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Coastal locations are highly influenced by input from freshwater river runoff, including sources of terrestrial carbon, which can be expected to modify the 14C reservoir age, or R (t), associated with marine water. In this Baltic Sea case study, pre-bomb museum collection mollusc shells of known calendar age, from 30 locations across a strategic salinity transect of the Baltic Sea, were analysed for 14C, δ13C and δ18O. R (t) was calculated for all 30 locations. Seven locations, of which six are within close proximity of the coast, were found to have relatively higher R (t) values, indicative of hard-water effects. Whenever possible, the Macoma genus of mollusc was selected from the museum collections, in order to exclude species specific reservoir age effects as much as possible. When the Macoma samples are exclusively considered, and samples from hard-water locations excluded, a statistically significant correlation between Macoma R (t) and average salinity is found, indicating a two end-member linear mixing model between 14Cmarine and 14Crunoff. A map of Baltic Sea Macoma aragonite R (t) for the late 19th and early 20th centuries is produced. Such a map can provide an estimate for contemporary Baltic Sea Macoma R (t), although one must exercise caution when applying such estimates back in time or to 14C dates obtained from different sample material. A statistically significant correlation is found between δ18Oaragonite and Macoma R (t), suggesting that δ18Oaragonite can be used to estimate Macoma palaeo-R (t), due to the δ18Oaragonite signal being dominated by the salinity gradient of the Baltic Sea. A slightly increased correlation can be expected when δ18Oaragonite is corrected for temperature fractionation effects. The results of this Baltic Sea case study, which show that R (t) is affected by hydrographic conditions and local carbon inputs, have important consequences for other coastal and estuarine locations, where R (t) is also likely to significantly vary on spatial and temporal bases.

Seasonality estimates based on stable isotope analyses of shellfish remains has been an important thrust of settlement pattern reconstruction, allowing researchers to place people on the landscape at points in space at different times of the year. In exposed coastal settings seasonality reconstructions are typically dependent on annual changes in water temperature. This paper has two goals. First, we continue development of a method for determining shellfish harvest seasonality in estuarine environments where annual salinity changes, not temperature, drive isotopic variation. Second, we contribute to settlement pattern studies by showing how small and large sites can be linked into a single system by examining different site types and shellfish species. Our case study focuses on the Late Prehistoric period of the San Francisco Peninsula, includes a large shellmound (CA-SMA-6) and an ephemeral camp (CA-SFR-171), and examines clam (Macoma spp.) and mussel (Mytilus spp.) harvesting. In this case, data support a fission-fusion settlement pattern, with periods of dispersal during late winter through early summer and aggregation in late summer through early winter.

