Age structure,growth, and morphometric variations in the Atlantic surf clam,Spisula solidissima,from estuarine and inshore waters

The surf clam, Spisula solidissima (Dillwyn), population in the estuarine waters of Long Island Sound, New York, USA, was characterized in 1984 and again in 1988 by an age structure restricted to just two age-classes, and an apparent lifespan of only about 10 yr. In the inshore coastal waters off Fire Island, New York, a wide age range from 2 to 22 yr old was present. The age structure at Long Beach, New York, a third geographic region which is coastal but influenced by the Hudson River estuary, was similar to Long Island Sound. Juvenile surf clams grew at similar rates in all three geographic regions. However, adults from the Long Island Sound population grew significantly slower and reached an asymptotic maximum size which was 37% smaller than Fire Island adults. Long Beach adults had intermediate growth rates and maximum sizes. The shells of Long Island Sound clams were also 25% thinner than those from the other two regions. Density dependent effects on growth, evaluated over abundances ranging from 0.5 to 294 ind. m^–2, were present but were too small to account for observed regional differences. Results suggest that adult surf clams may be physiologically stressed by the reduced salinity and more extreme temperatures found in estuarine waters.     

Density-dependent growth in the Atlantic surfclam, Spisula solidissima, off the coast of the Delmarva Peninsula, USA

Spisula solidissima (Dillwyn, 1817) is a large, suspension-feeding bivalve, whose range extends from Nova Scotia to South Carolina. This species is harvested commercially. Shell length and age data were collected for this species from 1980 to 1994 during surveys of population size and structure. These data were used to examine the relationship between the growth rate of S.␣solidissima and intraspecific density. The null hypothesis was that density (represented by number of individuals per tow) would have no effect on rate of growth. A negative relationship would support the alternative hypothesis, that intraspecific competition had taken place. This analysis focused on the surfclam population offshore from the Delmarva Peninsula, USA because: (1) a major recruitment event occurred in 1977, (2) clam fishermen had reported “stunted” surfclams in that area, (3) a wide range of local densities were available to examine, and (4) the existence of a closed area within the study area set up an interesting contrast with areas left open to harvesting. Maps of surfclam abundance across the Delmarva region demonstrate that areas of highest density have generally remained in the same location through time. The results suggested that intraspecific competition has been important in structuring this population. Based on data from 1980 to 1992, shell length was significantly reduced at high density, and a significant interaction between age and density was observed. Growth modeling indicated decreased asymptotic lengths and growth rates with increasing density. In nine out of ten pairwise randomization tests, fitted von Bertalanffy growth curves, representing different densities, were significantly different from each other. High densities of clams have persisted in the area that was closed to harvesting for 11 years (1980 to 1991). In 1994, length at age was significantly less in this closed area compared to that in the surrounding area. This effect was apparent in clams from 3 to 17 years of age, and most pronounced in the cohort that recruited to the Delmarva region in high numbers in 1977. Lower growth rates within the closed area have management implications for the optimal duration of closures. Received: 16 May 1997 / Accepted: 7 October 1997     

Growth of the Atlantic surfclam,Spisula solidissima,from Georges Bank to the Delmarva Peninsula,USA

Age/shell length data for offshore surfclam, Spisula solidissima (Dillwyn, 1817), populations were used to estimate the parameters of the von Bertalanffy growth model by time period and region, from Georges Bank in the north to the Delmarva Peninsula in the south. Randomization tests were used to compare curves. We tested the a priori hypothesis that growth curves would change over time in the south (i.e., New Jersey and Delmarva) but remain constant in the north (i.e., Long Island and South New England). This hypothesis was proposed because surfclam population structure in the south had been altered by the hypoxic event of 1976, and possibly by intense, long-term commercial harvesting. Northern regions, unaffected by these factors, served as natural controls. Based on a comparison of data collected in 1980 with pooled data from 1989 and 1992, the hypothesis was supported. Both the growth coefficient (k) and maximum shell length (L ) declined between two time periods in the two southern regions, while during the same time interval, no change occurred in the two northern regions. Differences in growth between regions were often statistically significant. For example, compared with the southern regions, the growth coefficient on Georges Bank was larger, and those clams attained a smaller maximum length. In a comparison of adjacent regions from Delmarva to S. New England, k increased from south to north. This could imply faster growth in cooler water, as well as no relationship between growth and primary productivity. Alternatively, size-selective mortality, imposed by the commercial fishery, was discussed as a mechanism that might account for this unexpected pattern.     

