Prevalence, patterns, and effects of shell damage on Geukensia demissa in South Carolina estuarine habitats

Shelled molluscs frequently exhibit a record of damage on exterior surfaces that can evidence past predation attempts and may affect survival and growth. In South Carolina populations of the ribbed marsh mussel, Geukensia demissa, >90% of the individuals and up to 60% of the total shell area are damaged. A trend toward greater amounts of damage occurred on mid-marsh compared to oyster reef mussels from the barrier beach side of inlets. Shell damage effects on survivorship and shell and tissue growth were assessed seasonally during multi- and single-season field experiments. Mussels from a common mid-marsh site were divided into size classes (~50 or 70 mm), treated to create two damage levels (undamaged and damaged), and replaced within mid-marsh exclusion cages to minimize additional shell damage. In both multi- and single-season experiments increased shell damage resulted in significantly greater mortality. Linear shell growth was unaffected by increased damage, but 50 mm mussels grew twice as fast. Shell mass increased 16–50% in the multi-season and single-season winter period, but decreased 7–12% during the single-season summer period. Tissue mass significantly decreased 31–43% in 50 mm damaged mussels, but increased by 33% for 70 mm mussels in both multi-season and the single-season winter period experiments. Shell damage did reduce tissue mass 43% in 70 mm single-season summer mussels. Experimental results indicate shell damage from a simulated increase in predation can affect negatively both survival and growth of marsh mussels. Seasonal timing of shell damage and initial mussel size also influenced the effects of sublethal predation on shell and tissue growth. The previously unrecognized importance of sublethal predation and the resultant significant negative effects of shell damage on survival and growth will affect the distribution and population dynamics of G. demissa in coastal marshes and will influence the overall contribution of ribbed mussels to estuarine ecosystems.     

Eumelanin-based coloration and fitness parameters in birds: a meta-analysis

Although melanin is the most common pigment in animal integuments, the adaptive function of variation in melanin-based coloration remains poorly understood. The individual fitness returns associated with melanin pigments can be variable across species as these pigments can have physical and biological protective properties and genes involved in melanogenesis may vary in the intensity of pleiotropic effects. Moreover, dark and pale coloration can also enhance camouflage in alternative habitats and melanin-based coloration can be involved in social interactions. We investigated whether darker or paler individuals achieve a higher fitness in birds, a taxon wherein associations between melanin-based coloration and fitness parameters have been studied in a large number of species. A meta-analysis showed that the degree of melanin-based coloration was not significantly associated with laying date, clutch size, brood size, and survival across 26 species. Similar results were found when restricting the analyses to non-sexually dimorphic birds, colour polymorphic and monomorphic species, in passerines and non-passerines and in species for which inter-individual variation in melanism is due to colour intensity. However, eumelanic coloration was positively associated with clutch and brood size in sexually dimorphic species and those that vary in the size of black patches, respectively. Given that greater extent of melanin-based coloration was positively associated with reproductive parameters and survival in some species but negatively in other species, we conclude that in birds the sign and magnitude of selection exerted on melanin-based coloration is species- or trait-specific.     

Determination of lethal dissolved oxygen levels for selected marine and estuarine fishes, crustaceans, and a bivalve

