocky shores, correlating with the greater variations in these conditions where wave action is reduced. Exoskeletons and pereopods are stouter in the species of more exposed shores, except where the microhabitat affords protection from wave action. In Isocladus armatus, geographic and seasonal variations in relative frequencies of colour forms correlate with differences in environmental temperatures. Under laboratory conditions, the colour forms show differences in tolerances to temperature and salinity.This work was carried out in the Zoology Department, University of Canterbury, New Zealand; the text was completed in Zoologisches Institut der Universität Kiel, Germany.     

Co-ordinated rhythms of digestion,absorption and excretion in Mytilus edulis (Bivalvia: Mollusca)

Sequential alternation of extracellular digestion in the stomach and intracellular digestion in the diverticula appears widespread among bivalves. The present study documents some physiological consequences of such processes in Mytilus edulis L. collected during 1981 from Whitsand Bay, Cornwall, England. Pronounced temporal fluctuations in faecal deposition are described that relate, in terms of amplitude and period, to both sinusoidal rhythmicity established for ammonia excretion and changes in the morphology of digestive tubules. Although at least partially synchronised among replicate groups of mussels, these cycles bore no consistent relationship with exogenous influences. Hourly fluctuation in the net absorption efficiency for nitrogen, as evidenced by the mean percentage ±2 SE, measured over 24 h sampling periods, was considerable (16.0±53.7, 49.3±10.9 and 52.8±6.6 for mussels acclimated in March, June and October, respectively). This variation in absorption derived from an inverse relationship between the percentage nitrogen within faeces and the rate of faecal egestion. Accordingly, peaks of faecal deposition presumably represented the pulsed remnants of intracellular digestion. Co-ordinated rhythms of digestion, absorption and excretion were thus evident in M. edulis. These processes displayed seasonally dependent periodicities of approximately 8, 3 and 4 h in March, June and October, respectively. It was concluded that, at least for M. edulis, this previously unquantified rhythmicity of physiological processes warrants careful consideration during assays commonly undertaken in the complication of nutrient and energy budgets.     

An ultrastructural study of Crassostrea angulata (Mollusca,Bivalvia) spermatogenesis

The cytoplasm of Crassostrea angulata (collected from the Atlantic coast of Northern Spain, Galiza, between 1992 and 1993) spermatogonia contains a few proacrosomal dense vesicles and intermitochondrial round granulo-fibrillar dense bodies. In primary spermatocytes, large cytoplasmic spaces appear occupied by large round and more scattered granulo-fibrillar dense bodies which are intermingled with cord-like dense structures and are in close association with mitochondria and the rough endoplasmic reticulum. As a result of Golgian activity, the number of dense proacrosomal vesicles increases and a flagellum arises from the centrioles. During the second meiotic division, several round dense bodies surrounding cord-like dense structures as well as dense proacrosomal vesicles, were observed between chromosomes. During spermiogenesis, no Golgian activity was noticed, and only the rough endoplasmic reticulum was observed in close association with proacrosomal vesicles. Similar results were observed for Ostrea edulis (collected from the Atlantic coast of southern Portugal, Algarve, between 1992 and 1993). Spermatozoa of both species are of the aquasperm type, and although their ultrastructure is quite similar, silver and phosphotungstic acid-chromic acid stainings enabled us to clearly distinguish both species at the ultrastructural level. The different morphological aspects of spermatogenesis in the Pteriomorphia and the use of silver and PTA stainings in the Mollusca are reviewed and discussed.     

Functional and environmental anaerobiosis in the razor-clam Ensis directus (Mollusca: Bivalvia)

In Ensis directus (Conrad) metabolic changes accompanying muscular activity and experimental anoxia were investigated. Octopine dehydrogenase, strombine dehydrogenase, alanopine dehydrogenase and lactic dehydrogenase were shown to be present in different parts of the body, but the activity of ODH always exceeded that of the other enzymes by far. Muscular activity connected with escape digging or evoked by electrical stimulation resulted in a decrease of stored phosphagen and in an accumulation of octopine. During recovery the normal concentrations of energy-rich phosphates were restored quickly, whereas octopine was removed more slowly. In experimental anoxia only minor quantities of L-and D-alanine, succinate, propionate, and acetate were formed. Quantitatively the most important endproduct was strombine, which rose linearly with the duration of anoxic incubation.     

