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     

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     

Digestion in a subtidal population of Mercenaria mercenaria (Bivalvia)

The digestive processes of Mercenaria mercenaria (Linn.) were studied in a natural subtidal population. Hourly sampling of 4 to 5 quahogs was conducted over one 25 h period in August, 1978, at Woods Hole, Ma., USA. Crystalline style length did not vary significantly with time, tide, particulate C, N or suspended material in the water. Changes in secretory activity of the style sac epithelium were not evident from histological sections. Based on cellular morphological differences, digestive tubules were individually classified into one of 4 categories, indicative of the state of intracellular digestion within the digestive cells. All 4 tubule types were present in individuals from each sample hour, signifying that intracellular digestion occurs continually within the digestive gland. A 3 h time lag is evident between times of peak C, N or suspended materials and apparent peak levels of absorptive (Type II) tubules.Contribution No. 76 Marine Science Institute, Northeastern University; Nahant, MA 01908, USA     

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.     

Heterozygosity,growth, and survival of the hard clam,Mercenaria mercenaria,in seagrass vs sandflat habitats

Populations of the hard clam,Mercenaria mercenaria L., sampled from dissimilar, adjacent habitats (October and November 1987), were used to assess environmental effects on associations among multi-locus heterozygosity, growth, and survival. Individuals were collected from three widely separated localities along the east coast of North America with each consisting of a seagrass bed and an adjacent, unvegetated sandflat. Demographic differences between adjacent populations were attributed to habitat type. Samples from intertidal sandflats at two of the localities were dominated by younger individuals than those from seagrass beds. Differential growth between adjacent populations was detected at only one locality wherein seagrass individuals grew faster than those from the sandflat. Allelic frequencies revealed adjacent populations that were genetically homogeneous. Nearly all of the genetic variance (98%) was within populations, yet a small (1.7%), but significant, portion occurred between localities. Observed and expected heterozygosities revealed a deficiency of heterozygotes in all six samples. Inbreeding and small-scale population subdivision were discounted as causative factors because deficiencies were heterogeneous across loci. Multi-locus heterozygosity was not correlated with growth rate in samples from any locality. Using a consensus test, multilocus heterozygosity was positively associated with ageclass in sandflat, but not seagrass, samples. We suggest that heterozygosity-fitness trait associations in marine bivalves are more likely to occur in populations inhabiting more stressful, fluctuating environments     

Behavior and growth ofMercenaria mercenaria during simulated storm events

Experiments were conducted in April–August 1989 on juvenileMercenaria mercenaria (L.) in an oscillatory water tunnel to simulate resuspension of bottom sediments by waves and to determine the effects of shortterm storm events on particle ingestion, pseudofeces production, and shell growth. Juveniles (mean length = 19.2 mm) were subjected to identical concentrations of algae in both low-flow, gentle waves (maximum velocity = 7 cm s^–1) and high-velocity storm waves (maximum velocity = 22 cm s^–1). Suspended sediment levels reached 193 mg 1^–1 at 1 cm above the bed during storms. Shell growth decreased by a maximum of 38% during the storm when levels of phytoplankton were high (average cell concentration = 43 × 10⁶ cells 1^–1), and by 18% when phytoplankton levels were low (av cell conc = 6 × 10⁶ cells 1^–1). Orientation of clam siphons was not related to flow direction. Significantly more pseudofeces were produced when the clams were subjected to increased sediment resuspension under waves, and in troughs of sand ripples. The size of sediment grains ingested did not vary significantly among the flow treatments. The decrease in shell growth during storms may be due to a reduction in filtration rate coupled with a decrease in net energy gained from filtration due to costs of pseudofeces production. The magnitude of the decrease seems to be related to concentration of algae, water temperature, age of clams and sediment transport mode (bed load or suspended load). Thus, the interpretation of growth increments must be made in the context of these environmental variables.     

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.     

Growth and survivorship in a tidal-flat population of the bivalve Mercenaria mercenaria from Southampton water

A monthly sampling programme was conducted to investigate aspects of the biology of a population of Mercenaria mercenaria (L.) Smooth growth curves were constructed for each year class, and the factors affecting growth rate were considered. Seasonal cycles in flesh weight and calorific content appear to be related to gonal proliferation and subsequent spawning. Recruitment is sporadic and probably depends on the spawning of upstream populations, where conditions are more favourable for larval development. Survivorship curves were constructed for each year class; predation by crabs and gulls appears to account for a large proportion of the observed mortality.     

Transport of cadmium and other metals in the blood of the bivalve mollusc Mercenaria mercenaria

The distribution of metals (Ba, Cr, Cu, Fe, Mn, Ni and Zn) in the blood plasma and the circulating hemocytes was determined for a eulamellibranch bivalve, the quahog Mercenaria mercenaria (L.), collected from a relatively clean site at Falmouth, Massachusetts, USA. Whole blood volume was an exponential function of quahog length (Y=5.71×10^-5 X^3.0678; Y=ml of whole blood; X=mm length). Of this volume, 1.2±0.3% ( ) was attributable to blood cells. Total metal content (g metal, or g metal normalized per g of whole blood) was much higher in blood plasma than in hemocytes. In quahogs exposed in the laboratory to 100 ppb ¹⁰⁹Cd, 93.0% of the total accumulated blood Cd was in the plasma rather than in the circulating hemocytes (7.0%), irrespective of the length of exposure (1 h to 31 d). Less than 5% of the plasma Cd was either Cd²⁺, small inorganic Cd complexes or bound to organic molecules with a molecular weight smaller than 1 000. Cadmium was primarily bound to high molecular weight protein(s) (>60 000 daltons) within the plasma. This plasma protein-Cd complex has a low affinity constant (approximately 10⁴ M ^-1), indicating non-specific Cd binding, although the capacity for Cd-binding in the plasma is great (as high as 200 g Cd per ml of plasma). Blood plasma may be far more important in metal transport than has previously been realized.     