The objective of this study was to provide a database of the incipient lethal concentrations for reduced dissolved oxygen (DO) for selected marine and estuarine species including 12 species of fish, 9 crustaceans, and 1 bivalve. All species occur in the Virginian Province, USA, which is a cold temperate region. The study period was August 1987 to September 1995. Standard bioassay procedures were employed, with most tests being of 4-day duration. Up to eight lethal concentrations (LCs) between LC₀₅ and LC₉₅ were estimated. The study provides four general conclusions about determining lethal thresholds of low DO for these organisms. First, the concentration response curve of most species did not change greatly beyond day 1 of the exposure with the exception of crustacean larvae, which were usually more sensitive on day 4, possibly due to molting. Second, acute LC₅₀ values (1- to 4-day) for low DO were influenced by life-stage and habitat, with pelagic larvae generally being the most sensitive and benthic juveniles the least. Species mean LC₅₀ values ranged from 1.4 to 3.3 mg l^-1 for larvae, 1.0 to 2.2 mg l^-1 for postlarvae, and 0.5 to 1.6 mg l^-1 for juveniles. No intraspecific differences in LC₅₀ were detected between larval stages in crustaceans or with age in larval fishes. The response range between LC₀₅ and LC₉₅ was narrowest for the least sensitive organisms (0.6 mg l^-1), and broadened with sensitivity. The mean LC₁₀:LC₅₀ ratio for all species was 1.32 for larvae and juveniles, and 1.36 for postlarvae. The ratio for postlarvae represents only four species, and hence is not considered different from the other life stages. Third, variability increased with increased species and life stage sensitivity to low DO, and with endpoints of LC₁₅ and below, which reduces the certainty of some of these results. Lastly, no influence of temperatures between 20°C and 30°C was detected in a small set of tests with thermally acclimated crustacean larvae. This data set has been used to describe protection limits for juvenile and adult survival, and for larval recruitment for the case of persistent (₄ h) low DO for estuarine and coastal waters of the Virginian Province, USA.     

Bathymetric patterns of genetic variation in a deep-sea protobranch bivalve, Deminucula atacellana

The origin of the deep-sea benthic fauna is poorly understood and represents an enormous gap in our understanding of basic evolutionary phenomena. One obstacle to studying evolutionary patterns in the deep sea has been the technical difficulty of measuring genetic variation in species that are typically minute, rare, and must be recovered from extreme depths. We used molecular genetic techniques to quantify variation in the 16S rRNA mitochondrial gene within and among populations of the common protobranch bivalve Deminucula atacellana (Schenck, 1939). We analyzed 89 individuals from nine samples collected in the 1960s along a depth gradient from 1100 to 3800 m in the western North Atlantic. Genetic variability within populations is much lower than between populations, and peak haplotype numbers occur near the center of its depth distribution. Continental slope (<2500 m) and rise (>2500 m) populations were genetically distinct despite the lack of any obvious topographic or oceanographic features that would impede gene flow. These findings indicate that the deep-sea macrofauna can have strong population structure over small (134 km) spatial scales, similar to that observed in shallow-water and terrestrial organisms. This surprisingly high biodiversity at the genetic level affords the potential for adaptation and evolutionary diversification, the ultimate historical causes of high species diversity in the deep-sea benthos. Received: 24 July 1997 / Accepted: 26 January 1998     

Studies on the causes of mortality of the estuarine bivalve Macoma balthica under conditions of (near) anoxia

Survival of the bivalve Macoma balthica in (near) anoxic seawater was studied in a static system and a flow-through system and compared with emersed exposure to air and N₂. In the static system, a decrease in pH and exponential accumulation of sulphide in the incubation medium were observed, indicating excessive growth of (sulphate-reducing) bacteria. These changes in the chemical environment were prevented by the use of a flow-through system. However, this treatment hardly affected survival time. Median mortality times were 8.3 and 9.0 days for the static and flow-through incubation, respectively. Addition of the antibiotic chloramphenicol strongly increased survival time in both systems with corresponding values of 17.9 and 23.0 days. A similar value was obtained for survival in air (LT₅₀= 21.7 days). In a second experiment (1 year later), we obtained much lower values for anoxic survival in a static system, although laboratory conditions, season and temperature were similar. The pH values were adjusted to 6.5, 7.2 and 8.2 by buffering the media (25 mM Tris-HCl), and the corresponding LT₅₀ values were 5.5, 5.7 and 4.7 days, respectively. In the presence of chloramphenicol the values were 10.8, 10.9 and 9.5 days, respectively. These values show that a slightly acidic medium increased survival time. Exposure to an atmosphere of N₂ resulted in a survival time close to that in anoxic seawater without chloramphenicol (LT₅₀= 6.4 days). Overall the results indicate that proliferation of anaerobic bacteria associated with the bivalves was the main cause of death. Since chloramphenicol also displayed a strong positive effect in the flow-through system, which prevented the accumulation of released waste products and a decrease of pH, bacterial damage must have been by injury of the tissues of the clams and not by the release of noxious compounds to the medium. Bacterial outbreaks are a part of every anoxic event (eutrophication), and therefore, in their habitats, direct bacterial infection may also be the cause of clam mortality. It is concluded that laboratory studies on anoxic tolerance, or impact of sulphide, may produce artefacts when no precautions are taken to suppress bacterial proliferation. Received: 4 July 2000 / Accepted: 6 December 2000     

