Genetics and shell morphology in a hybrid zone between the hard clams Mercenaria mercenaria and M. campechiensis

Populations of Mercenaria mercenaria (L.) from South Carolina, USA, and M. campechiensis (Gmelin) from the Gulf of Mexico, Florida, USA, were sampled in 1987. The two species differed at all of seven enzyme loci tested, as well as in the thickness of shell ridges and nacre color. The difference in lunule shape was not great, although differences in relative shell width, shell weight, and lunule size make morphometric discrimination between the species possible. Shell ridges, nacre color, and multivariate morphometrics in a sample of clams collected from the Indian River Lagoon on the Atlantic coast of Florida in 1985 do not assort independently. Individuals with thick ridges, white nacre, and/or campechiensis-like morphometrics have significantly different allele frequencies at most enzyme loci from individuals with thin ridges, purple nacre, and/or mercenaria-like morphometrics. The deviations are in the direction predicted from the analysis of the allopatric populations of M. mercenaria and M. campechiensis. M. mercenaria outnumber M. campechiensis in the Indian River sample, but the majority of the clams seem to be hybrids.Contribution No. 235 from South Carolina Marine Resources Center     

Mitochondrial gene variation in Mercenaria clam sibling species reveals a relict secondary contact zone in the western Gulf of Mexico

We investigated phylogeographic relationships among American Mercenaria taxa by assessing variation in a 444 nucleotide fragment of the mitochondrial 16S ribosomal gene in clams sampled from four representative sites in January to November 1994. Three of these sites were in the Gulf of Mexico, one was on the Atlantic coast in South Carolina. Direct sequencing of this amplified gene fragment in 85 individuals revealed 21 haplotypes. Phylogenetic analyses consistently resolved this variation into three well supported clades, and within-clade genetic divergence levels were markedly lower than among-clade values. One of the clades, A, was taxon-specific, in that it solely and exclusively contained specimens of M. mercenaria (Linnaeus, 1758) sampled in South Carolina. The other two clades, B and C, were the most divergent and both encompassed specimens of M. campechiensis (Gmelin, 1791) and of M. campechiensis texana (Dall, 1902), sampled from the three Gulf of Mexico sites. Clade B was found at high frequencies at all three Gulf sites, whereas Clade C occurred at low frequencies at two western Gulf sites. We interpret this pattern as resulting from the secondary contact and introgression of two allopatrically differentiated Mercenaria taxa in the western Gulf of Mexico. Clade C haplotypes may represent relict mitochondrial lineages from original Gulf Mercenaria spp. populations that predate massive mitochondrial introgression by M. campechiensis. We further propose that the M. campechiensis texana nuclear genome is a mosaic, heavily weighted toward M. campechiensis, but containing some relict alleles inherited from the precontact population, especially those governing shell characteristics, which may be adaptive in cohesive sediments of bays and estuaries in the northwestern Gulf of Mexico.     

Sclerochronological records of temperature and growth from shells of Mercenaria mercenaria from Narragansett Bay,Rhode Island

Annual internal growth increments in shells of hard clams (Mercenaria mercenaria) provide an accurate record of both growth history and some types of environmental change. These increments were used to determine age and growth rate of hard clams collected in 1984–1985 from ten sites in Narragansett Bay, Rhode Island, USA, and to assess geographic variation in growth within the bay. The regional comparisons were facilitated by modeling the clam growth using the von Bertalanffy equation and the parameter of Gallucci and Quinn. The optimum region for hard clam growth was near the head of the bay, with areas further north (Providence River) and south (lower bay) not as productive. The relationship between size and age was similar to that reported for hard clams from northern New Jersey. Using dendrochronological techniques, variations in growth were analyzed on a temporal (year-to-year) basis. The comparative longevity of the hard clams (individuals with 40 annual bands were encountered) demonstrated that sclerochronologies of several decades were possible. A 26 yr growth record based upon 100 individuals was established, covering the years 1959–1984. Significant temporal variations occurred in bivalve growth, and a broad trend of increasing growth indices over the last two decades of the record was noted. The yearly standardized growth index values were highly and positively correlated with mean annual water temperatures in Narragansett Bay for the same time interval. The incremental shell records of these bivalves provide a quantitative indication of marine climatic (temperature) variability and growth which can be used in hindcasting the effects of natural or anthropogenic environmental perturbations.     

