Analysis of growth rate in Mya arenaria using the Von Bertalanffy equation

Field studies were conducted in Gloucester, Massachusetts, USA, to determine linear shell growth rates for Mya arenaria. These rates were then compared with those reported for the same species from other locations. Most shell deposition occurred from March through November of each year. Winter interruptions in growth were not as marked in the small clams as in the larger ones (>60.0 mm). Annual variations in growth were slight during the period 1973–1974. Growth of mature clams (>35.0 mm) slowed during the spawning season. No significant sexual dimorphism in mean annual growth rates was detected. Winter rings were shown to be a reliable method for determining age in clams from Gloucester. Age-size relationships, based on two independent measures of annual growth, winter rings and tagging experiments, were computed using the Von Bertalanffy growth equation. No well-defined latitudinal patcerns in growth could be established for M. arenaria.     

Diet and food preferences of the adult horseshoe crab Limulus polyphemus in Delaware Bay,New Jersey,USA

Adult horseshoe crabs Limulus polyphemus (L.) feed on a wide variety of infaunal and epifaunal invertebrates during their spring spawning migration in Delaware Bay, New Jersey, USA. Comparison of the gut contents with estimates of available prey showed that the most abundant potential prey item, the bivalve Gemma gemma, was avoided. The thinner shelled but comparatively scarce clam Mulinia lateralis was a preferred prey item. In the laboratory, crabs fed on G. gemma when it was the only available item but not when M. lateralis or soft-shell clams, Mya arenaria, were offered in conjunction. Large M. lateralis (>10mm) were preferred to small M. lateralis; there was no discrimination between M. lateralis and M. arenaria of the same size. Male and female horseshoe crabs had similar gut contents and laboratory feeding preferences, despite the fact that females are larger than males. Crabs spawning later in the summer contained more food than did crabs collected at the peak of spawning activity.     

Induced triploidy in the soft-shelled clam Mya arenaria: energetic implications

Given that triploid adult bivalves reportedly grow larger and faster than their diploid siblings, such differences should be traceable to variation in energy allocation. In one proposed mechanism, retarded gametogenesis found in triploid adults may allow them more energy for somatic growth. Another hypothesis states that triploids are more heterozygous; increased heterozygosity has been positively correlated with enhanced growth. Juvenile soft-shelled clams, Mya arenaria, were treated with cytochalasin B to induce triploidy and examined with respect to components of a balanced energy equation (C=P+R+E). The variables measured were oxygen uptake (V _{o 2}), filtration rate (FR), dry tissue weight, shell length, shell height and shell inflation. Energy budgets were constructed and diploid and triploid groups compared. Few significant differences were found between diploid and triploid juvenile clams with respect to energy budget components. However, at seven loci assayed electrophoretically the triploid individuals were nearly twice as heterozygous as their diploid siblings. Moreover, tripoloid variances were less than diploid variances for every variable measured. Increased heterozygosity has been correlated with the decreased variance of morphological parameters. This study is believed to be the first to show decreased variance of physiological properties as well as morphological characters. Overall the data clearly indicate that energy allocation in juvenile M. arenaria is not related to ploidy.     

Disease progression and mortality in neoplasticMya arenaria in the field

Four 6-mo mark-and-recapture experiments conducted in Long Island Sound, USA, from 1988 to 1990, involving approximately 2250 individual observations, demonstrated that under natural conditions significantly higher mortality (p0.001, chi-square test) occurred amongMya arenaria (L.) with hematopoietic neoplasia than those diagnosed as non-neoplastic. Using a blood-screening technique, the clams were diagnosed and placed in one of three diagnostic groups based on the severity of the disease (the percentage neoplastic cells per total number of blood cells): non-neoplastic (NN), 0%; low-severity neoplastic (LSN), < 50%; high-severity neoplastic (HSN), > 50%. Mortality of those clams initially diagnosed as HSN ranged from 48% to 78%, depending on the test period, as compared to 3% to 21% for the non-neoplastic. Mortality in the LSN treatment varied from 8% to 34%. Both progression and remission were evident in clams at all stages of the disease. Mortality and rates of progression and remission in individuals appeared to be linked to water temperatures. Differential mortality may be responsible for the apparent seasonal cycle of prevalence in populations.     