Seasonal variation in reproduction and population structure of the bivalve Limopsis tajimae in the upper bathyal zone of Suruga Bay, Japan

Seasonal variation in reproduction and population size structure was investigated for the suspension-feeding bivalve Limopsis tajimae Sowerby inhabiting the upper bathyal zone (300 m deep) of Suruga Bay, central Japan. The bivalve was collected at 1- to 4-month intervals for a period of 22 months, and bottom environment was monitored concurrently to detect factors affecting seasonality in the bivalve. Bottom water temperature, organic carbon and nitrogen contents in the sediments did not exhibit seasonal variation. Size-adjusted soft-tissue weight varied slightly, but statistically significantly between stations and months. However, its seasonal pattern was not obvious, and the pattern of temporal variation was totally different between stations. The sex ratio did not deviate from 1:1, and there was no significant difference between shell lengths of females and males. Females possess both immature small oocytes and large developed oocytes in their ovaries throughout the year, suggesting that they can potentially undergo year-round continuous reproduction. The proportion of developed oocytes in each female varied greatly from month to month, although no seasonal cycle was obvious. Population size structure of L. tajimae was polymodal. A mode of the smallest size class occurred in most months, suggesting long periods of bivalve recruitment. These findings indicate that seasonal variation in reproduction of the bivalve was negligible, probably reflecting constant physical and nutritive conditions of the bottom environment. Received: 6 January 1998 / Accepted: 19 May 1998     

Contrasting pattern of mitochondrial population diversity between an estuarine bivalve, the Kumamoto oyster Crassostrea sikamea, and the closely related Pacific oyster C. gigas

The Kumamoto oyster (Crassostrea sikamea) shows a spatially restricted distribution, favoring estuarine tideland environment. On the other hand, the Pacific oyster (C. gigas) has a broader range of habitat. The present study compared the mitochondrial population structure between the two closely related species. For accurate species identification of oysters sampled from Japanese and East Asian continental coasts, we performed sequencing analysis of the mitochondrial DNA (mtDNA) and PCR-RFLP assay of the first internal transcribed spacer of nuclear rRNA genes. Then, we estimated the extent of population differentiation within each of C. sikamea and C. gigas based on the mtDNA data. Few haplotypes were shared among the sites of sampling in C. sikamea, which contrasted with an extensive haplotype sharing among C. gigas samples. We discuss the mechanisms of elevated population differentiation observed in C. sikamea in light of the ecology and the ancient ocean geography around the present-day habitats.     

Influence of environmental parameters on intraspecific variation in Fucus vesiculosus

The vesiculation and branching of Fucus vesiculosus L. were examined on plants from three different locations on the Maine Coast, USA. These localities varied in exposure to wave action and salinity. Statistical analysis of population differences was computed for 10 morphometric characteristics. It was found that decreased salinity correlated with increased vesiculation and branching. Increased wave action correlated with decreased vesiculation. Position within vertical range was not a factor in either vesiculation or branching.     

Allozyme and physiological variation in the scallop Placopecten magellanicus and a general model for the effects of heterozygosity on fitness in marine molluscs