Effect of the hard clam Mercenaria mercenaria on benthic fluxes of inorganic nutrients and gases

The effect of the hard clam Mercenaria mercenaria (L.) on the exchange of dissolved nutrients (silicate, phosphate, ammonium, nitrate+nitrite) and gases (oxygen, carbon dioxide) across the sediment-water interface was examined in 1983 and 1984 using experimental mesocosms (13 m³), designed to simulate a shallow coastal ecosystem, that allow for reciprocal biogeochemical interactions between water column (5 m) and bottom sediments (30 cm deep). Benthic, fluxes, measured during a spring-summer warming period, were compared for mesocosms maintained either with added M. mercenaria (16 per m², treatment) or without M. mercenaria (control) as a component of the benthic community. Fluxes were within the range observed for the (control) community in situ in mid-Narragansett Bay and exhibited the pattern of increasing flux with increasing temperature observed in nature. For dissolved inorganic (DIN=ammonia+nitrate+nitrite), analytical problems allowed comparison of fluxes only at the higher temperatures (17° to 22°C); average DIN fluxes were 57% higher in mesocosms with clams. Fluxes of other nutrients and gases were modelled as exponential functions of temperature (9° to 22°C) using a linear regression of the natural logarithm of flux upon temperature in both treatment and control situations. Differences between regression slopes and intercepts for conditions with and without clams were assessed by analysis of covariance. Slopes of the linearized temperature-flux relationships were not significantly different between treatments for any flux measured, suggesting that presence of the clams did not alter the basic relationship of flux and temperature normally observed with this community. For oxygen consumption and silicate production, the regression intercepts were significantly higher in the clam treatment, suggesting that the level of flux was generally elevated in the presence of M. mercenaria by about 20% (oxygen) and 86% (silicate) at any given temperature. Data for carbon dioxide and phosphate showed similar trends to oxygen and silicate, but the variance in fluxes was larger and regression intercepts for the two treatments were not significantly different.     

Effect of tidal currents,seston, and bottom sediments on growth of Mercenaria mercenaria: results of a field experiment

Tidal currents, seston, and sediments separately influence growth of the hard clam, Mercenaria mercenaria, but it is uncertain how these factors may interact. A 3×3 factorial field experiment, carried out in Great Sound, a coastal lagoon in Southern New Jersey, USA, between May and September 1986, determined the relative effects of three sediment types and three site-specific seston/tidal current regimes on the individual growth of M. mercenaria. Analysis of variance of the change in shell length after 15 wk (differences in initial and final lengths) demonstrated a significant difference (P=0.0064) in growth among sites, but no significant differences (P=0.1331) for growth in different sediments, although trends were evident. Effects of sites were independent of sediment type (P=0.2621). Shell growth rates differed by 10.7% between the slowest and fastest sites, but only differed by 5.7% between sediment types, with fastest growth in sand and slowest in mud. Tidal current speeds and four measures of seston (chlorophyll a, particulate inorganic and organic matter, PIM and POM, and energy content) were measured >20 times in near-bottom waters at each site. Horizontal fluxes of POM exhibited higher correlation coefficients with growth rates, than did seston concentrations or current speeds alone. We attribute significant site differences to differences in horizontal seston fluxes fluxes among sites. We suggest that horizontal seston fluxes may be a major factor affecting individual growth of suspension-feeding bivalves.No. 216 of the Jackson Estuarine Laboratory Contribution Series     

Influence of algal and suspended sediment concentrations on the feeding physiology of the hard clam Mercenaria mercenaria

