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.     

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.     

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.     

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.     

Hematopoietic neoplasia inMya arenaria: Prevalence and indices of physiological condition

A field survey of hematopoietic neoplasia (Hn) in the soft shell clamMya arenaria (L.) was undertaken using an immunoperoxidase diagnostic technique. Monthly collections ofM. arenaria were made at two sites: Little Buttermilk Bay and New Bedford Harbor, both in Buzzards Bay, Massachusetts, USA, from May 1986 to October 1987. Clams were diagnosed for leukemia and analyzed for soft-tissue dry weight, condition index, and carbon and nitrogen content of the soft tissue. Prevalence of leukemia inM. arenaria exhibited a seasonal fluctuation with a maximum prevalence in fall (September to October) and a minimum prevalence in early summer (March to July). A second maximum peak in late winter (January to March) was observed at one site. Leukemia primarily affected clams that were 3 to 4 yr post-settlement. Lower prevalence levels were observed in both younger and older clams. LeukemicM. arenaria with advanced stages of the disease, were in poorer physiological condition based on dry weight of the soft tissue, condition index, and carbon content of the tissue. Nitrogen metabolism appeared to be unimpaired. Significant differences were observed between the two sites with respect to prevalence of Hn and the physiological condition of clams. Differences in disease prevalence between the two sites may be the result of unknown environmental factors that facilitate initiation of the disease or, that compromise the defense mechanisms of the clams.     

Introduction History and Invasion Patterns of Ammophila arenaria on the North Coast of California

European beachgrass ( Ammophila arenaria [L] Link), introduced to stabilize sand, is fully naturalized in central and northern California and has supplanted populations of native dune plants in many areas, including the North Spit of Humboldt Bay. We interpreted air photos of the North Spit for the presence of Ammophila arenaria at three points in time: 1939/1942, 1962, and 1989. To quantify the spread and to detect invasion patterns of Ammophila , we then compiled maps using a geographic information system. We documented introduction dates and locations to improve the accuracy of our photo interpretation. Interpretation of the three photo series revealed invasion and expansion of Ammophila in both foredunes and inland dunes. The most dramatic increase in cover was found in the foredunes, whereas documented and presumed plantings were found mostly on inland dunes. Adjusting for intentional eradication, Ammophila cover on the North Spit was found to have increased between 1939 and 1989 by 574%. The square root of the area occupied by Ammophila on the North Spit increased linearly with time, conforming to biological invasion theory.     

Age determination of the soft-shell clam Mya arenaria using shell internal growth lines

Shells of known-age Mya arenaria have revealed an internal annual growth pattern in this bivalve. The number of external growth rings on the shell exterior does not correspond to the age of the clams. This study provides an alternative method to using external annual rings for age determination. The internal growth lines are visible in thin sections when the shells are cut from the umbo to the ventral margin. The lines are formed by late spring of each year prior to spawning and are probably in response to decreased winter growth. When the 7 year olds were examined it was difficult to distinguish the most recent growth line and the inner shell layer. This could result in ages being underestimated. This annual pattern of internal growth lines could be useful to biologists and paleoecologists engaged in population studies.     

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.     

The effect of sea-water submergence on rhizome bud viability of the introducedAmmophila arenaria and the nativeLeymus mollis in California

Ammophila arenaria, an invasive European beach grass, dominates most United States Pacific coast beaches north of San Francisco Bay, and it appears to severely reduce opportunities for regeneration of native plant species, including American beach grass,Leymus mollis. The knowledge of how longAmmophila rhizomes can survive in sea-water is important for long-tern management strategies, which must consider the probability of reinvasion of areas whereAmmophila has been eradicated. The bud viability of bothAmmophila arenaria andLeymus mollis remained high following submergence in sea-water for 7 days, andLeymus bud viability was still high after 13 days submergence. In fact,Leymus bud viability appears to be enhanced slightly by submergence for 7 days in sea-water. SinceAmmophila rhizomes retain a mean bud viability of >50% following submergence for 7 days, there is clearly the potential for long distance dispersal to other beaches. Even after 13 days of submergence,Ammophila rhizomes still had a mean bud viability of 8.5%. Assuming near-shore current speeds of 5–45 cm/sec, viableAmmophila rhizomes can be transported up to 505 km in 13 days.     

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.     

The capacity of the filter-feeding bivalve Mya arenaria L. to affect water transport in sandy beds

Benthic filter feeding macrofauna organisms may be an important factor linking sediment and water column. Transport of water and concomitantly of suspended matter is directly related to the size of the benthic filter feeding population. This paper aims to determine the potential for water transport by the bivalve Mya arenaria along a coastal stretch of roughly 100 km length in the southern Baltic Sea. Quantification of population filtration rates specific to the area is based on distribution, abundance and biomass of M. arenaria and calculated according to previously published filtration rate–biomass relations. Calculated rates range up to >8 m³ m^–2 day^–1 (at 5–20 m water depth in sandy sediment) with the potential to locally process a volume of water equivalent to the water column within <1 day. Data from 1991–2002 at one site suggest that the area-specific potential population filtration rate remains remarkably constant in time despite changes in population structure of M. arenaria. The related impact on pore water exchange within the permeable sediment associated with the leakage of water from the gape of M. arenaria valves is discussed.Communicated by L. Hagerman, Helsingør     