Population dynamics of the soft-shell clam Mya arenaria

A life table was constructed for Mya arenaria from Gloucester, Massachusetts, USA, based on schedules of age-specific fecundity and mortality determined under natural conditions. Mortality rates decrease with size and age in this species, with the period of maximum mortality occurring during the summer months. Mortality rates during the fall and winter were considerably lower, perhaps due to the inactivity of natural predators. The survivorship curve for M. arenaria approximates the Type 3 curve of Deevey (1947). Mean life expectancy is low in recently-settled clams, peaks when the individual reaches 30.0 to 34.9 mm (1 year of age) and remains fairly high for most of the remainder of life. The intrinsic rate of natural increase (r _max) is very high: 4.74. This enormous rate of potential increase is offset by high rates of larval mortality in the plankton. Unlike the reproductive values of most animals studied, those in M. arenaria peak late in life, well after the know age of first reproduction. This is probably the result of increased fecundity with age. The implications of this work in the area of resource management are discussed.     

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     

Tissue distribution and excretion of 99mTc-disofenin in three marine species: Pleuronectes americanus (winter flounder), Homarus americanus (lobster), and Mya arenaria (soft-shell clam)

To determine the pharmacokinetics of a small lipophilic molecule in vivo, the distribution and accumulation of ^99mTc-radiolabelled disofenin (diisopropylacetanilide iminodiacetic acid) were traced during 1991–1992 by scintigraphy and gamma well counting in winter flounder (Pleuronectes americanus collected from Boston Harbor and Long Island Sound in 1992), lobsters (Homarus americanus collected from Massachusetts Bay in 1991), and soft-shell clams (Mya arenaria purchased in 1991). The agent was distributed throughout the bodies of lobsters within 12 s, throughout flounder within 40 s, and throughout clams within 2 min. It was concentrated most strongly by the liver of flounder, which contained 61.2±7.8% of the injected dose within 1 h of injection, and by the lobster hepatopancreas. Accumulation also occurred in the flounder kidney, lobster antennal glands, and the kidney and pericardial glands of clams. The compound was rapidly excreted from the flounder liver into the gall bladder, and from the lobster hepatopancreas into the stomach. The data suggest its excretion from the lobster antennal glands and clam kidneys. The rate of clearance of disofenin from the body varied among species: 99±2.1% of the initial dose remained in flounder sampled 16 to 24 h after injection, compared to 80.5±7% remaining in the lobster after 15 h, and 87.4±5.9% remaining in clams after 27 h. The clearance rates in flounder and lobster are considered to be minimum values because of the lack of gut activity in unfed individuals. Overall, these in vivo tracer studies establish the utility of scintigraphy for assessing the uptake and excretion of a lipid soluble compound in different taxa, and may be applicable for evaluating disease and/or environmental effects on organ function in marine animals.     

Gill pump characteristics of the soft clamMya arenaria

Pumping rates in the soft clamMya arenaria, collected in June 1987 in the Great Belt, Denmark, were determined as rates at which clams cleared suspensions of algae,Dunaliella marina. The frictional resistance of the siphons to water flow was estimated by studying the effect of excision of the siphons at the base on the rate of water pumping. The frictional resistance was also calculated from the Poiseuille equation. Excision of the siphons had no measurable effect on the pumping rate, and calculations indicated pressure losses ranging between 0.3 to 1.2 mm H₂O. This is consistent with the conception that the capacity of filter feeding bivalves to process large amounts of water depends upon a low resistance to water flow in the siphons.     