Hypoxia is the depletion of dissolved oxygen below 2 mg O2/L. Relatively few studies on hypoxia and its effects on benthic macrofauna have been done in tropical marine ecosystems. This study describes the temporal response of the water column, sediments and macrofauna to seasonal hypoxia in a semi-enclosed bay (Cienfuegos, Caribbean Sea). The Calisito site was sampled monthly from June 2010 until February 2012, yielding 21 sampling times. At each sampling event water and sediment samples were collected for measuring the abiotic variables (temperature, salinity, dissolved oxygen, nutrients, redox potential discontinuity, silt/clay and organic matter content) and macrofauna (abundance and species richness). Temperature and surface salinity followed a typical temporal pattern during the summer/rainy and the winter/dry periods. Salinity stratification occurred in the rainy period, lasting three months in 2010 and six months in 2011. The bottom water dissolved oxygen indicated hypoxic and anoxic events during the wet periods of 2010 and 2011 associated with salinity stratification, low hydrodynamics and oxidation of the accumulated organic matter. Over the study period, 817 individuals were collected and identified. Polychaetes were the dominant group in terms of abundance (57 % of total) followed by mollusks (41%). Hypoxia (and occasionally anoxia) caused strong deleterious effects on the abundance and species richness of macrofaunal communities in the study site. The most abundant polychaetes were opportunistic species with high tolerance to hypoxic conditions: Prionospio steenstrupi, Polydora sp.and Paraprionospio pinnata. Most of them colonized relatively fast once hypoxia ended. Persistent species such as Caecum pulchellum and Parvanachis obesa were present during hypoxia with fluctuating densities and apparently recover to higher abundances when normoxic conditions are re-established. Macoma tenta and Tellina consobrina colonized approximately 1-2 months later than the first polychaete peak during normoxia. Probably, the deleterious effects of hypoxia on the macrofauna were intensified by negative interspecific relationships such as competition by suitable space and predation. The recolonization of macrofauna depended possibly on local transport by currents within the bay because the connection with the Caribbean Sea is relatively limited. In summary, seasonal hypoxia in Cienfuegos Bay influences the water and sediment geochemistry and reduces both the abundance and diversity of macrofauna. ResumenLa hipoxia es la disminución del oxígeno disuelto por debajo de 2 mg O2/L. Relativamente pocos estudios sobre la hipoxia y sus efectos en la macrofauna bentónica han sido realizados en ecosistemas marinos tropicales. Este estudio describe la respuesta temporal de la columna de agua, los sedimentos y la macrofauna a la hipoxia estacional en una bahía semicerrada en Cienfuegos, Mar Caribe. La estación de Calisito fue muestreada mensualmente de Junio 2010 a Febrero 2012, resultando en 21 eventos de muestreo. En cada oportunidad se recolectaron muestras de agua y sedimento para la medición de las variables abióticas (temperatura, salinidad, oxígeno disuelto, discontinuidad del potencial redox, contenido de limo/arcilla y materia orgánica) y macrofauna (abundancia y riqueza de especies). La temperatura y la salinidad superficial presentaron un patrón temporal típico durante los periodos de verano/ lluvia e invierno/seca. Ocurrió estratificación por salinidad en el periodo lluvioso, que duró tres meses en 2010 y seis en 2011. El oxígeno disuelto de fondo indicó eventos de hipoxia y anoxia durante los periodos lluviosos de 2010 y 2011 asociados con la estratificación de la columna de agua, la hidrodinámica lenta y la oxidación de la materia orgánica acumulada. Esto causó efectos deletéreos en la abundancia y diversidad de la macrofauna. Los poliquetos más abundantes fueron especies oportunistas con alta tolerancia a condiciones hipóxicas: Prionospio steenstrupi, Polydora sp. y Paraprionospio pinnata. La mayoría de estos colonizaron relativamente rápido una vez que terminó la hipoxia. Especies persistentes como Caecum pulchellum y Parvanachis obesa estuvieron presentes durante la hipoxia con densidades fluctuantes y aparentemente recuperaron sus abundancias cuando las condiciones normóxicas fueron re-establecidas. Macoma tenta y Tellina consobrina colonizaron 1-2 meses después que el primer pico de poliquetos durante la normoxia.Probablemente, los efectos deletéreos de la hipoxia en la macrofauna fueron intensificados por relaciones interespecíficas negativas como la competencia por espacio y la depredación. La recolonización de la macrofauna dependió probablemente del transporte local por las corrientes dentro de la bahía pues la conexión con el mar Caribe es relativamente limitada. En resumen, la hipoxia estacional en la bahía de Cienfuegos influencia la geoquímica del agua y los sedimentos y reduce la abundancia y diversidad de la macrofauna.

The chapter highlights the comprehensive account of congener profiles of polycyclic aromatic hydrocarbons (PAHs) in intertidal bivalve mollusks (Meretrix meretrix, Macoma birmanica, and Sanguilonaria (Soletellina) acuminata) of Indian Sundarban mangrove wetland. The main aim of this work was to use the bivalves as bioindicators of the contamination of the 16 USEPA PAH. The PAH profile in bivalves is largely dominated by a petrogenic fingerprint, with over-imposition of pyrolytic PAH sources, as evidenced by diagnostic molecular ratios. Bioaccumulation factors (BAF) of individual compounds from the sediments were calculated, and it reveals overall higher values in the visceral mass of the bivalves. S acuminata showed significantly higher levels of PAHs, especially the high-molecular-weight (HMW) PAHs, compared to the other two species as a sensitive indicator of trace organic stress in future monitoring programmes.