Short-term laboratory feeding experiments were conducted to determine the response of the hard clam Mercenaria mercenaria (L.) (32 mm in mean shell length) to increasing sediment concentrations. Clams were fed mixed suspensions of Pseudoisochrysis paradoxa (50 and 150 cells l^-1) and bottom sediments (0 to 44 mg l^-1). Algal ingestion rate deelined with increasing sediment loads. This resulted primarily from a reduction in clearance rate, which declined by 0.08 l h^-1 g^-1 (1.3%) for every 1 mg l^-1 increase in sediment load. This reduction was of similar magnitude for juvenile (13 mm) clams. At the algal concentrations tested, pseudofaeces production was intermittent and inconspicuous below about 10 mg silt l^-1. Loss of algae in pseudofaeces increased with increasing sediment loads; however, even at the highest silt and algal concentrations, clams lost a maximum of only 18% of the algae cleared from suspension. Thus, pseudofaeces production is not expected to cause significant loss of algal food at the sediment concentrations normally encountered in the natural environment ( ca 40 mg silt l^-1). Absorption rate of total organic matter remained constant, at least up to silt concentrations of 20 mg l^-1. Experiments using dual ⁵¹Cr:¹⁴C-formaldehyde-labelled sediment indicated that clams were able to counteract the dilution of algae by absorbing a considerable fraction (21 to 22%) of detrital sedimentary organics. Absorption efficiency of pure P. paradoxa ranged from 82% at 50 cells l^-1 to 58% at 150 cells l^-1. Integration of physiological rate measurements suggests that at moderate to high algal concentrations (300 g Cl^-1), growth improvement by the addition of silt, documented in mussels, surf clams and oysters, is unlikely to occur in M. mercenaria. It is suggested that a suspension-feeding bivalve's success in maximizing its energy gain in a turbid environment depends on the combination of two features: a high selection efficiency and a high rate of pseudofaeces production. It is proposed that species which regulate ingestion primarily by producing pseudofaeces are better adapted to cope with high suspended sediment loads than species such as M. mercenaria, which control ingestion mainly by reducing clearance rate.Contribution No. 451 from the Marine Sciences Research Center, State University of New York at Stony Brook, USA     

Mortality and growth of juvenile hard clams Mercenaria mercenaria during brown tide

Blooms of the picoplankton-sized alga Aureococcus anophagefferens Hargraves et Sieburth, also known as "brown tide," frequently recur in Great South Bay, New York, USA. A field study compared mortality and growth rates of juvenile hard clams, Mercenaria mercenaria (L.), raised at a site that experienced a bloom of A. anophagefferens (>1᎒⁶ cells ml^-1) to clams grown in a nonbloom site from May to September 2000 at two sites on the north and south shores of Long Island. High (67%) cumulative clam mortality was observed for clams raised at the brown tide site (south shore) during the bloom (June-July), and individuals exhibited no measurable growth during that period. After the bloom subsided (mid-July), clams that survived the brown tide suffered low (<5%) cumulative mortality over the remaining sampling period (July-September). These survivors exhibited rapid growth for 4 weeks shortly after the bloom's decline. Subsequently, their growth slowed to rates comparable with those observed at the nonbloom site (north shore). By contrast, a low (<5%) cumulative mortality was observed for M. mercenaria raised at the site with no brown tide, and positive growth occurred throughout the study. These data suggest that some juvenile clams not only survive brown tide but also recover following the bloom.     

Growth of juvenile Mercenaria mercenaria and the effect of resuspended bottom sediments

The influence of silt on growth of juvenile hard clams Mercenaria mercenaria (L.) (9 mm in mean shell length) was investigated in the laboratory using mixed suspensions of algae (50x10⁶ Pseudoisochrysis paradoxa cells l^-1) and fine-grained bottom sediments (0 to 44 mg l^-1). Growth rates, expressed as percent increase in ash-free dry tissue weight, were not significantly affected by sediment concentrations up to 25 mg l^-1. Significant reduction in growth (by 16% relative to controls fed only algae), and condition of clams, occurred at 44 mg silt l^-1. The results of the 3-week growth experiment agree well with predictions made in an earlier study by integrating results of shortterm physiological measurements. Growth rates obtained with experimental algal-silt diets at 21°C (2.6 to 3.3% increase in dry tissue weight d^-1) were comparable to those determined at ambient concentrations of Great South Bay particulates at 20°C (0.9 to 4.0% d^-1). Levels of particulate inorganic matter in seawater from Great South Bay, New York, exhibited pronounced daily changes, and ranged from 6 to 126 mg dry weight l^-1. Growth enhancement by the addition of silt to an algal diet, reported in mussels, surf clams and oysters, was not found in M. mercenaria. It is suggested that these three species are better suited than hard clams for culturing efforts in inshore turbid waters above uncompacted, muddy bottoms.Contribution No. 452 from the Marine Sciences Research Center, State University of New York at Stony Brook, USA     