Toxicity evaluation of a complex metal mixture to the softshell clam Mya arenaria

Adults of the softshell clam Mya arenaria were continuously subjected to a flowing raw seawater solution containing a mixture of salts of manganese, zinc, lead, nickel, copper, and cadmium. Final calculated concentrations, in g l^-1, of the toxicant solution were 7200 Mn, 2500 Zn, 70 Pb, 50 Ni, 50 Cu and 1 Cd; these concentrations approximated highest measured levels within surficial interstitial sediment waters from mid-Narragansett Bay, Rhode Island. M. arenaria were also subjected to a 20% solution, i.e., 1440 Mn, 500 Zn, 14 Pb, 10 Ni, 10 Cu, and 0.2 g l^-1 Cd. One study was conducted for 112 days in winter at 0° to 10°C and another for 16 days in summer at 16° to 22°C. In the winter study, all clams exposed to a 100% solution died between the 4th and 10th week; soft parts of survivors at 6 weeks contained about 19 times more Pb, 15 x more Zn, 12 x more Cu, 10 x more Mn, 3 x more Ni and 0.1 x more Cd than controls; relatively minor changes in whole body elemental content of Ca, Cr, Fe, K, Mg, Na, Sr, and V were observed. Clams exposed to a 20% solution during winter survived the 112 day study; at that time these contained about 5 x more Cu, 4 x more Mn, 3 x more Zn and about 2 x more Pb than controls; comparatively minor changes were observed in other elements examined. In the summer study, all M. arenaria subjected to the 100% solution died between 6 and 14 days; survivors from this group at 7 days contained about 25 x more Pb, 13 x more Cu, 11 x more Zn, 7 x more Mn, and 3 x more Ni than controls; other changes in elemental content were not as pronounced. Mortality in the 20% group during summer was slightly higher than controls during the 16 day study; at 14 days survivors from this group contained about 12 x more Mn, 7 x more Pb, 7 x more Zn, 4 x more Cu, and 3 x more Ni than controls. Survival and bioaccumulation patterns were not altered through feeding a supplemental diet of algae. The significance of these findings are discussed in terms of potential environmental perturbations, especially local dredging practices.     

PCR-based detection of mtDNA haplotypes of native and invading mussels on the northeastern Pacific coast: latitudinal pattern of invasion

Frequencies of mitochondrial haplotypes characteristic of native Mytilus trossulus and introduced M. galloprovincialis were determined in populations along the west coast of North America from San Diego, California, to the Aleutian Islands, Alaska. We also identified the haplotypes of mussels cultured from larvae arriving in Coos Bay, Oregon, during 1988–1990 from sites in Japan in the seawater ballast of ocean-going ships. All mussels from ballast-water samples were M. galloprovincialis. We found that sampled populations north of San Francisco Bay, including Coos Bay, were entirely composed of mussels with the M. trossulus haplotype, while haplotypes of both species were present in all sites in and south of San Francisco Bay. The presence of M. trossulus in southern sites is contrary to evidence from allozyme studies, and we suggest that mtDNA introgression from M. trossulus to M. galloprovincialis may explain this discordance. This study demonstrates that, despite continued transport and release, M. galloprovincialis has not become established in northern sites. Failure to invade the north coast of North America may reflect environmental unsuitability for M. galloprovincialis. However, invasion success may be probabilistic, and the continuing release of M. galloprovincialis larvae may foreshadow a future successful invasion.     

Siphon nipping facilitates lethal predation in the clam Mesodesma mactroides (Reeve, 1854) (Mollusca: Bivalva)

In soft sediment marine communities, fishes frequently bite off extended siphons of buried clams; the consequential shortening of the siphon is known to reduce burial depth of the clams, secondarily increasing their vulnerability to lethal excavating predators. In this study, siphon nipping on the yellow clam, Mesodesma mactroides, was simulated by removing the top 6.6–30% of siphons. This caused a burrow reduction in 25–75%, respectively, compared to control individuals with intact siphons, in field and laboratory trials. To examine subsequent consequences of reduced burial depth, we exposed nipped and intact clams to potential predators in the laboratory simulating the observed natural clam abundance. Artificially nipped clams were consumed twice as much as control clams. The present results suggest that sympatric croppers contribute to the stock recovery failure by facilitation of lethal predation and that re-seeding to increase the local abundance of M. mactroides should be an essential aspect of conservation efforts in South America.     