Behavior has a predictive power that is often underutilized as a tool for signaling ecological change. The burrowing behavior of the deposit feeding bivalve Macoma balthica reflects a typical food-safety trade-off. The choice to live close to the sediment surface comes at a risk of predation and is a decision made when predation danger, food intake rates or future fitness prospects are low. In parts of the Dutch Wadden Sea, Macoma populations declined by 90% in the late 1990s, in parallel with large-scale mechanical cockle-dredging activities. During this decline, the burrowing depth of Macoma became shallow and was correlated with the population decline in the following year, indicating that it forecasted population change. Recently, there has been a series of large recruitment events in Macoma. According to the food-safety trade-off, we expected that Macoma should now live deeper, and have a higher body condition in association with this change in depth of living. Indeed, we observed that Macoma now lives deeper and that living depth in a given year forecasted population growth to the next year, especially in individuals larger than 14 mm. As living depth and body condition were strongly correlated in individuals larger than 14 mm, larger Macoma could be living deeper to protect their reproductive assets. Our results confirmed that burrowing depth signals impending population change and, together with body condition, can provide an early warning signal of ecological change. We suggest that population recovery is being driven by improved intertidal habitat quality in the Dutch Wadden Sea, rather than by the proposed climate-change related effects. This shift in ecosystem state is suggested to include the recovery of diatom habitat in the top layer of the sediment after cockle-dredging ended.

Despite the great importance of the siphons for infaunal bivalves, only a few studies have examined their tissues using histology techniques or scanning electron microscopy. In the present study, the siphons of Tellina lineata Turton, 1819 and Macoma biota Arruda & Domaneschi, 2005 were investigated. The siphon walls are composed by a series of muscle sheets of longitudinal ("L"), circular ("C") and radial ("R") fibers, with a clear pattern common to both species: there is a main median longitudinal layer (Lm), and two peripheral circular layers, one inner (Ci) and one outer (Co), near the epithelia. A median circular layer (Cm) separates an internal (Lmi) from an outer (Lmo) median longitudinal layer. Further, the Co is split by a thin outer longitudinal layer (Lo), forming Coi and Coo layers, the former being obliquely oriented. Thin radial fibers (R) delimit clear packages of Lmi and Lmo fibers. In each siphon, there are six longitudinal nerve cords, running within the Lmi layer, adjacent to the Cm. The inhalant and exhalant siphons of M. biota are very similar in structure, but the Lmo of the exhalant siphon is almost twice as thick as its Lmi, while in the inhalant siphon these layers have similar thicknesses; the Coi is very thick, especially in the exhalant siphon. The inhalant siphon of T. lineata is very similar to that of M. biota, differing only with respect to the thickness of the Coi, which in the former species is not as well developed as in the latter. The Lmo of the exhalant siphon of T. lineata is by far the most developed layer, with the Lmi represented only by uniseriate small cells; in the vicinities of the nerve cords, the Cm is split in two layers. The epithelia of both siphons of M. biota and T. lineata bear ciliated receptors, which were difficult to observe as they were frequently covered with mucus. It was possible to observe that cilia are present in both species, differing in length and in the number per receptor between the inhalant and exhalant siphons, and between the species. Detailed comparisons among the siphons of M. biota and T. lineata and other bivalve species are very difficult, because of at least two reasons. First, each investigator has used different methods to prepare and observe the siphons through histological sections; and second, different nomenclatural schemes are used to describe the musculature of the siphons, causing confusion when the same layers are compared among different species. In order to unify the nomenclature of tissue layers of the bivalve siphons, we now propose a scheme to name these layers based on topological homology.

Ecological and taxonomic study of the mollusk-rich fauna of the Golfe d'Arguin, North Mauritania, investigates the various environmental influences affecting this tropical shelf. The upwelling of nutrient-rich waters leads to a highly productive environment under tropical conditions. The resulting mixed carbonate-siliciclastic sediment contains a large portion of calcareous components produced by heterotrophic organisms—e.g., mollusks, foraminifers, worms, barnacles—that are reworked on the open shelf. On the basis of mollusk assemblages, six taphocoenoses are defined, all being characterized by a mixed fauna of tropical (e.g., Tellina densestriata), subtropical (e.g., Macoma cumana) and temperate (e.g., Spisula subtruncata) species. Differences between the assemblages are related to the medium—grain size ranging from mud to gravel—that results from local hydrodynamic conditions and water depth. Among carbonate grains, Donax burnupi shells are very abundant in the swell-exposed, northern part of ...