Age-structure, recruitment, and adult mortality in populations of the Atlantic surfclam, Spisula solidissima, from 1978 to 1997

The Atlantic surfclam, Spisula solidissima (Dillwyn, 1817), is a dominant member of the benthic community on the continental shelf from Georges Bank to North Carolina, USA. This bivalve has supported a major fishery, primarily off New Jersey and the Delmarva Peninsula, since the 1960s. Early papers documented that these populations were at historical lows in the mid-1970s owing to commercial harvesting and a hypoxic event off New Jersey. It was also shown that major recruitment took place off New Jersey in 1976 and off the Delmarva Peninsula in 1977. Because the size frequencies of surfclams from federal surveys do not show distinct year classes, there has been uncertainty about the number of year classes in these populations throughout the 1980s and 1990s. The present study describes changes in population age- and size-structure from 1978 to 1997 in federal waters (≥5.5 km from shore) of the USA. Given the 30 to 35 year life span of S. solidissima, these populations could be composed of many year classes. Yet, these populations were composed of only two to three year classes in 1978. Through annual recruitment, the number of year classes increased over time, and populations off New Jersey and the Delmarva Peninsula contained at least 19 year classes in 1997. This major change in population structure over time was not evident from examination of available size-frequency data, and could only be inferred from data on age-composition. Throughout the 1980s and 1990s, the surfclam fishery was supported by multiple year classes. The mean and variance of recruitment to the New Jersey region, as indicated by the abundance of 4-year-olds over time, was greater than that off Delmarva, particularly between 1980 and 1986. The instantaneous rate of adult mortality, which includes the effect of harvesting, was approximately 0.26 yr⁻¹ in each region. Received: 11 September 1998 / Accepted: 1 February 1999     

Studies on the digestive proteases of the milkfish Chanos chanos

The protease activity of crude extracts from various organs of the digestive tract of two groups of milkfish was determined. One group (Sample A) derived their food from ponds that had predominantly unicellular algae while the other group (Sample B) were reared on ponds dominated by the filamentous green algae Chaetomorpha brachygona. In general, crude extracts from Sample A fish had a higher protease activity than Sample B fish. In both samples, high protease activity was observed in crude extracts from the pyloric caeca, intestines and pancreas. Chymotryptic activity was observed in crude extracts of pancreas, intestines and pyloric caeca of both fish samples. Tryptic activity was, however, observed only in fish grown on unicellular algae. Experimental evidence suggests that a powerful trypsin inhibitor in Chaetomorpha brachygona may account for the absence of tryptic activity in all crude extracts of Sample B fish. The presence of this inhibitor may also explain the widely observed poor growth rate of milkfish reared on this natural food.SEAFDEC contribution no. 112     

Studies on the carbohydrases in the digestive tract of the milkfish chanos chanos

Crude extracts from various regions of the digestive tract of pond grown milkfish were tested forttheir ability to catalyze the hydrolysis of various carbohydrates. The most active carbohydrases were those involved in the hydrolysis of -glucosidic bonds. Maltose, trehalose, dextrin, starch and glycogen were rapidly hydrolyzed in the presence of crude extracts from the intestines and the pyloric caeca. High amylase activity was observed in extracts from the intestines, pancreas, pyloric caeca and liver. The intestinal amylase had optimum activity at pH 6.2 and at a temperature of about 50°C. It was active at a chloride concentration of 10 to 40 ppt. The amylase activity in the intestines consistently peaked daily at about noon when the milkfish gut was full. In contrast, enzyme activity was significantly lower at 0030 hrs when the gut was empty. These results are consistent with earlier observations that the milkfish is a daytime feeder and suggest further that intestinal amylase secretion is in phase with the feeding activity of the milkfish. Although the fishes used in this study fed mostly on the naturally occurring algae in the ponds, no cellulase activity was detected in any region of the digestive tract. Less active carbohydrases that were detected include a -glucosidase and -galactosidase, both of which were of limited substrate specificity.SEAFDEC Contribution No. 71     

Blooms of surf-zone diatoms along the coast of the Olympic Peninsula,Washington. XI. Regeneration of ammonium in the surf environment by the Pacific razor clam Siliqua patula

A 20 month field study was conducted on ammonium excretion rates of the Pacific razor clam Siliqua patula Dixon along the beaches of Washington State, USA. Excretion rates of all those nutrients likely to be regenerated in sufficient quantity to affect surf diatom growth were measured; ammonium appeared to be the most important metabolite. Excretion of ammonium by razor clams far exceeded that by other beach fauna. Ammonium excretion rates of razor clams were positively correlated with shell length, but no correlation between ammonium excretion rate and water temperature was evident. This may be an artifact or may represent some degree of seasonal acclimation of the species to temperature. Weight-specific ammonium excretion rates were negatively related to clam size, indicating a possible large (and unknown) contribution of regenerated ammonium by smaller clams in their first year of growth; smaller clams were rarely captured during this study.Contribution No. 1048 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA.     