Stable isotopes changes in the adductor muscle of diseased bivalve Ruditapes philippinarum

In this article, we show how a disease could bias stable isotope analyzes of trophic networks and propose a strategy in the choice of tissues to be analyzed. In the past few years, a new pathology (brown muscle disease or BMD) affecting the posterior adductor muscle of Ruditapes philippinarum has emerged in Arcachon Bay. BMD induces a necrosis of muscle tissues which become infused by conchiolin and hence calcified. As muscle of mollusks are often used for trophic food webs studies through stable isotopic analyzes, this work investigated the effect of BMD on carbon and nitrogen isotopic ratios of anterior and posterior adductor muscles of clams collected in February and August 2007. Infected clams displayed a lower condition index and a posterior adductor muscle δ¹³C enrichment of 1.2‰ in February and 0.7‰ in August. δ¹⁵N of posterior muscles was however not affected by the disease. Anterior muscle of diseased clams remained healthy and displayed the same isotopic signature as both posterior and anterior muscular tissues of healthy clam. Acidification significantly depleted δ¹³C in posterior muscles of infected clams, suggesting calcification, contrary to anterior muscles of infected clam and to both muscles of healthy clams, where no effect was observed. An X-ray diffractometry analysis confirmed the presence of CaCO₃ (aragonite). Trophic food web studies relying on stable isotope ratios should utilize only healthy animals or anterior adductor muscles when expertise in mollusk pathology is lacking.     

Potential effects of a non-indigenous predator in its expanded range: assessing green crab, <Emphasis Type="Italic">Carcinus maenas</Emphasis>, prey preference in a productive coastal area of Atlantic Canada

The non-indigenous green crab (Carcinus maenas) is an important predator on bivalve wild beds in coastal areas worldwide. This study explored size-dependent green crab prey preference on American oysters (Crassostrea virginica), blue mussels (Mytilus edulis), and soft-shell clams (Mya arenaria) in a productive coastal system of Atlantic Canada. Using two sizes of prey and three different experimental manipulations, small, medium, and large green crabs were given a choice among these three bivalves, and their daily feeding rates were monitored over the course of 3 days. For both prey sizes, green crabs showed an early feeding preference for soft-shell clams and, only as they declined in numbers, a switch toward mussels and subsequently toward oysters. We found that such changes in the timing (order) of prey preference are related to prey differences in shell thickness, a fairly reliable indicator of prey shell strength.     

Population differences in nerve resistance to paralytic shellfish toxins in softshell clam, <Emphasis Type="Italic">Mya arenaria</Emphasis>, associated with sodium channel mutations

The softshell clam, Mya arenaria, is a commercially important bivalve with wide latitudinal distribution in North America. Populations of clams with a history of repeated exposure to toxic Alexandrium spp. have developed a natural resistance to the paralytic shellfish toxins (PSTs) produced by these algae. An association between PST resistance in individual clams and a single mutation in the saxitoxin (STX) binding region of the α-subunit of the voltage-gated sodium (Na⁺) channel gene was previously identified. Here we establish that more than one mutation associated with nerve resistance to STX occurred at this locus. Both cDNA from mRNA and genomic DNA sequences from individual clams are identical demonstrating that both alleles are expressed simultaneously. In addition, one resistant allele per individual is sufficient to confer neural resistance to STX even though heterozygous individuals show an intermediate level of resistance to STX in in vitro nerve trunk assays.     