Numerous shells of bivalves, which study attributed to 37 species, were collected from samples of ground taken with a Van Veen dredge sampler at the northeast coast of Sakhalin Island at depths of about 10–60 m in June–October 2003–2009. Tellina lutea, Mactromeris polynyma, Serripes groenlandicus, Macoma spp., Liocyma fluctuosum, and Siliqua alta occurred most often in the samples. The assessment of growth in 18 bivalve species revealed common patterns of age variations that were irrespective of their way of life, maximum body size, and maximum age. The greatest annual increment in shell length was observed in the first 4 years of life, and the asymptotic size of the mollusks was reached with a 10-fold decrease in the annual growth rate. The life span of the mollusks varied from 2 up to 41 years and sometimes significantly differed from that of the studied species in other areas of their range, clearly displaying the ecological conditionality of this parameter. Species with maximum ages of less than 10 years were the most numerous in the bivalve community of the region. The results of the research are discussed in relation to the nature of the environment at the northeast coast of Sakhalin Island.

The first occurrence and presence of the bivalve Macoma balthica in the Pleistocene shallow marine sediments (Crags) of East Anglia has long been used to characterise the youngest “Weybourne” or “Wroxham” Crag Formation of Early to early Middle Pleistocene age. A review of recent work on the morphological variation and genetic lineages of living populations of the species and of work on fossil discoveries in East Greenland suggests that the current palaeontological species definition is too narrow, that the species arrived in the Atlantic about 2.4 Ma and much earlier than is envisaged from the studies of the North Sea Basin, and that there were several trans-Arctic migrations of the species during the Pleistocene. This paper reviews these findings and concludes that: (i) detailed morphometric analyses of fossil Macoma species from the Crag need to be undertaken and the species definitions comprehensively reviewed to take account of the morphological and genetic variation seen in modern populations; (ii) detailed morphometric analysis of modern M. balthica should be undertaken and tied to the different genetic lineages, and (iii) the current palaeontological definition of Macoma balthica should not be relied on as the basis for a First Appearance Datum (FAD) in the Southern North Sea basin.

Recent reductions in sea ice cover and thickness in the Arctic will lead to changes in food supplies for benthic consumers. We experimentally assessed responses of two common Arctic macrobenthic species, Macoma balthica (Bivalvia) and Monoporeia affinis (Crustacea) from Kotzebue Sound (Alaska, USA) to varying food materials (phytoplankton and ice algae). Phytoplankton and 13C-labelled ice algae were added separately or together to pre-sieved sediment cores containing known numbers of Macoma or Monoporeia. After 8 d, sediments and animals were analyzed for bulk C/N contents, isotopic signatures, fatty acid compositions and compound-specific δ13C values. Our results showed that the two species responded differently to varying food materials. Loss of characteristic fatty acids and changes in δ13C values in the surrounding sediments indicate that Macoma preferentially consumed ice algae compared to phytoplankton, while Monoporeia non-selectively ingested both food materials. Feeding behaviors also differed between the two species: Macoma fed primarily on material from the sediment surface, and did not appear to mix material to depth, while Monoporeia mixed some fresh food material from the surface down to subsurface sediments (1–2 cm). Analysis of animal biomass showed more 13C-labelled organic carbon from ice algae was assimilated by individual Macoma compared to individual Monoporeia. Moreover, δ13C values of fatty acids in Macoma were much higher than those in Monoporeia, suggesting that Macoma directly assimilates fatty acids from their food source while Monoporeia might de novo biosynthesize fatty acids in their biomass. Additionally, both species had higher fatty acid contents and proportion of polyunsaturated components when they fed on ice algae compared to phytoplankton, implying that ice algae may be a better food than phytoplankton for Arctic benthos. Finally, analysis of bacteria-specific fatty acids in sediments (from both types of animal cores) showed that the δ13C values were enriched up to 400‰ relative to natural background values (−25% to −30‰), suggesting that organic carbon from fresh ice algae could be rapidly incorporated into bacteria biomass. The collective results of this experimental study suggest that the changes in food supplies due to reduction of ice coverage will have a greater impact on those benthic organisms that prefer ice algae to phytoplankton. This may lead to the potential changes in community structure favoring those organisms that can best utilize the fatty acids contained in both phytoplankton and ice algae.