Genetic differentiation in three Sepia species (Mollusca: Cephalopoda) from Galician waters (north-west Iberian Peninsula)

The systematics of the genus Sepia is not yet clear. Morphological evidence has led to S. officinalis Linnaeus, 1758 being considered as belonging to the subgenus Sepia sensu stricto, and S. orbignyana Férussac, 1826 and S. elegans Blainville, 1827 as belonging to the subgenus Rhombosepion. Samples of 30 individuals of S. officinalis and S. orbignyana from both sides of an oceanographic boundary off the north-west Iberian Peninsula, and a sample of S. elegans from the northern side, were collected in 1993–1994. Allozyme electrophoresis for 32 presumptive loci revealed low levels of genetic variability for the three Sepia species (mean expected heterozygosity estimates were <0.052). No significant differences in allozyme frequencies were detected among populations of either S. officinalis or S. orbignyana. The genetic identities (I) of S. officinalis and S. orbignyana (I=0.12) and of S. elegans (I=0.13) were significantly different from that of S. orbignyana and S. elegans (I=0.49). The former are typical of values for confamilial genera, and a new generic status is proposed for the latter two species, which become Rhombosepion orbignyana (Férussac, 1826) and R. elegans (Blainville, 1827).     

Levels of genetic polymorphism at two enzyme encoding loci in eight species of the genus Macoma (Mollusca: Bivalvia)

Levels of genetic polymorphism were surveyed at two enzyme loci (LAP, PGI) in 2 intertidal and 6 subtidal species of the bivalve genus Macoma living in the waters of the San Juan Islands (Washington and Canada). The temporal environmental variability-genetic variability hypothesis predicts that intettidal species should have greater levels of genetic polymorphism than subtidal species. This is not true for the genus Macoma. However, at the PGI locus, genetic polymorphism was proportional to niche breadth in both the intertidal and subtidal species, but only for intertidal species at the LAP locus. These results support the contention that temporal environmental variability is not necessarily important in maintaining genetic variability, but that environmental heterogeneity may select for increased polymorphism at some loci.     

Variability of the frequency of male neoteny in Ostrea puelchana (Mollusca: Bivalvia)

Ostrea puelchana (D'Orbigny), a larviparous oyster from the Atlantic coast of South America, comprises two types of individuals: (1) adults which live for several years and become sexually developed after one year, these show a rhythmic successive hermaphroditism with male, hermaphroditic and female phases; (2) neotenic individuals settled on the adults, which live for only about one year and function exclusively as males. Adult oysters were collected in February 1982 from a natural bed in the San Matias Gulf, Argentina, and were transferred to two sites 200 km to the south of their original habitat, where they were maintained under culture conditions. At the beginning of the study, 17% of adult oysters were observed bearing neotenic individuals. After two years, the number of adults in the female phase had decreased, and neotenic individuals had entirely disappeared. The presence of female-phase adults would seem to be necessary for the existence of neotenic males. Further analysis of the results revealed high adult density to be directly associated with greatly decreased numbers of adults in the female phase; this may play a role in the decrease of neotenic individuals.     

Freshwater Mussel,Lamellidens Marginalis (Lamarck) (Mollusca: Bivalvia: Unionidae) As an Indicator of River Pollution

Abstract

Analyses were made of heavy metals, manganese, nickel, copper, zinc and lead in water samples and soft body, shell and different tissues (gills, digestive glands, mantle and viscera) of the Unionid mussel, Lamellidens marginalis collected from two tributaries of the Cauvery river. Water samples from Station I contained higher concentrations of the metals than those from Station II. the concentration of metals in water at both stations were in the descending order: Mn > Zn > Pb > Ni > Cu. However, the concentrations of metals in the soft body were in the descending order: Zn > Mn > Pb > Ni > Cu at both stations in all size groups of mussels tested. the concentration of zinc maintained a linear relationship with the size of the mussels, but manganese showed a reverse trend. Small size (4-5 cm) mussels accumulated more manganese (105.5 μg.g^?1 dry wt.) than larger ones (7-8 cm; 6.5 μg.g^?1 dry wt.). Both young and old is accumulate the same level of lead, copper and nickel in the soft body. the order of concentrations of metals (Mn, Pb, Zn, Ni and Cu) in the shell of mussels from both stations coincided with the order of concentrations of background water except for lead. the accumulation of lead was higher in shell (30.4-36.2 μg.g^?1 dry wt.) than in soft body (6.4-12.0 μg.g^?1 dry wt.). the pattern of concentration of metals in the various tissues reveal that the digestive glands have greater ability than other tissues to concentrate most metals under study. the concentration factors for soft body, shell and different tissues are presented. the advantages in using the common mussel for biomonitoring of contaminants in water is also discussed.     