Heterozygosity and growth rate have been correlated in many molluscs, although the phenomenon is not universal. Enhanced growth of heterozygous molluscs has been attributed to lower energetic requirements for basal metabolism. We have investigated heterozygote deficiency, and the correlation between single-locus and multiple-locus heterozygosity and phenotype in juveniles of the scallop Placopecten magellanicus (Gmelin). Six samples were collected between 22 November 1984 and 11 December 1985 at Passamaquoddy Bay, New Brunswick, Canada. On average, heterozygote deficiency was small, although somewhat higher at the octopine dehydrogenase (Odh) locus, and decreased with age. No correlation was observed between genotype and growth rate. This result and published records indicate that allozyme heterozygosity of pectinids does not influence growth to the same degree as in other bivalves. However, we have observed in P. magellanicus a correlation between allozyme heterozygosity and octopine accumulation, a trait that relates to functional anaerobiosis, and may be related to the scallop's scope for movement. We combine these results and results from the literature into a model that relates the hypothesis of associative overdominance (at the population genetics level) with the hypothesis of energetic efficiency (at the physiological level). The model assumes that energy savings attributed to heterozygosity are used to maximize fitness. In freely moving molluscs this results in enhanced activity (such as searching for prey or swimming away from a predator), and in sessile molluscs either in enhanced somatic growth in juveniles or gonadal growth in adults.     

Influence of seasonal variation in temperature, salinity and food availability on module size and colony growth of the estuarine bryozoan Conopeum seurati

Zooid size and colony growth of the estuarine bryozoan Conopeum seurati (Canu) (order: Cheilostomatida; suborder: Malacostegina) were examined over 15 mo at Avonmouth Dock, Avon, England. Data were analysed in conjunction with synchronous measurements of temperature, salinity and food availability. Zooid length, width and area were strongly temperature-dependent, while both food availability and colony growth rate had no significant effect on zooid length, width or area. Salinity and the interaction of temperature and salinity significantly influenced zooid length and area, suggesting that changes in zooid size may result from oxygen limitation in warm waters. The validity of a number of other mechanisms proposed to account for temperature-related changes in zooid size is discussed. The results support the use of zooid size as an indicator of both long-term trends and seasonal variations in temperature in Recent and fossil assemblages as long as data sets are large and the effects of other factors on zooid size are considered. Colony growth rate was found to be significantly influenced by both the amount of food available to the colonies and the combined effect of temperature and food availability, suggesting that growth rate increases as food increases, but that the former may be limited at low temperatures when metabolic rates are low. Received: 30 March 1999 / Accepted: 24 September 1999     

Feeding and cleaning mechanisms in the suspension feeding bivalve Mytilus edulis

Feeding and pseudofeces formation were studied in intact Mytilus edulis clearing suspensions of graphite particles, and the processes were compared with activities observed in mussels with a severed adductor muscle. Most particulate material in the stomach was present in the suspended state. Ingestion of particles in suspension could take place concurrently with the production of pseudofeces. Severing of the adductor muscle resulted in profuse secretion of mucus that gradually subsided to a constant low level. Addition of graphite particles at concentrations that did not cause mucus secretion in the intact mussels strongly stimulated secretion in the operated mussels, resulting in the accumulation of highly viscid, mucusparticle aggregates. These aggregates were mechanically stable, in contrast to the fragile pseudofeces. It is indicated that normal feeding depends upon hydromechanical mechanisms that produce highly concentrated suspensions of particles for ingestion, and that mucociliary mechanisms serve to clean the gills and other organs of the mantle cavity for excess particulate material. It is further indicated that intact mussels secrete mucus only in the amounts needed to consolidate excess particulate material, and that lesions affect the normal balance between particles in suspension and mucus secretion by enhancing the sensitivity of the mechanisms that control mucus secretion.     

Selectivity in feeding by the deposit-feeding bivalve Macoma nasuta

Macoma nasuta Conrad is primarily a deposit-feeding bivalve sucking the top millimeter of the sediment surface. Growth experiments show that surface sediment supports growth better than detritus falling from the water column. Gut clearance time is between 1 and 9 h (12°C). Fecal pellets are ejected in a regular rhythm. However, the total amount of feces per unit time shows considerable individual variation. Due to sorting in the mantle cavity, about 97% (dry weight) of the surface material is ejected again as pseudofeces. Selectivity by the bivalve is estimated by comparison of particle size and organic composition of sandy and muddy sediments and compared with feces produced by clams fed these sediments. Fecal pellets are in all cases richer in organic components than the sediment, indicating a high degree of selectivity. Ingestion and digestion of small animals (meiofauna) occur, but many of the ingested specimens survive. It is not possible to estimate the assimilation of organic matter by simple difference between the ingested sediment and the ejected feces. The difficulties in calculating energy budgets which arise from selective feeding and associated bacteria are discussed.     

