Phylogeography of the ocean quahog (Arctica islandica): influences of paleoclimate on genetic diversity and species range

The ocean quahog, Arctica islandica (Linnaeus, 1767), is a commercially important bivalve found on continental shelves throughout much of the North Atlantic. To assess genetic subdivision in this species, we sequenced 385 nucleotides of the mitochondrial cytochrome b (cyt b) gene from 83 specimens collected from 12 localities between September 1998 and July 1999 (based on preliminary data, the Internal Transcribed Spacers, ITS, of the nuclear ribosomal repeat were not useful). The cyt b data delimited 11 haplotypes with 0.26 to 8.1% nucleotide difference (coded by 36 variable nucleotide positions) among them. Only three haplotypes were detected in 39 specimens collected along the USA coastline, compared to five haplotypes from nine Icelandic individuals. The western Atlantic populations ranging from Penobscot Bay (Maine, USA) to southern Virginia showed relatively low diversity and appeared genetically similar in that region. Based on the presence of shared haplotypes, AMOVA analyses, and phylogenetic reconstructions, Icelandic populations appear to be more genetically similar to western Atlantic populations than eastern Atlantic populations. Specimens from the Faroe Islands (n=4) show mixed affinities. These data are consistent with the hypothesis that a warm Holocene climatic optimum (ca. 7,500 years BP), and not glacial refugia, shaped the present-day genetic structure in A. islandica. Received: 18 January 2000 / Accepted: 26 June 2000     

Cell turnover in tissues of the long-lived ocean quahog Arctica islandica and the short-lived scallop Aequipecten opercularis

Cell proliferation and apoptosis were investigated in tissues of two bivalve species, Arctica islandica from the German Bight (age of bivalves: 33–98 years) and Iceland (7–148 years) and Aequipecten opercularis from the English Channel (2–4 years). High proliferation rates (10% nuclei dividing) and apoptosis in tissues of A. opercularis were in line with high-energy throughput and reduced investment into antioxidant defence mechanisms in the scallop. In contrast, cell turnover was slow (<1% nuclei dividing) in A. islandica and similar in mantle, gill and adductor muscle between young and old individuals. In the heart, cell turnover rates decreased with age, which indicates less-efficient removal of damaged cells in ageing A. islandica. Cell turnover rates, mass specific respiration and antioxidant enzyme activities were similar in German Bight and Iceland ocean quahog. Variable maximum life expectancies in geographically separated A. islandica populations are determined by extrinsic factors rather than by fundamental physiological differences.     

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.     

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.     

Molecular data from the 16S rRNA gene for the phylogeny of Veneridae (Mollusca: Bivalvia)

Veneridae, a cosmopolitan and ubiquitous family in world-wide littoral environments, lists more than 500 species, many of which are often subject to intensive commercial exploitation due to their numerical dominance in benthic communities. Historically, the family has been divided into 12 subfamilies by M. Keen and, even though this taxonomical arrangement was adopted for convenience and does not necessarily reflect genetic relationships, as suggested by Keen herself, this classification is still accepted. To contribute to clarify the systematics and phylogeny of Veneridae, the portion of the nucleotide sequence corresponding to domains IV and V of the gene encoding the large subunit of mitochondrial ribosomal RNA was analysed in 14 species representative of 10 genera belonging to 6 different subfamilies. The results obtained using the maximum-parsimony and neighbour-joining methods indicate that the current placement of the genera into subfamilies does not always reflect a natural subdivision. In addition, the six species of Tapetinae studied, though confirmed to be a monophyletic clade, do not exhibit a correct attribution at the genus level.     

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     

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.     

Le caryotype d'Aulacomya ater regia (Mollusca: Bivalvia: Mytilidae) des Iles Kerguelen

In 1983, specimens of the antarctic mussel Aulacomya ater regia Powell, 1957 were collected from the Kerguelen Islands and transported alive to the Zoological Station in Villefranche-sur-Mer, France. The chromosome complement of the mussel, as determined from branchial tissue, was found to consist of 13 pairs of chromosomes with terminal or subterminal centromeres. This species differs from the other species of Mytilidae studied up to the present in the number and the morphology of the chromosomes. A. ater regia is also unique among the known bivalves in displaying only terminal or subterminal centromeres.     