Preliminary report on the digestive enzymes present in the digestive gland ofPerna viridis

Quantitative data on digestive enzymes of bivalves are very limited and so this study was performed to obtain such information, and to find out whether some of these enzymes are well adapted to conditions in the digestive gland. The green musselPerna viridis was obtained from Clementi West Market in 1985. The results show that various carbohydrases are present in the digestive gland. The activities of-amylase, cellulase and-glucosidase are higher than that of other carbohydrases, whilst the activities of both lipase and protease, especially the former, are low. The pH optima of the following enzymes are also given:-amylase = 5.8; laminarinase = 5.5;-glucosidase = 5.8; trehalase = 4.8;-glucosidase = 5.5. The only enzyme which was not well adapted to conditions in the digestive gland (pH 6.2) was trehalase. The wide spectrum of enzymes present in this bivalve indicates that it has the ability to utilize a wide range of nutrients efficiently.     

Blooms of surf-zone diatoms along the coast of the Olympic Peninsula,Washington. X. Chemical composition of the surf diatom Chaetoceros armatum and its major herbivore,the Pacific razor clam Siliqua patula

The surf diatom Chaetoceros armatum T. West, collected from its natural habitat along the Washington coast (USA), had a large inorganic component in the form of a clay coat (consisting of clay minerals, illite and montmorillonite) surrounding the cell chains: 63% of the dry weight of the natural material collected in November was inorganic. The organic fraction was composed of 67.6% lipid, 29.7% protein, and 1.3% carbohydrate. Culture cells were likewise high in protein and lipid and low in carbohydrate. Traces of chitin found in field samples were probably a contaminant, since chitin was absent from cultured cells. This diatom species serves as a major food source throughout the year for the Pacific razor clam Siliqua patula Dixon, which inhabits these same beaches. Protein constituted 47% and lipid 42% of the dry weight of razor clam tissue. The fatty acid distribution in the diatom lipid resembled that previously reported for other diatom species; similarly, the fatty acid distribution of the clam lipid was similar to that previously reported for other bivalve molluscs. The clam fatty acids differed in chain length and degree of saturation from those extracted from its food source, indicating an active fatty acid metabolism in the clam.Contribution No. 995 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA.     

Studies on digestion and the fine structure of digestive caeca in Eurydice pulchra (Crustacea: Isopoda)

In the carnivorous isopod Eurydice pulchra Leach the mid-gut caeca have a pH of 5.6 and produce a strong acid proteinase, a lipase and a carbohydrase. Electron microscope studies of the digestive caeca of starved and fed animals show that secretion is merocrine and may originate from a complex involving the basal cell membrane and associated mitochondria, producing lysosomes. Absorbed materials, in fed animals, appear to take the form of neutral lipid bodies, glycogen, and proteinaceous crystals, the latter forming within the golgi complex. Digestion of both lipid and protein inclusions was observed to occur intra-cellularly by fusion with free lysosomes. Secretion and absorption may occur in the same cell, and the scheme for the sequence of epithelial cell stages is proposed.     

Persistent blooms of surf diatoms along the Pacific coast,USA

Productivity was studied in two diatom species, Chaetoceros armatum T. West and Asterionella socialis Lewin and Norris, which form persistent dense blooms in the surf zone along the Pacific coast of Washington and Oregon, USA. Past observations have shown that surf-diatom standing stock usually declines in summer along with concentrations of nitrate and ammonium. Using the ¹⁴C method, photosynthetic rates in natural surf samples were measured monthly for one year (October 1981 through September 1982) at a study site on the Washington coast. Also measured were temperature, salinity, dissolved nutrients, particulate carbon and nitrogen (used as estimates of phytoplankton C and N), and chlorophyll a. Assimilation numbers (P _max) were higher in summer (5 to 8 g C g^-1 chl a h^-1) than in winter (3 to 4gC). Specific carbon incorporation rates (µ_max) showed no obvious seasonality, mostly falling within the range of 0.09 to 0.13 g C g^-1 C(POC) h^-1. The discrepancy between the seasonal trends for chlorophyll-specific and carbon-specific rates reflects a change in the carbon-to-chlorophyll ratio. Because of seasonal differences in daylength and light intensity, daily specific growth rates () are thought to be higher in summer than in winter. Neither ammonium enrichment assays nor particulate carbon-to-nitrogen ratios provided convincing evidence for nitrogen limitation during summer, and the observed changes in diatom abundance cannot be explained on this basis. Both the high diatom concentrations and their seasonal variations probably are due mainly to factors other than growth rates; two factors considered important are diatom flotation and seasonal changes in wind-driven water transport. C. armatum usually dominates the phytoplankton biomass in the surf zone, and evidence suggests that this species is strongly dominant in terms of primary production.Contribution No. 1391 of the School of Oceanography, University of Washington, Seattle, Washington, USA     