The persistence of chronically accumulated hydrocarbons in the hard shell clam Mercenaria mercenaria

Hard shell clams, Mercenaria mercenaria, from the chronically polluted environment of the Providence River, USA, were transferred to a clean laboratory system located in lower Narragansett Bay, Rhode Island. The hydrocarbon contents of these transplanted clams were monitored for 120 days after transfer. After this period only slight depuration had occurred. A 41.9 g g wet weight^-1 average initial hydrocarbon burden decreased to 29.3 g g^-1, a decrease that became statistically significant (P<0.1) only after 120 days. At most, only 30% of the clams' hydrocarbons were lost. These findings indicate that the duration of the exposure period and the chemical nature of the source hydrocarbons are both instrumental in determining the persistence of these compounds in filter-feeding bivalves. Chronically accumulated petroleum hydrocarbons are strongly retained by the organisms and are only very slowly depurated.     

Effects of suspended and sedimented clays on juvenile hard clams,<Emphasis Type="Italic"> Mercenaria mercenaria</Emphasis>, within the context of harmful algal bloom mitigation

Increased interest in using ecologically inert clays to flocculate, sediment, and thus mitigate harmful algal blooms at nearshore mariculture sites has prompted studies on the effectiveness of this method on prolific blooms, such as those caused by the neurotoxic dinoflagellate Karenia brevis in the Gulf of Mexico. Potential repercussions of this control strategy revolve around the increased flux of suspended particles to the benthos. Juvenile suspension-feeding bivalves are potentially vulnerable as they could suffer burial, a decrease in clearance rates, and/or an increase in pseudofeces production in response to suspended clay, leading to reduced growth and delay in attaining size refuge from predators. Here we assess lethal and sublethal effects on juvenile hard clams, Mercenaria mercenaria, in a 2-week flume application of phosphatic clay to simulated blooms of the nontoxic dinoflagellates Heterocapsa triquetra and Prorocentrum micans. Flow regimes simulated two contrasting, worst-case field conditions where (1) low flow allowed complete sedimentation and formation of a benthic sediment layer, and (2) high flow allowed complete particle suspension. No clam mortalities occurred in either treatment. The sedimentation treatment showed variable growth inhibition in shell and/or tissue, but effects were not apparent compared to controls (no sediment layer), and clams rapidly resumed siphon contact with the overlying water column. In contrast, a strong growth effect (~90% reduction in shell and tissue growth) occurred in particle-suspension trials compared to no-clay controls. These results suggest that repeated clay applications in the field are likely more detrimental to clams in a high-energy environment leading to prolonged in situ resuspension of clay than in an environment favoring sedimentation.Communicated by R.J. Thompson, St. Johns     

Toxicity of ammonia,nitrite ion,nitrate ion,and orthophosphate to Mercenaria mercenaria and Crassostrea virginica

The 96-h lethal tolerance limits of the hard clam Mercenaria mercenaria (Linné) and the oyster Crassostrea virginica (Gmelin) to ammonia, nitrite ion, nitrate ion, and orthophosphate were defined. Sublethal effects of the chemicals upon the rates at which the shellfish removed algal cells from suspension were also studied. In comparison with other marine and aquatic species which have been studied, hard clams and oysters are extremely tolerant.     