Vegetation development in coastal foredunes in relation to methods of establishing marram grass (Ammophila arenaria)

In coastal foredunes marram grass (Ammophila arenaria) is used to stabilize windblown sand. The development of traditionally plantedAmmophila into a more natural foredune vegetation may take 5 – 10 yr. For economic reasons, traditional planting may be replaced by alternative techniques such as planting seeds or disk-harrowing rhizome fragments. In this paper, we compare the initial vegetation development of traditionally planted stands with stands established from seeds and from rhizomes. The experiments were conducted on an artificial foredune originating from dredged sea sand. The total experimental area covered more than 100 ha and the vegetation development was studied for 6 yr. The data were analysed bya priori grouping of plant species according to their ecology, as well as by Principal Components Analysis (PCA) and Redundancy Analysis (RA) of the percentage ground cover per plant species. Comparing ecological groups of plants showed that all planting methods delivered equal numbers of plant species that are indicative for coastal dunes. PCA and RA showed that methods based on the use of rhizome material resulted in a higher percentage cover of clonal perennials (Calammophila baltica, Festuca rubra ssp.arenaria, Carex arenaria andCirsium arvense) than the traditionally planted stands and the stands obtained from seeds. The latter two were characterized by the dominance of annuals, bi-annuals and (mostly nonrhizomatous) perennials. Initially, the rates of succession were highest in the stands obtained from rhizomes. However, after 3 – 6 yr there were no differences between the various stands. During the first four years, the percentage cover by rhizomatous foredune plants developed faster than that of annuals, bi-annuals and perennials. After 6 yr, the latter contributed almost as much to the percentage cover as the clonal species. Nomenclature: van der Meijden et al. (1990) for plant species. The plant species were classified into ecological groups according to van der Meijden et al. (1991) and Mennema et al. (1980).     

Reduction in carbon flux in Mya arenaria caused by a spill of No. 6 fuel oil

Rates of respiration, assimilation and filtration have been determined on a monthly basis for two populations of Mya arenaria in Casco Bay, Maine (USA), one of which was heavily oiled by a spill of No. 6 fuel oil. Monthly estimates of carbon flow have been calculated from these data. The same general trends were seen in each population; small negative carbon flow in winter, large negative carbon flow during the spring; large positive carbon flow during summer. However, the oiled population gained carbon at only 50% of the rate seen in the unoiled population. This difference results from the uptake of hydrocarbons by the oiled population.Contribution No. 75009 to the Bigelow Laboratory.     

Influence of various concentrations of orthophosphate on the division rate of an estuarine benthic diatom,Navicula arenaria,in culture

A hydrolysis procedure along with a high-pressure liquid chromatographic procedure is given enabling simple and reliable thymine determinations in the nanogram range in different fractions of sea-water samples taken from three different locations in the Northern Adriatic Sea. The levels corresponded to 1–3 g DNA per liter. From total polyanionic thymine, which had been precipitated as the cetyltrimethylammonium salt, the highest percentage was linked to the particulate fraction, with a definite subsurface minimum at 10 to 15 m. There was a corresponding maximum of a high molecular non-particulate thymine-containing fraction at the corresponding depth. From the bottom at 30 m upwards to about 20 m, a low molecular thymine-containing material has been found. Remarkably these basic features were common to all three locations, one of which was supposed to be in clean water, one near a thickly settled, touristic area, and the other in front of a large river delta coming from industrial hinterland.     

Pre-growth mortality of Abies cilicica trees and mortality models performance

In this study, we compared tree-growth rates (basal area increment) from recently dead and living Taurus fir (Abies cilicica Carr.) trees in the Kovada lake Forest of Isparta, Turkey. For each dead tree, tree-growth rates were analyzed for the presence of pre-death growth depressions in the study area (number of sample plots = 11) in 2006. However, we compared both the magnitude and rate of growth prior to death to a control (living) group of trees. Basal area increment (BAI) averaged substantially less during the last 10 years before death than for control trees. Trees that died started diverging in growth, on average, 50-60 years before death. About 18% of trees that died had chronically slow growth, 46% had pronounced declines in growth, whereas 36% had good growth up to death. However, tree-ring-based growth patterns of dead and living Taurus fir trees were compared and used 12 mortality models that were derived using logistic regression from growth patterns of tree-ring series as predictor variables. The four models with the highest overall performance correctly classified 43.8-56.3% of all dead trees and 75.0-87.5% of all living trees, and they predicted 25.0-43.8% of all dead trees to die within 0-15 years prior to the actual year of death.     