Sclerotized protein in the shell matrix of a bivalve mollusc

The epsilon-amino groups of lysine and phenolic groups of tyrosine are most heavily concentrated in the newlydeposited organic matrix of the shell of the bivalve Mercenaria mercenaria. A phenoloxidase enzyme which oxidizes L-dihydroxyphenylalanine is present only in this new area of the shell matrix. Scanning electron micrographs of calcified secretions of the shell show that accretion lines, thought to be layers of organic matrix separating diurnal acceretions of calcium carbonate, are not developed until up to 4 d after deposition of shell material. These results suggest that the shell matrix is hardened by some kind of polymerization, and that lysine and tyrosine residues in the matrix are involved in the process. Accretion lines in polished and etched sections become visible only after complete hardening of the polymer occurs.     

Phenotypic variations in the gills of the symbiont-containing bivalve Lucinoma aequizonata

The marine bivalve Lucinoma aequizonata (Lucinidae) maintains a population of sulfide-oxidizing chemoautotrophic bacteria in its gill tissue. These are housed in large numbers intracellularly in specialized host cells, termed bacteriocytes. In a natural population of L. aequizonata, striking variations of the gill colors occur, ranging from yellow to grey, brown and black. The aim of the present study was to investigate how this phenomenon relates to the physiology and numbers of the symbiont population. Our results show that in aquarium-maintained animals, black gills contained fewer numbers of bacteria as well as lower concentrations of sulfur and total protein. Nitrate respiration was stimulated by sulfide (but not by thiosulfate) 33-fold in homogenates of black gills and threefold in yellow gill homogenates. The total rates of sulfide-stimulated nitrate respiration were the same. Oxygen respiration could be measured in animals with yellow gills but not in animals with black gills. The cumulative data suggest that black-gilled clams maintained in the aquarium represent a starvation state. When collected from their natural habitat black gills contain the same number of bacteria as yellow gills. Also, no significant difference in glycogen concentrations of the host tissues was observed. Therefore, starvation is unlikely the cause of black gill color in a natural population. Alternative sources of nutrition to sulfur-based metabolism are discussed. Denaturing gradient gel electrophoresis (DGGE) performed on the different gill tissues, as well as on isolated symbionts, resulted in a single gill symbiont amplification product, the sequence of which is identical to published data. These findings provide molecular evidence that one dominant phylotype is present in the morphologically different gill tissues. Nevertheless, the presence of other phylotypes cannot formally be excluded. The implications of this study are that the gill of L. aequizonata is a highly dynamic organ which lends itself to more detailed studies regarding the molecular and cellular processes underlying nutrient transfer, regulation of bacterial numbers and host–symbiont communication. Received: 1 September 1999 / Accepted: 1 February 2000     

Petroleum hydrocarbons in the marine bivalve Venus verrucosa: accumulation and cellular responses

Cellular and subcellular responses of the marine burrowing bivalve Venus verrucosa collected from the north-eastern coastline of Malta from January to June 1985, after exposure to petroleum hydrocarbons (PHC) were investigated. After long-term exposure to 100 gl^-1 of water-accomodated fractions (WAF) of crude oil, PHC were found to accumulate most rapidly in the digestive gland and then in the gills, with saturation levels being reached within 100 d of exposure in both cases. PHC accumulation, both in the mantle and muscle tissues, was more gradual and consistent throughout the whole exposure period. After 150 d of exposure, the digestive cells of the digestive gland were significantly reduced in height (atrophy) and exhibited reduced lysosomal membrane stability. After 144 h of exposure to higher concentration of PHC (820 and 420 gl^-1), several cytological effects were recorded, including an increase in cell volume and activity of gill mucocytes as well as in the number of haemocytes in gill blood sinuses. There was also evidence of damage to the epithelial lining of the foot, stomach and style sac and marked atrophy of the digestive cells of the digestive gland. The significance of such responses is discussed.     