Oxygen transport by the hemocyanin of the protobranch mollusc Solemya reidi

Oxygen transport by the hemocyanin of the protobranch mollusc Solemya reidi Bernard was studied in native hemolymph samples. Clams were collected from two different reducing environments, beneath log booms in Alberni Inlet, British Columbia, Canada, and from the sewage effulent in Santa Monica Bay, California, USA. The hemocyanin concentration in a pooled hemolymph sample (n = 10 individuals) was 33.5 mg ml⁻¹. The mean hemolymph pH of five Alberni clams maintained for 3 wk in mud was 7.96 ± 0.06. No significant variation in hemocyanin oxygen-affinity or cooperativity was found for hemocyanin in whole hemolymph samples from these five individually studied clams. There was a significant difference only at 15 °C in the oxygen affinity of hemocyanin in pooled whole-hemolymph samples from S. reidi collected from Alberni Inlet compared with clams collected from Santa Monica Bay. Little effect of temperature on hemocyanin oxygen-affinity was found for temperatures below 20 °C; above 20 °C, the oxygen affinity was reduced for clams from both sites. Temperature and pH had no apparent affect on hemocyanin cooperativity. Moderate Bohr shifts were found at all temperatures examined. The presence of physiologically relevant concentrations of thiosulphate in hemocyanin samples resulted in a decrease in hemocyanin oxygen-affinity, opposite to the effect on hemocyanin found for the hydrothermal vent crab Byth-ograea thermydron, but thiosulphate had no effect on hemocyanin cooperativity. Received: 2 September 1997 / Accepted: 6 February 1998     

Effects of Lunatia heros predation on the population dynamics of Mya arenaria and Macoma balthica in Maine,USA

The interaction between the naticid snail predator Lunatia heros and 2 iteroparous, infaunal, intertidal bivalves was investigated in Lubec, Maine, USA. The Mya arenaria population consists primarily of young, small individuals. M. arenaria survivorship is low when young (3.5% yr^-1 for the first 5 yr), then increases. M. arenaria can attain a length of 60 mm, but it is susceptible to L. heros attack only until it is 30 mm long. It delays reproduction until it is 20 mm long (3.8 yr) and diverts its resources instead into rapid early growth (4.9 mm yr^-1 for the first 5 yr). The Macoma balthica population has a larger proportion of older individuals than does that of M. arenaria. Survivorship is higher for M. balthica than for M. arenaria (76.3% yr^-1 for the first 5 yr). Unlike M. arenaria, M. balthica attains a final length of only 25 mm and all sizes are susceptible to L. heros attack. M. balthica grows slowly (2.7 mm yr^-1 for the first 5 yr) and diverts its resources into earlier reproduction at a length of 10 mm (2.9 yr). These contrasting life-history patterns and the possible relationship between bivalve resource allocation and refuges from predation are discussed.     

Comparative life histories of three species of planktonic calanoid copepods in Dabob Bay,Washington

The life history patterns of Calanus marshallae Frost, C. pacificus Brodsky, and Metridia lucens Boeck were determined in Dabob Bay, Washington, USA, during 1973 and October 1981–September 1982. C. marshallae emerged from diapause and moulted to adults in January–February. One major generation was observed, having been produced mainly during early to mid-March. Most surviving individuals spawned in March, arrested development at the C5 stage, and were in diapause by late May. However, some individuals developed to adults by late April and produced a very minor second generation. C. pacificus emerged from diapause and moulted to adults in February–March. The first generation produced during the spring lagged slightly behind the C. marshallae generation. C. pacificus produced two additional generations, one in late spring and one in the fall. The production of the three generations coincided with or closely followed phytoplankton blooms. Some C. pacificus arrested development at the C5 stage and entered diapause during the summer, but the majority of the population did not do so until fall. M. lucens did not appear to enter a diapause state. The fall and winter population was chiefly adult females which mostly remained at depth and were reproductively immature. Nevertheless, naupliar M. lucens were found on all sampling dates, indicating that reproduction never ceased as it did for the two Calanus species. M. lucens produced generations in late winter/early spring, late spring, and late summer/early fall. Interspecific comparisons between the life histories observed in Dabob Bay and intraspecific comparisons between the life histories in Dabob Bay and other locations are made. The advantages or disadvantages conferred to the copepods by their differing life histories are also discussed.     