Heritability of juvenile growth for the hard clamMercenaria mercenaria

The heritability of juvenile growth in the hard clamMercenaria mercenaria (L.) sampled from Wellfleet Harbor, USA, in 1987, was analyzed in two experiments using standard quantitative genetic methods. In the first experiment, culture density was not controlled and was found to have a significant impact upon the non-additive genetic component of growth, possibly through genotype-environment correlation. Variable clam density appeared to have little impact on the additive genetic variance component. Estimates of the narrow sense heritability, calculated from the additive genetic component, ranged from 0.72 (±0.32) to 0.91 (±0.17). Culture density was controlled in the second experiment. The narrow sense heritability estimate from this experiment was 0.37 (±0.13). These estimates suggest that selection, either natural or artificial could bring about rapid change in the mean phenotype(s) of juvenile growth characteristics in hard clams.Contribution no. 793 from the Belle W. Baruch Institute for Marine Biology and Coastal Research     

Clam predation by whelks (Busycon spp.): Experimental tests of the importance of prey size,prey density,and seagrass cover

In 57 l-m² samples within a meadow of Halodule wrightii in Bogue Sound, North Carolina, USA, densities of the clams Mercenaria mercenaria and Chione cancellata were positively associated with seagrass cover. Where seagrass was experimentally removed, marked individuals of both clam species exhibited high rates of mortality in fine sand sediments during two successive experiments spanning 13 months. In the unaltered (control) seagrass meadow, M. mercenaria density remained constant over 13 months and C. cancellata density declined at a slower rate than in the unvegetated plots. Seagrass provides these clams with a refuge from whelk (Busycon carica, B. contrarium, and B. canaliculatum) predation, the major cause of mortality and population decline in experimentally unvegetated plots. In 2 factorial field experiments in unvegetated substratum in which densities of M. mercenaria and C. cancellata were varied independently, first over 5 levels (0 X, 1/2X, 1 X, 2 X, 4 X) and subsequently over 4 levels (0 X, 1/4 X, 1 X, 4 X), there was no repeatable intra- or interspecific effect of density on percent survival, or on the rate of any mortality type. Whelk predation fell preferentially on larger size classes of both species, whereas factors which contribute to clam disappearance usually acted more intensely on smaller sizes. Experimental exclusion of large predators by caging demonstrated that even in unvegetated substratum survivorship of both clam species was high in the absence of whelks and other predators. Individuals of C. cancellata live closer to the sediment surface than those of M. mercenaria, which may explain why seagrass does not serve as effectively to protect them from whelk predation. The mechanism of whelk inhibition may depend upon sediment binding by the H. wrightii root mat, which produces a demonstrable decrease in the physical penetrability of surface sediments.     

Invertebrate introductions in marine habitats: two species of hydromedusae (Cnidaria) native to the Black Sea,Maeotias inexspectata and Blackfordia virginica,invade San Francisco Bay

The hydrozoans Maeotias inexspectata Ostroumoff, 1896 and Blackfordia virginica Mayer, 1910, believed to be native to the Black Sea (i.e. Sarmatic) and resident in a variety of estuarine habitats worldwide, were found as introduced species in the Petaluma River and Napa River, California, in 1992 and 1993. These rivers are mostly-estuarine tributaries that flow into north San Francisco Bay. Both species appeared to be well-established in this brackishwater habitat. Salinities at the collection sites were about 11 during the summer, rising to nearly 20 in the early autumn and falling to near 0 in the winter. Large numbers of all sizes of both species of medusae were observed and collected, indicating that the hydroid stages of the life cycles of the two are also well-established in these rivers. In the Petaluma River, populations of both species were at maximum in late July, with numbers of individuals declining through August and into September; the Napa River was sampled only in October, and at that time only B. virginica was found. Examination of full guts of M. inexspectata and B. virginica medusae revealed that both species had fed nearly exclusively on small crustaceans, principally barnacle nauplii, copepods and their eggs and nauplii, and crab zoea larvae (M. inexspectata only). All the M. inexspectata medusae were males, indicating that the population has probably developed from the introduction of perhaps only a single male polyp or polyp bud. In spite of its inability to reproduce sexually, this population appears to be maintained by the prodigious ability of the polyp to bud and reproduce asexually, and is fully capable of invading additional low-salinity habitats from its present Petaluma River site. Male and female B. virginica medusae were collected in both the Petaluma River and the Napa River, indicating that B. virginica may have been introduced by either the polyp or medusa stage (or both), but that multiple individuals (of both sexes) must have arrived from another port in one or more invasions. As indicated for M. inexspectata, the B. virginica population will also probably seed new populations in San Francisco Bay and elsewhere. Based on its cnidome as well as the morphology of both medusa and polyp, M. inexspectata has been reclassified by moving it from the family Olindiidae, Limnomedusae, to the family Moerisiidae, Anthomedusae.     