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.     

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.     

Some aspects of the thermal tolerance of Trichomya hirsuta (Mollusca: Bivalvia) of Eastern Australia

The upper temperature tolerance range for Trichomya hirsuta L. is described for acclimation temperatures of 10° to 32°C at 30% S. Changes in thermal resistance with time are described along with the incipient lethal temperatures. The eurythermicity for the species is 240.7 C°-squared. The ultimate upper incipient lethal temperature is 35.1°C. Definition of the incipient sublethal temperature is given, and is equal to 32°C. Diet effects are shown not to be significant in short-term lethal experiments, but indicate that unfed mussels reduce their resistance after prolonged exposure to high temperatures. Size was also shown to have no effect on resistance and tolerance. A comparison of the tolerance triangle of T. hirsuta with those of the temperate species Mytilus edulis and Mya arenaria revealed that the 24-h LT₅₀ minus 2.2 C° approximates the incipient lethal temperature. The eurythermicities of the temperate species are Mytilus edulis, 363 C°-squared; and Mya arenaria, 415 C°-squared.     

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.     

Sperm ultrastructure of the giant clam Tridacna maxima (Tridacnidae: Bivalvia: Mollusca) from the Great Barrier Reef

Using transmission electron microscopy, spermatozoa from a member of the Tridacnidae, or giant clams, are described for the first time and compared with spermatozoa of other bivalves, especially other heterodonts. The acrosomal vesicle of Tridacna maxima (Röding, 1798) is short (0.37?μm), blunt-conical, and exhibits a prominent basal ring. A narrow apical elaboration of the nucleus, the nuclear peg, projects deep into the basal invagination of the acrosomal vesicle. Aside from this specialization, the nucleus is a solid elongate-cylindrical structure (7.66?μm) that exhibits several small irregular lacunae. Four or occasionally three round-ovate mitochondria surround a pair of orthogonally-arranged, triplet-substructure centrioles. The proximal centriole is connected to a small indentation of the nuclear base by a thin layer of granular pericentriolar material, whereas the distal centriole is anchored to the plasma membrane by nine terminally-forked satellite fibres. The 9?+?2 pattern axoneme of the tail is continuous with the distal centriole. Comparison with other bivalves indicates a very close relationship between tridacnids and cardiids based on sperm ultrastructure. Specifically, the presence of a nuclear peg links Tridacna spp. with the cardiid genus Cerastoderma, but further information on the many unstudied genera is required to test the exact nature of this relationship. The sperm ultrastructure provides additional support for the recently proposed hypothesis that the Tridacnidae may be no more than a specialized subfamily of the Cardiidae.     

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.     

Seasonal variation in the balance between physiological mechanisms of feeding and digestion in Mytilus edulis (Bivalvia: Mollusca)

Seasonal variations in the balance between physiological mechanisms of feeding and digestion are considered among 45 to 57 mm shell length Mytilus edulis L. from southwest England. Mussels had been acclimated to standardised conditions of food availability in March, June and October, 1981. Results indicate that despite showing a negative relationship (P<0.001) with the efficiency of ¹⁵N absorption, coincident alterations of both ingestion rate (0.17 to 0.54 mg total Phaeodactylum tricornutum h^-1) and organic gut content (1.00 to 1.80 mg) represent significant mechanisms effecting the seasonal regulation of nutrient acquisition. Concurrent changes in absorption efficiency alone clearly exerted a relatively minor influence upon seasonal rates of absorption. Associated gut residence times of ¹⁵N-labelled material varied between 4 and 10 h. Furthermore, despite the ensuing deleterious effects, together with increased gut contents, absorption efficiencies among mussels that had been starved throughout acclimation were consistently maintained equivalent to those among individuals feeding normally. Such constancy was at least partially achieved by an increased residence time of material within diverticulae, relative to the digestive system as a whole. Finally, comparison of isotopic absorption efficiencies with those recorded for total organics and organic nitrogen has demonstrated differences of as much as 30% between gross and net uptake due, we suggest, to excretion of metabolically derived material within the alimentary canal.     

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.