Growth,reproduction and gross biochemical composition of the Manila clam Ruditapes philippinarum in the Bay of Arcachon,France

Growth, reproduction and gross biochemical composition of the Manila clam Ruditapes philippinarum were studied for one oceanic and two inner stations in the Bay of Arcachon, France, from March 1989 to March 1991. During this period, sea-water temperature, salinity and chlorophyll a were also recorded. A marked increase in length occurred during the first year in all areas, after which growth rates decreased. In contrast, weight increased more steadily. The Manila clam exhibited best development in the oceanic area, but there was no difference in growth of clams between the two inner stations. Differences in growth between oceanic and inner stations may result from differences in fluctuations of environmental conditions such as temperature and salinity. Except for higher carbohydrate contents in clams recovered in autumn from the oceanic station Le Ferret, biochemical components differed little between stations. During the second winter, glycogen levels were relatively low, but no mortalities were recorded. On the other hand, sowing spat in autumn instead of spring or sowing larger-sized spat did not reduce the time required for culture of R. philippinarum.     

Possibilities for management of coastal foredunes with deteriorated stands ofAmmophila arenaria (marram grass)

Ammophila arenaria (marram grass) is the most important plant species for sand stabilization in European coastal foredunes. Stand degeneration due to poor supply of wind-blown sand enhances the susceptibility for wind erosion when successional species do not become established. ReplantedA. arenaria often fails to become established. In the present study we examined whether management practices such as mowing, fertilizing, burning or below-ground cutting of plants may be applied to re-establish the vigour ofA. arenaria. Field experiments performed at exposed sites, where naturally succeeding plant species are not supposed to become established due to salt spray, showed that none of the applied methods resulted into enhanced tiller density ofA. arenaria. Thus, further studies are necessary to solve this type of management problem. At the leeward side of foredunes, the successional speciesFestuca rubra ssp.arenaria andElymus athericus could be established successfully as both pre-grown seedlings and planted bundles of cuttings (all further experiments were fertilized). When planted in spring, cuttings of successional plant species established less than those planted in early winter. Water repellency of the sand surface is supposed to be the main cause for this. Pre-grown seedlings were less susceptible for the season of planting. Direct sowing was not effective. Rabbit browsing had to be omitted to obtain successful establishment. When living rhizomes of successional plant species were still present in the soil profile, fencing alone turned out to be effective to re-establish vegetation at initially bare sites.     

The role of European marram grass in dune stabilization and succession near Cape Agulhas, South Africa

The coastline near the southern tip of Africa is characterized by large mobile dunes that are driven along wide beaches by strong winds throughout the year. This results in the blockage of the river mouths causing severe flooding of the low-lying farmland of the Agulhas Plain during the rainy winter season. Large parts of the driftsands were stabilized with the European dune pioneer species Marram grass (Ammophila arenaria), which has proved highly invasive along the North American west coast. In order to establish the potential invasiveness ofA. arenaria in South African coastal dune systems and its role in the succession of a large stabilization area, studies were carried out on De Mond Nature Reverve. Using aerial photos, maps and planting records, the vegetation of sites of various ages were sampled. By means of this chronosequence of stands, there is clear evidence that succession takes place at De Mond. Four communities are distinguished, varying from recent plantings ofA. arenaria to mature dune thicket or dune fynbos (heath) vegetation. These relate to four different stages of succession,A. arenaria occurring in reduced abundance in the older communities. After 50 years, formerA. arenaria areas are usually covered by dense dune scrub and in some places even in asteraceous dune fynbos. Succession is most rapid in sheltered, moist dune slacks, butA. arenaria remains vigorous in conditions favourable for its growht, i.e. on exposed, steep dune slopes with strong sand movement.A. arenaria does not appear to spread unaidedly at De Mond and has been successfully used for temporary dune stabilization.