Changes in morphology and physiology of an East Mediterranean sponge in different habitats

The sponge Tetilla sp. (Tetractinomorpha: Tetillidae) is a common species in the eastern Mediterranean. This sponge inhabits four different habitat types differing in wave impact and irradiance levels. Two of these habitats (a shallow cave and deep water) are characterized by relatively calm water, whereas the other two (shallow exposed site and tide pools) are in turbulent water with high energy flow. The present study examined the influence of physical (depth, illumination and water motion) and biotic factors on morphology, skeletal plasticity and reproductive traits among the four spatially separated populations. Sponges from tidal pools had significantly larger body volume than sponges from deep water and from shallow caves (ANOVA: tidal-deep P<0.0001; tidal-shallow caves P<0.05). Sponges from exposed habitats were significantly larger than deep-water sponges (ANOVA: P=0.01). In addition, individuals from tide pools and from the exposed habitat had a significantly higher proportion of structural silica than sponges from the calmer deep water and from the cave sites. Oxea spicules in sponges from the calm habitats were significantly shorter than in those from the tidal pools and the exposed habitats. The percentage of spicules out of a sponges dry weight in individuals transplanted from deep (calm) to shallow (turbulent) water significantly increased by 21.9±12.9%. The new spicule percentage did not differ significantly from that of sponges originally from shallow water. Oocyte diameter differed significantly between habitats. The maximal size of mature eggs was found in deep-water sponges in June (97±5 m). In the shallow habitats, a smaller maximal oocyte diameter was found in the cave, in May (56.5±3 m). Furthermore, oocyte density in shallow-water sponges was highest in May and decreased in June (with 88.2±9 and 19.3±9 oocytes mm^–2, respectively). At the same time (June), oocyte density of deep-water sponges had just reached its maximum (155±33.7 oocytes mm^–2). The difference in oocyte size and density between deep- and shallow-water individuals indicates an earlier gamete release in the shallow sponge population. The results suggest that plasticity in skeletal design of this sponge indicates a trade off between spicule production and investment in reproduction.Communicated by O. Kinne, Oldendorf/Luhe     

Assemblage shift following population collapse of a non-indigenous bivalve in an urban lagoon

The quahog Mercenaria mercenaria has been introduced repeatedly to the Pacific coast of North America, but only one population is known to have become established. In the 1970s, the population of M. mercenaria at Colorado Lagoon, in Los Angeles County, California (33^o46′16″N, 118^o08′05″W), was estimated at more than 300,000 individuals. To determine the current status of this non-indigenous species (NIS), in 2009, we sampled 57 intertidal and 20 shallow subtidal plots, identifying and quantifying collected bivalves. No quahogs were found among the 2,490 living bivalves in our plots, though two were found intertidally outside of our plots. The M. mercenaria population has thus collapsed since 1980, but the native community has not recovered. Six of the fourteen living bivalve species we encountered were NIS; three are new records for the location, including the clam Venerupis philippinarum, which made up 87.6% of collected individuals. Though M. mercenaria is likely on its way to extinction on the US Pacific coast, the bivalve assemblage at this location remains heavily dominated by NIS.     

Mechanical clam dredging in Venice lagoon: ecosystem effects evaluated with a trophic mass-balance model