Water hardness and cardiovascular mortality in Finland

In Finland, where mortality rates of cardiovascular diseases are as a rule among the highest in the world, significant areal differences in these figures are encountered especially between the eastern and western parts of the country. To test the hypothesis that these differences,( the reason of which has long been a subject for a lively debate in Finland), may result from geochemical factors such as variations in the hardness or calcium or magnesium content of well water a correlation analysis was carried out in which the geochemical properties in each of the country's individual administrative districts, or communes (444 in all, excluding Åland) were compared with the percentage of heart diseases among the causes of death from diseases in 1991. The results show a poor or extremely low correlation nation-wide (r ranging from –0.039 to +0.045), indicating a virtual absence of any causal relationship. On the other hand, taken pairwise, the extreme western and eastern provinces, Vaasa andNorthern Karelia, show marked differences both in their geochemical and mortality data, the percentage of deaths caused by cardiovascular diseases being markedly lower in the province of Vaasa, where the hardness and the magnesium content of well water (but not the calcium content) are significantly higher (p < 0.001 and 0.002 respectively). This inverse relationship is in line with the supposition that in addition to other factors, geochemical differences may have an effect on mortality.     

Copper-induced differential mortality in the mussel Mytilus edulis

Copper toxicity experiments were performed with 3 000 mussels (Mytilus edulis L.) from a population that is a mixture of individuals originating in the Kattegat and in the Baltic. Subsequently, the electrophoretic patterns of the enzymes phosphoglucose isomerase (PGI) and phosphogluco mutase (PGM) were analysed in the dead and surviving mussels in order to detect possible differential mortality. Significant differences in allele frequencies between dead and surviving mussels were found for PGI. The results are discussed in relation to genotypic differences in the PGI and in relation to the composite structure of the investigated population. It is concluded that the differential mortality is due to genotypic differences in the PGI locus and/or to different susceptibility to copper by mussels originating in the two geographic areas.     

Climate change, humidity, and mortality in the United States

This paper estimates the effects of humidity and temperature on mortality rates in the United States (c. 1973-2002) in order to provide an insight into the potential health impacts of climate change. I find that humidity, like temperature, is an important determinant of mortality. Coupled with Hadley CM3 climate-change predictions, I project that mortality rates are likely to change little on the aggregate for the United States. However, distributional impacts matter: mortality rates are likely to decline in cold and dry areas, but increase in hot and humid areas. Further, accounting for humidity has important implications for evaluating these distributional effects.     

Incidental capture,direct mortality and delayed mortality of sea turtles in Australia's Northern Prawn Fishery

The species composition, catch and mortality rates of sea turtles captured incidentally by the tiger prawn fishery on Australia's northern coast in 1989 and 1990 were estimated by monitoring the fishery's catch. In 1990, the delayed rate of mortality from damage was estimated and the size composition was measured. Five species of turtles were captured: the flatback (Natator depressa, 59% of the total), loggerhead (Caretta caretta, 10%), olive ridley (Lepidochelys olivacea, 12%), green turtle (Chelonia mydas, 8%) and hawksbill (Eretmochelys imbricata, 5%). The turtle catches varied with water depth: the highest catch rates (0.068±0.006 turtles per trawl) were from trawls in water between 20 and 30 m deep, relatively few turtles (10%) were captured in water deeper than 40 m (25% of trawls). Catch rates varied with time of year: the highest catch rates were 0.098 (±0.013) turtles per trawl in winter. There was no significant difference in the overall catch rate (²= 0.047; p=0.8111; df=1) but a significant difference in mortality rate (²= 3.99; p<0.05; df=1) between the two years. The incidence of capture in the commercial fishery was 0.051 (±0.003) turtles per trawl towed for about 180 min, with 0.007 (±0.001) turtles per trawl drowning in the nets. There were no significant differences in the catch and mortality rates between the two years for any of the turtle species except the loggerhead, which had a significantly (² = 11.029; p=0.0013; df=1) lower catch rate in 1990 (0.002±0.001 turtles per trawl) than in 1989 (0.008±0.002 turtles per trawl), and a significantly higher mortality in 1990 (33%) than in 1989 (19%). Catch rates and mortality varied between the species: the flatback had the highest catch rate (0.030±0.002 turtles per trawl) but the lowest mortality (10.9%); the loggerhead had a catch rate of 0.005±0.001 turtles per trawl, and high mortality (21.9%); the olive ridley had a catch rate of 0.006±0.001 turtles per trawl and a low mortality (12.5%); the green turtle's catch rate was 0.004±0.001 per trawl and mortality 12.0%; the hawksbill had the lowest catch rate (0.002±0.001 turtles per trawl) but highest mortality (26.4%). Based on the fishing effort (27 049 d for 1989 and 25 746 d for 1990), we estimate that 5 503 (±424) turtles were caught and returned to the sea in 1989 and 5 238 (±404) in 1990, of which 567±140 drowned in 1989 and 943±187 in 1990. In 1990, an estimated 25% of all captured turtles suffered some non-lethal damage; an estimated 21% of turltes were captured comatose and 4% were injured. We conclude that, considering other threats, trawl-induced drowning is not the major impact on turtle populations in northern Australia, but that measures to reduce drowning and delayed mortality would be desirable.