Sublethal effects of chronic oil exposure on the intertidal clam Macoma balthica

When exposed to Prudhoe Bay crude oil in flowing seawater for 180 days, the small intertidal clam Macoma balthica showed behavioral, physical, physiological and biochemical changes. At a high concentration of oil in seawater (3.0 mg l^-1) burrowing rate decreased, respiration rate increased, growth was inhibited, and very high mortalities resulted. The lowest concentration of oil in seawater (0.03 mg l^-1) inhibited growth and caused reabsorption of gametes. One group of adverse oil effects which was related to sluggishness and disorientation of the clams appeared after a week of exposure to oil; another group related to a negative energy balance was not observed until 60 days. We conclude that chronic exposure of M. balthica to oil-in-seawater concentrations even as low as 0.03 mg l^-1 will, in time, lead to population decreases.Please address requests for reprints to Dr. D. G. Shaw at the Institute of Marine Science     

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.     

Influence of field-based nutrient enrichment on the photobiology of the giant clam Tridacna maxima

Nutrients were added to 12 microatolls in One Tree Island lagoon every low tide for 13 mo to an initial concentration of 10 μM (ammonium, N) and 2 M (phosphate, P). These concentrations remained above background for 2 to 3 h after addition. The addition of ammonium (N and N+P but not P alone) significantly increased P _g (gross photosynthesis) P _n (net photosynthesis) and R (respiration) per unit wet-tissue weight and α (photosynthetic efficiency) in Tridacna maxima after 3 mo nutrient enrichment. These responses to small and transient changes in ammonium concentrations suggest that symbiotic clams are not nutrient-replete, and that even subtle changes in nutrients can have a measurable effect on photosynthesis. The same clams did not show significant differences in photosynthetic parameters 6 mo after the beginning of nutrient enrichment, suggesting that their previous responses had either been seasonal or that symbiotic clams such as T. maxima are able to adjust their photophysiology following external changes in nutrient concentrations. Received: 26 August 1997 / Accepted: 11 December 1998     

Growth,mortality and recruitment of the yellow clam Mesodesma mactroides on Uruguayan beaches

We studied some aspects of the population dynamics of the yellow clam Mesodesma mactroides (Deshayes, 1854) which inhabits the eastern sandy beaches of Uruguay. Observations were made from March 1983 through March 1985 (harvesting season), and additional data from January 1988 to December 1989 (closed season) were also included in the analyses. Parameters of growth and mortality were estimated by methods that make use of age or length information. An age/length relationship was constructed by counting growth rings. Results were consistent with those obtained with modal class progression analysis and other length/frequency methods. The growth curve obtained from the age/length relationship did not account for seasonality in growth rate, and a seasonal growth model explained growth oscillations in a better way. The instantaneous coefficient of natural mortality (M) estimated for the closed season was 1.64, whereas an increase in fishing mortality (F) was observed during 1984. Age composition did not change during the 2 yr of study, and the first 6 mm class accounted for over 50% of the population. The main contribution of recruits was observed between late summer and early fall (February to April), and the recruitment pattern suggested one main settlement period per year. Growth and recruitment success appeared to be regulated by density-dependence processes. Some implications for management are considered in this context. Methodological aspects concerning the reliability of empirical equations and programs for estimating growth and mortality are discussed.     