New observations of the gills of Placopecten magellanicus (Mollusca: Bivalvia), and implications for nutrition

The internal anatomy and microanatomy of the gill of Placopecten magellanicus Gmelin collected in May and November 1985 from Chamcook Bay, New Brunswick, Canada, was studied using thin-section light microscopy and transmission electron microscopy. Most of the spurs show no evidence of organic union, and hence do not participate in vascular exchange. However, the dorsal bend shows both ciliary and organic interfilamentar union. The internal structure and the hemocytes of the dorsal respiratory expansion are presented. The epithelium consists of three distinct cell types, bounded by apical microvilli. All regions of the gill contain an epithelial basal membrane, which is greatly convoluted in the interconnecting vessels of the dorsal respiratory expansion. The significance of these observations is discussed in relation to possible roles in respiration, transmembrane transport and nutrition. The apical surface of all ciliated cells is covered with an acellular matrix composed of clear spherical vesicles, which may serve a mechanical function for which mucus would be unsuited. The significance of the abundance of mucocytes on the abfrontal surface of the principal filaments is discussed in terms of the escape response of pectinid bivalves.     

New observations of the gills of Placopecten magellanicus (Mollusca: Bivalvia), and implications for nutrition

The organization, general anatomy, and surface microanatomy of all regions of the gills of a representive bivalve mollusc, Placopecten magellanicus Gmelin, were studied using stereo-microscopic, histological, and scanning electron microscopic techniques. Individuals were collected in May and November 1985 from Chamcook Bay, New Brunswick, Canada. In addition to correcting earlier accounts of this structure, a number of new observations are reported. The orientation of the ciliated spurs appears to be responsible for the sinusoidal arrangement of the gill filaments. Micrographs showing the structure of the dorsal respiratory expansion are presented. The entire abfrontal surface of the principal filament, including the dorsal respiratory expansion, is densely ciliated and mucosecretory. These characteristics may aid in the establishment of a respiratory current and in the prevention of gill damage during escape responses. All nonciliated regions of the gill filaments are covered with microvilli, thus greatly increasing the surface area of the gill. The feeding mechanism is discussed in relation to the dorsal and ventral ciliated tracts. Symbiotic ciliate protozoans are constantly dislodged from the gill filaments and transported via the ventral mucus string to the buccal region. The nutritional implications of these observations are discussed.     

Temperature-time relationships for survival of embryos and larvae of Mulinia lateralis (Mollusca: Bivalvia)

The influence of temperature over 8 time intervals on survival of cleavage stages, trochophore larvae, and straighthinge veliger larvae of Mulinia lateralis (Say) was investigated using a thermal gradient apparatus. There was a direct relationship of mortality with increased period of exposure. As the clams aged, temperature tolerance increased, with cleavage stages being most sensitive to higher temperatures and straight-hinge larvae least sensitive. Multiple-regression equations were developed to allow prediction of percentage mortalities under different conditions of temperature and time exposure. Entrainment of M. lateralis embryos and larvae in the cooling-water systems of steam-electric power plants should be as short as possible to keep mortality toa minimum.Contribution No. 550 of the Natural Resources Institute, University of Maryland, and Contribution No. 531 of the Virginia Institute of Marine Science.     

Peribuccal organs ofPlacopecten magellanicus andChlamys varia (Mollusca: Bivalvia): Structure,ultrastructure and implications for feeding