Harvesting of the invasive Manila clam, Tapes philippinarum, is the main exploitative activity in the Venice lagoon, but the mechanical dredges used in this free-access regime produce a considerable disturbance of the lagoon ecosystem. An ecosystem approach to study the complex effects of clam harvesting was implemented using a trophic mass-balance model. The trophic relations in the ecosystem were quantified with a mixed trophic impact analysis and further evaluated by considering different explanations for the " Tapes paradox", which consists of the apparent population enhancement of Manila clams by dredging and the apparent nutritional advantages that this species receives from re-suspended organic matter. The key-role played by this introduced species is highlighted by a network analysis that indicates a "wasp-waist control" of the system by Manila clams. The model constructed to characterise the present state of the Venice lagoon ecosystem is compared with models produced for a reconstructed past lagoon and a projected future lagoon. The future model was obtained by simulating the elimination of clam dredging in 10 years. The three different models were compared using thermodynamic and informational indices. Simulating the elimination of clam dredging produced a 33% increase in artisanal fishery catches, carried out by means of static gears, even with no change in fishing effort. These simulations also forecast an increase in the mean trophic level of the artisanal fishery catches as a positive effect of eliminating mechanical clam harvesting.     

Sea otter foraging on deep-burrowing bivalves in a California coastal lagoon

Sea otter, Enhydra lutris, predation had no detectable effect on abundance and size distribution of deep-burrowing bivalve prey in the Elkhorn Slough, California, USA. Up to 23 otters were present for 6 mo of the study period (March 1984 through April 1985). This is in contrast to previous studies of sea otter predation, especially on the shallow-burrowing Pismo clam Tivela stultorum, which can be found along the wave-exposed coast near the slough. The deep-burrowing clams Tresus nuttallii and Saxidomus nuttalli made up 61% of the prey taken in the slough, and are more difficult for otters to excavate than Pismo clams. The occurrence of foraging otters was highest in an area where the two bivalve prey were extremely abundant (18 individuals m^–2). However, the otters did not selectively prey on the largest clams available within the study sight, but foraged preferentially in a patch of smaller individuals where bivalve burrow depth was restricted by the presence of a dense clay layer. This foraging strategy maximized the amount of prey biomass obtained per unit volume of sediment excavated. Our findings suggest that in soft-sediment habitats deep-burrowing bivalves may be more resistant to otter predation than shallower burrowers.     

Chromosomal mapping of major ribosomal rRNA genes in the hard clam (<Emphasis Type="Italic">Mercenaria mercenaria</Emphasis>) using fluorescence in situ hybridization

Karyotype and chromosomal location of the major ribosomal RNA genes were studied in the hard clam (Mercenaria mercenaria Linnaeus) using fluorescence in situ hybridization (FISH). Metaphase chromosomes were obtained from early embryos. Internal transcribed spacers (ITS) between major RNA genes were amplified and used as FISH probes. The probes were labeled with digoxigenin-11-dUTP by polymerase chain reaction and detected with fluorescein-labeled anti-digoxigenin antibodies. FISH with the ITS probes produced two to four signals per nucleus or metaphase. M. mercenaria had a haploid number of 19 chromosomes with a karyotype of seven metacentric, four metacentric or submetacentric, seven submetacentric, and one submetacentric or subtelocentric chromosomes (7M + 4M/SM + 7SM + 1SM/ST). Two ITS loci were observed: one located near the centromere on the long arm of Chromosome 10 and the other at the telomere of the short arm of Chromosome 12. FISH signals on Chromosome 10 are strong and consistent, while signals on Chromosome 12 are variable. This study provides the first karyotype and chromosomal assignment of the major RNA genes in M. mercenaria. Similar studies in a wide range of species are needed to understand the role of chromosomal changes in bivalve evolution.     

Measurement of calcium carbonate deposition in molluscs by controlled etching of radioactively labeled shells

Rates of calcium carbonate removal from shell pieces of Argopecten irradians (Say) and Mercenaria mercenaria (L.) were dependent on the type of etching fluid used and not shell origin. Etching was uniform over the entire shell surface, but surface morphology differed with etching fluids. Peak radioactivity was in early eluant fractions of shells etched immediately after radioactive labeling, and in later fractions when individuals were placed in isotope-free seawater after labeling. The etching technique can measure growth during the labeling period and subsequently by estimating the amount of calcium in fractions prior to the radioactive peak. Geometry of shell layers influenced the pattern of radioactivity seen in fractions. Peak location varied inside and outside the pallial line of individual M. mercenaria. A significant portion of the inorganic carbon used in shell formation by M. mercenaria was derived from metabolic CO₂.