A stochastic bioeconomic model for the management of clam farming

The Manila clam Tapes philippinarum is one of the most important commercial mollusc species in Europe. Intensive clam farming takes place in several coastal lagoons of the Northern Adriatic Sea, supporting local economy but raising the problem of the environmental sustainability of this activity. In this work, we propose a bioeconomic model that provides guidelines for an efficient management of intensive clam farming. Clam demography is described by a stochastic model of growth and survival, accounting for the effect of water temperature, seeding substratum and density dependence of vital rates. The model is calibrated on and applied to the case of Sacca di Goro, a lagoon located in the Po River Delta (Northern Italy). We consider two distinct management criteria: the optimisation of the marketable yield and the optimisation of monetary benefits, respectively. The use of a stochastic formulation allows us to reveal the existing trade-off between maximizing the median yield or profit and minimizing its variance. A Pareto analysis shows that seeding in spring or fall on sandy substrata and harvesting 18 months later provides the best compromise between these two contrasting objectives, maximizing profits while minimizing the associated uncertainty level. Finally, we show that seeding clams at high densities (more than 750 clams m⁻² on muddy substrata and more than 1500 elsewhere) can have not only a potentially negative impact on the ecological sustainability of clam farming, but also a negative economic effect.     

Reproductive ecology of the brooding,hermaphroditic clam Lasaea subviridis

The reproductive ecology of a small intertidal clam was studied for two years (September 1980-November 1982) at two mid-intertidal sites on the northern shore of Monterey Bay, California. Lasaea subviridis is hermaphroditic and incubatory; the young are released as juveniles. Brooding adults are found throughout the year. All embryos in a clam are at the same stage of development, but the number and stage of embryos varies among clams at any one time. Individuals are iteroparous and produce small numbers of oocytes and sperms simultaneously. All stages of gonad development are found in different individuals at any one time. There was no evidence of seasonality. Young reared in the laboratory after being dissected from the parent reached brooding size (shell length of 1.7 mm) within 6 mo. Laboratory experiments indicated that juveniles crawl away soon after release from the parent. These reproductive characteristics may contribute to this taxon's ability to colonize new environments rapidly.     

Host glutamine synthetase activities in the giant clam—zooxanthellae symbiosis: effects of clam size,elevated ammonia and continuous darkness

Host tissues and zooxanthellae of the giant clam Tridacna gigas contained glutamine synthetase, with the highest transferase activities present in the gill, followed by the kidney, mantle, zooxanthellae, foot, heart and adductor muscle, in that order. Synthetase activities of glutamine synthetase in host tissues and zooxanthellae were in a similar order, but the differences were not so marked. Host tissues also contained hexokinase, glucose-6-phosphate dehydrogenase and malate dehydrogenase. Highest hexokinase activities were present in the heart, followed, in order, by the gill, mantle, adductor muscle and foot. Highest glucose-6-phosphate dehydrogenase activities were present in the gill, followed by the mantle, heart, adductor muscle and foot. All tissues assayed contained high malate dehydrogenase activities. There was no detectable glutamate dehydrogenase activity. Glutamine synthetase activity in gill and mantle tissue decreased by 1.6% with every 1 cm increase in clam size. Host glutamine synthetase activity decreased by 80% in gill tissue and by 45% in mantle tissue in clams which were maintained for 8 d in continuous darkness. Similar effects were found when clams were kept in light in the presence of elevated ammonia concentrations. It is suggested that both host and symbionts are nitrogen-deficient in small clams and that host glutamine synthetase plays a role in ammonia assimilation by the intact association.     

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.     

Mate guarding behavior in clam shrimp: the influence of mating system on intersexual conflict

Mate guarding is a male strategy to gain access to receptive females but often results in antagonistic interactions between the sexes because of different costs/benefits of guarding. In addition to social, morphological, and physiological parameters, the type of mating system should also affect the strength of the conflict and thus the guarding duration. Specifically, when compared to females, self-compatible hermaphrodites might have reduced benefits of outcrossing. We investigated mate guarding in dioecious (co-presence of females and males) and androdioecious (co-presence of hermaphrodites and males) branchiopod crustaceans. Both sexes in androdioecious systems should shift their guarding times to lower values relative to dioecious systems because (1) androdioecious males are present in lower percentages than dioecious males and thus encounter rates with receptive mates are relatively greater for them; and (2) hermaphrodites should have low incentive to incur high costs of mate guarding, having the alternative of self-fertilization, and thus should be highly eager to resist. While females preferred short guarding times, when allowed to control the guarding duration (males tethered), dioecious males did not increase their guarding duration when females (treated with muscular relaxant) could not resist, in contrast to what has previously been found for androdioecious males. This indicates that hermaphrodites are more willing to resist mate guarding than females. The among-species comparisons supported our hypotheses: compromised guarding times were significantly lower in androdioecious than in dioecious species. The introduction of a parameter (mating system) not previously investigated in mate guarding models resulted in a powerful test of mate guarding theory, adding a valuable contribution to our understanding of intersexual conflict.