In order to better understand the structure of bivalve peribuccal organs and relate this to existing functional paradigms of their role in feeding, the labial palps of two scallop species,Placopecten magellanicus from the Bay of Fundy, Canada (1985 and 1986), andChlamys varia from the Bay of Brest, France (1986), were examined using histological techniques and electron microscopy. The ridged palp surface displays a uniformly dense ciliation with relatively few mucocytes; these are essentially concentrated in the region of the secondary ledge and may, through their secretory activity, determine the fate of particle masses in this area. The mucus secretions of the ridged palp surface are qualitatively different from those of the smooth palp surface. Mucocytes are much more abundant on the smooth palp surface, where it is suggested that their homogeneous secretions attenuate the potentially adverse effects of anteriorly-directed cleansing and swimming currents. Two other cell types are found in the palp epithelia: ciliated cells, which are very numerous on the ridged surface and relatively rare on the smooth surface, and non-ciliated epithelial cells, which are very numerous on the smooth surface and rare on the ridged surface, where they are confined to the palp margin. In addition to the mechanical role of the ciliated cells and mucocytes, the ultrastructural characteristics of the ciliated and non-ciliated epithelial cells indicate a dichotomy of function between the ridged and smooth surfaces. The ridged surface epithelial cells present an ultrastructural specialization in the absorption of dissolved and colloidal matter, suggesting an accessory nutritive role, whereas the smooth surface simple epithelial cells show signs of active molecular synthesis. No specialised sensory cells were observed on the ridged surface; it is therefore not yet possible to conclude whether the labial palps are capable of selection based on individual particle characteristics.     

Advanced age for sexual maturity in the ocean quahog Arctica islandica (Mollusca: Bivalvia)

A study of gonadal state in 39 small specimens of Arctica islandica Linné collected in April 1976 from the Middle Atlantic shelf of North America suggests that sexual maturity is reached at a later age than has been reported for other shelf bivalves. Eight of the specimens had gonads in the undifferentiated state. On the basis of annual internal growth banding in the shells of these immature specimens, age ranged between 4 and 14 years. Average age was 9.38 years (standard deviation, s,= 3.54). Shell length ranged from 24 to 47 mm (average 38.85 mm, s=9.25). Differentiated gonads were found in specimens as young as 6 years. Age of maturity shows a wide range and may be dependent upon growth rate and locality.     

Rhythmicity of larval release in three species of intertidal brachyuran crabs (Crustacea: Brachyura) from Inhaca Island (Mozambique)

 A study of rhythmicity of larval release in three species of intertidal brachyuran crabs, based on laboratory and field experiments, was undertaken at Inhaca Island, southern Mozambique, using Leptodius exaratus and Macrophthalmus grandidieri from December 1994 to January 1995, and Arcotheres palaensis from April to July 1995. L. exaratus and M. grandidieri showed a semi-lunar cycle in larval release. The release of larvae for L. exaratus, a species having conspicuous larvae, occurred in the first half of the night, after the post-crepuscular high tide, which suggests maximisation of protection of larvae from visual predation. The larval release activity matched the late spring and early neap tides. Results from the field were similar to those from the laboratory. M. grandidieri, having inconspicuous larvae, did not show a pattern related to the light–dark cycle and hatched during spring tides (around full and new moons) to maximise larval dispersion. A. palaensis, living inside the host mussel which inhabits the lowest section of the intertidal zone, did not show a relation with moon phase, tidal or light–dark cycles. Received: 16 February 1999 / Accepted: 8 December 1999     

Burrow morphology,biometry, age and growth of piddocks (Mollusca: Bivalvia: Pholadidae) on the south coast of England

Biometry and growth of three piddock species Pholas dactylus, Barnea candida and B. parva, from chalk and clay substrata were investigated between 1999 and 2000 at five low shore locations along the south coast of England. Piddock burrow shape was significantly different (P<0.01) amongst the locations. Burrows at Lyme Regis showed the largest (height/maximum diameter) ratio (4.86±2.00) whilst those at Compton the lowest (3.73±1.62). Using the method of Bhattacharya, the population structure of P. dactylus, B. candida and B. parva was separated into eight, three and five modal size classes, respectively. Age and shell growth were determined from the number and spacing, respectively, of annual growth lines present in acetate peel replicas of shell sections. The von Bertalanffy growth (VBG) equation fitted the size at age data obtained for P. dactylus and B. candida (L and K were 79.3±13.8 mm and 0.0011±0.22 and 29.6±1.5 mm and 1.17±0.47, respectively) whilst the size at age data for B. parva were linearly related and did not fit the assumptions of the VBG equation. Male and female gonads were mature and piddocks competent to spawn between June and September 1999, with settlement of juveniles observed between November 1999 and February 2000. A significant relationship between burrow aperture diameter and age of the occupant piddock was established for the three species. Burrow morphology and spatial distribution of burrows were influenced by substratum hardness and population density. Based on estimates at Lyme Regis, piddocks are capable of removing up to 41% of the shore substratum to a depth of 85 mm over their lifespan (12 years), significantly compromising the structural stability of the soft rock shores they inhabit and contributing to bioerosion.Communicated by O. Kinne, Oldendorf/Luhe     