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     

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     

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.     

Rate of excretion of ammonia by the hard clam Mercenaria mercenaria and the American oyster Crassostrea virginica

Two species of bivalve molluscs, Mercenaria mercenaria Linné and Crassostrea virginica Gmelin, were maintained in an identical laboratory environment and fed the same diet for 1 month prior to measuring the quantity of ammonia which they excreted per gram dry weight. Data derived from juveniles and adults of these species were fitted to a log-log equation. M. mercenaria excreted more ammonia per gram body weight than C. virginica, and considerably more scatter is evident in the fit of the data from M. mercenaria to the log-log relationship. Experiments showed that neither temperature fluctuation during the experiments nor decomposition of organic nitrogen in the test water account for the scatter of points. Behavioral differences between the two species of organisms may explain these differences.This work is the result of research sponsored by NOAA Office of Sea Grant, Department of Commerce, under Grant No. 04-3-158-30.     

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     

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.     

Genotype-specific growth of hard clams (genus Mercenaria) in a hybrid zone: variation among habitats

Shell growth rate is an important component of fitness in bivalve molluscs. Using the parameter computed from the von Bertalanffy growth equation, we quantitatively compared rates of annual shell grwoth among the hard clams Mercenaria mercenaria, M. campechiensis, and their hybrids sampled from a variety of habitats in the Indian River lagoon, Florida, USA, a zone of species overlap and natural hybridization. Our results indicate that the classical paradigm describing hard clam growth, in which growth rate is fastest in M. campechiensis, intermediate in hybrids, and slowest in M. mercenaria is not supported in the Indian River lagoon. Instead, M. campechiensis has a growth advantage in deep-water habitats in the northern section of our study area. In the central and southern sections of our study area, hybrids have a growth advantage over M. mercenaria in shallow-water habitats, but M. mercenaria has a growth advantage over hybrids in deep-water habitats. In all other sampled habitats, either growth rate among genotype classes is equal, or M. mercenaria has a growth advantage. This complex relationship between genotype and habitat-specific growth provides a mechanism for selection to act on hard clams in the Indian River.     

Nucleic acids and growth of larval and early juvenile spider crab,Hyas araneus

The spider crabHyas araneus (L.) was collected from the North Sea in winter 1986–1987 and reared in the laboratory from hatching of the Zoea I (ZI) through the first juvenile instar (CI). Within a given moult cycle, individuals of the same age were sampled in intervals of 2 (ZI, ZII, CI) or 3 d (megalopa) for analysis of dry weight (W), carbon (C), nitrogen (N), hydrogen (H), protein, DNA, and RNA. Lipid was calculated from C. Biomass, growth rate and nucleic acid contents showed high variability during each moult cycle and between instars. Instantaneous growth rates of C were high in postmoult and intermoult, and low in the premoult period of each moult cycle. A shift was observed from high rates of lipid accumulation in the postmoult and intermoult stages to proportionally increasing protein accumulation during late premoult (ZI), or throughout a major part of the remaining moult cycle (in all other instars). DNA was accumulated throughout the ZI and ZII instars, but decreased in late premoult megalopa. It increased again from late intermoult through intermediate premoult in juveniles. RNA increased continuously during ZI and ZII, and decreased in the megalopa, almost to levels that had been found immediately after hatching. In juveniles, variation in RNA followed closely those in DNA. Cell multiplication (expressed by DNA increase) dominated over increase in cell size (defined by the C/DNA ratio) during the zoeal instars and in postmoult through early intermoult in the megalopa and CI. When specific (C-related) RNA values and RNA/DNA ratios were compared with instantaneous growth rates in C and N, no general correspondence was detected. The only significant relationship between specific RNA values and instantaneous C or N growth rates was found in the megalopa. The same held for the relationship between the RNA/DNA ratio and growth. Here, in addition to the megalopa, a correspondence with C growth was also found in the CI instar. Our results suggest that variation in nucleic acids may provide useful insights into mechanisms of growth on the cellular level (cell multiplication vs cell enlargement). However, lack of general correlation with variation in growth rates ofH. araneus larvae shows that the use of nucleic acids as a measure of growth is probably based upon too simplistic assumptions; it may not yield reliable predictions, when growth is associated with developmental events.     

Differential timing of larval starvation effects on filtration rate and growth in juvenile Crepidula onyx

This study demonstrates that the timing of larval starvation did not only determine the larval quality (shell length, lipid content, and RNA:DNA ratio) and the juvenile performance (growth and filtration rates), but also determine how the latent effects of larval starvation were mediated in Crepidula onyx. The juveniles developed from larvae that had experienced starvation in the first two days of larval life had reduced growth and lower filtration rates than those developed from larvae that had not been starved. Lower filtration rates explained the observed latent effects of early larval starvation on reduced juvenile growth. Starvation late in larval life caused a reduction in shell length, lipid content, and RNA:DNA ratio of larvae at metamorphosis; juveniles developed from these larvae performed poorly in terms of growth in shell length and total organic carbon content because of “depletion of energy reserves” at metamorphosis. Results of this study indicate that even exposure to the same kind of larval stress (starvation) for the same period of time (2 days) can cause different juvenile responses through different mechanisms if larvae are exposed to the stress at different stages of the larval life.