High levels of genetic subdivision of marine and estuarine populations of the estuarine catfishCnidoglanis macrocephalus (Plotosidae) in southwestern Australia

The cobblerCnidoglanis macrocephalus (Valenciennes) is an endemic marine and estuarine catfish from southern Australia. Conflicting views on the degree of isolation of the estuarine populations underscore general questions about genetic divergence in coastal species. Although estuaries are widely recognized as ecologically important, little work has been done on their role in favouring genetic divergence. In order to estimate the extent of genetic subdivision among nearshore marine and estuarine populations, electrophoretic variation of enzymes was examined in seven marine and six estuarine populations of cobbler from sites spanning 1500 km along the southwest Australian coastline. Among all populations, the mean standardized variance in allelic frequencies (F _ST) for six polymorphic loci was 0.277, a high value comparable to those of other shallow-water teleosts whose life-history characteristics and habitat preferences restrict their dispersal capability. The pattern of genetic identities between populations showed divergence between west and south coast sites. Within these regional groups, however, there was substantial heterogeneity, much of which was associated with estuaries. Among all six estuarine sites, the averageF _ST was 0.333, 40% higher than the value of 0.237 for the marine sites. Low estimates of the genetically effective number of migrants suggest population subdivision between marine and estuarine environments and between similar habitat types. This study indicates the importance of habitat in affecting the connectedness of populations, even in apparently open marine systems.     

Prodissoconch morphology is environmentally modified in the brooding bivalve Lasaea subviridis

Embryonic shell sculpture of intertidal and laboratory brooded individuals of the direct developer Lasaea subviridis was studied using scanning electron microscopy. Intertidally brooded individuals develop a distinct prodissoconch I (PI) and prodissoconch II (PII) of unusual morphologies. The PI is relatively small (50–100 m in length), oval in outline, lacks a radial sculpture and is restricted to the initial pitted zone of the prejuvenile shell. A larger PII is formed (510–680 m in length) and it is dominated by pronouced commarginal striae but also includes a belt of radial ridges extending from the PI/PII boundary. Laboratory brooded individuals kept constantly submerged do not develop the pronounced commarginal striae characteristic of PII. This implies that PII formation in L. subviridis is not a direct result of the brooding habit, and can be modified by environmental factors, which possibly include low-tide exposure. There is no single feature of external prodissoconch morphology that unambiguously indicates a direct development mode combined with mantle cavity brood protection. General guidelines are presented to help recognize this developmental mode, based on prodissoconch morphology. In comparing prodissoconch morphologies of brooding bivalves, the habitats as well as the phylogenies of species should be considered, especially in comparisons of intertidal and subtidal species.     

Seasonal variation in some population parameters of Euchaeta species (copepoda: Calanoida)

Studies of the biology of oceanic copepods are few relative to those of coastal species. Females of the genus Euchaeta have spermatophores attached to the genital somite by the male and carry their broods of eggs attached to this somite, so defining the breeding season. Populations of E. norvegica in the fjordic environment of Loch Etive (collected between 1971 and 1974 and from 1978 to 1979), Scotland and in the marginal oceanic region of the Rockall Trough, northeastern Atlantic Ocean (collected between 1973 and 1976) and of a further eight species of Euchaeta in the Trough are examined. Seasonal changes in the incidence of egg masses and spermatophores attached to the females, sex ratio and population numbers are determined. Sampling errors in the estimation of these parameters are larger in the oceanic Rockall Trough. The Loch Etive populations of E. norvegica produce two generations per year with a proportion, variable between years, of the population producing a third generation. This species produces a single annual generation in the Rockall Trough as do E. acuta and E. pseudotonsa. Two generations per year are probably passed through by E. gracilis in the Trough while the large bathypelagic E. sarsi may produce a single generation in the autumn of every second year. The other bathypelagic species, E. barbata, E. scotti, E. abbreviata and E. longissima, breed continuously throughout the year and no estimate of their generation times is possible. Consequently, this approach to population analysis is useful where breeding is seasonal, but resolving generation time in bathypelagic crustaceans remains a problem.