First evidence of gender-related differences in immune parameters of the clam Ruditapes philippinarum (Mollusca, Bivalvia)

For the first time, gender-related differences in some important functional parameters of haemocytes from the clam Ruditapes philippinarum have been studied. Clams were collected during the pre-spawning phase, when sex can be distinguished by microscopic observation of gonadal smears, in order to ascertain whether the two sexes reach the stressful spawning period with different degrees of immunosurveillance. Total haemocyte count (THC), haemocyte size frequency distribution, capability of haemocytes to assume the vital dye neutral red (NR, indicative of endocytotic activity), lysozyme-like and acid phosphatase activities in both haemocyte lysate (HL) and cell-free haemolymph (CFH), and superoxide dismutase (SOD) and catalase (CAT) activities in HL were all evaluated. No differences in THC values were found between the two sexes, but differing haemocyte size frequency distributions were observed: the fraction of larger haemocytes (7–9 μm diameter, 200 fl volume) was markedly increased in females and that of smaller haemocytes (<5 μm diameter, <200 fl volume) markedly increased in males. Interestingly, when haemocytes were observed under the light microscope, a significantly higher percentage of granulocytes was found in haemolymph from females, compared with that of males. Significantly increased NR uptake was recorded in female haemocytes. No significant variation in lysozyme-like activity was observed in HL, whereas in CFH, enzyme activity was significantly higher in females. HL acid phosphatase activity was significantly higher in males with respect to females; no significant variations in enzyme activity were observed in CFH. HL SOD and CAT activities were significantly higher in females than in males. Overall, on the basis of the cell parameters studied here, gender-related differences in immune parameters were found in R. philippinarum, indicating that females have more active haemocytes than males during the pre-spawning period.     

Gill function and particle transport in Placopecten magellanicus (Mollusca: Bivalvia) as revealed using video endoscopy

The technique of endoscopic video observation was used to study feeding processes of Placopecten magellanicus (Gmelin), collected from Bull Arm, Newfoundland in August 1991 and 1992, under near-natural feeding conditions. The fate of captured particles depended on the extent of ingestive or handling capacity saturation. Under low (1 to 10 particles l^-1) to medium (10 to 20 particles l^-1) particle concentrations, most particles were incorporated in continuous anteriorly directed slurries in the dorsal ciliated tracts of the gill arch and dorsal bends. As particle concentration or exposure time to the lower particle concentrations increased, four endogenous mechanisms of ingestion volume control were increasingly observed: (1) rejection of dense mucus-particle masses from the principal filament troughs onto the ventrally beating cilia and associated currents of the ordinary filament plicae, counter to and below the incoming pallial current maintained by the principal filament cilia; (2) intermittent stopping of the anteriorward flow in the dorsal ciliated tracts; (3) reduction or cessation of input from the principal filaments to the dorsal ciliated tracts; (4) detachment of the dorsal bends from the mantle to establish a shunt from the infrabranchial to the suprabranchial cavity. Chemical and histochemical tests of purified fluid withdrawn from the dorsal ciliated tracts indicate that mucus is present at all particle concentrations. Mucus therefore participates both in normal feeding and in ingestion volume control on the bivalve gill, although different mechanisms, and types of mucus, effect transport of material in the dorsal (feeding) and ventral (cleaning) ciliated tracts.     

Seasonal and tidal changes in the length of the crystalline style intertidally living Macoma balthica (Mollusca,Bivalvia)

Samples of Macoma balthica were collected during 1978 and 1979 from the westernmost part of the Dutch Wadden Sea. The length of the crystalline style in the stomach of this intertidal bivalve was studied through tidal and annual cycles. The length of the style was ca. half the shell length and changed slightly with the season. Maximum lengths were recorded in spring, the season of high food intake, and minimal values in autumn and winter. Changes in the length of the style also exhibited a tidal cycle, but with two maxima and two minima. Tidal fluctuations in style length amounted to ca. 5% only. The influence of enzymes and waste products from the digestive diverticula, the food intake as well as the influence of the pH on the length of the style are discussed.Communication No. 401 of the Delta Institute     

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.