The paper presents the first comprehensive account of congener profiles of polycyclic aromatic hydrocarbons (PAHs) in intertidal bivalve mollusks [Meretrix meretrix, Macoma birmanica, and Sanguilonaria (Soletellina) acuminata] of Sunderban mangrove wetland (India). The main aim of this work was to use the bivalves as bioindicators of the contamination of the 16 USEPA PAH. The PAH profile in bivalves is largely dominated by a petrogenic fingerprint, with over-imposition of pyrolytic PAH sources, as evidenced by diagnostic molecular ratios. Bioaccumulation factors (BAF) of individual compounds from the sediments were calculated, and it reveals overall higher values in the visceral mass of the bivalves. S acuminata showed significantly higher levels of PAHs, especially the high molecular weight (HMW) PAHs, compared to the other two species as a sensitive indicator of trace organic stress in future monitoring programs.

We studied the effect of crustacean predators on native/non-native Macoma balthica bivalves in aquarium experiments. North Sea M. balthica (NS Macoma) were recently observed in the southern Baltic Sea. They differ genetically and in terms of morphology, behaviour and evolution- ary history from Baltic Sea M. balthica (BS Macoma), and this may affect predation pressure and community structure. We hypothesised that predators consume more of the prey they co-exist with. NS Macoma and BS Macoma were exposed to crustacean predators common in the North Sea (Carci- nus maenas and Crangon crangon) and in the Baltic Sea (C. crangon and Saduria entomon). Contrary to our hypotheses, the North Sea predators ate more BS Macoma, and S. entomon ate more NS Macoma. The crush-limited C. maenas preyed more on globular BS Macoma, whereas S. entomon, which do not crush but pry open the bivalve shell, ate more NS Macoma, which have a lighter (thus probably thinner) shell than BS Macoma. When NS and BS Macoma were offered together, BS Cran- gon ate more NS Macoma. We also studied BS Crangon consumption of M. balthica to assess whether sizes offered fall within the size spectrum that C. crangon can eat. Small (20 to 40 mm long), medium (40 to 50 mm) and large (50 to 60 mm) C. crangon especially ate small M. balthica. Differences in shape, size and meat/shell weight ratio between the BS and NS Macoma partly explained the differences in the susceptibility to predation by native and non-native predators.

Distribution and abundance of the macrobenthic fauna was studied in the intertidal and subtidal soft-bottoms of the Ensenada de San Simón (north-western Spain). Depth and grain size were the most important factors in determining patterns of distribution of the macrozoobenthos. A reduced Macoma community was present in the inner area of the inlet structured in two facies subdivision in the community that does not affect its qualitative composition: the first one was in intertidal areas associated with meadows of the sea grasses Zostera noltii and Z. marina and defined by the dominance of Hydrobia ulvae; the second facies in intertidal sediments was characterized by the dominance of Tubificidae and nematodes and Pseudopolydora paucibranchiata. The poverty of these bottoms in number of species can be due to salinity fluctuations and human activities developed in the area. Subtidal bottoms were characterized by a Syndosmya(=Abra)alba community. Two facies were detected: one facies located in shallow subtidal bottoms with H. ulvae and Thyasira flexuosa as characteristic species, and a second facies in deeper areas, characterized by the polychaetes Ampharete finmarchica, P. paucibranchiata, Aphelochaeta marioni, Melinna palmata and Paradoneis lyra.

A long sequence of open-coast tidal flat deposits with a restricted, mainly boreal fauna, comprises the Tapes and Mactra Zones. Characteristic species are Venerupis aurea, Spisula arcuata, Arctic islandica and Lentidium complanatum. This stage is provisionally correlated with the Upper Pliocene of the North Sea Basin. During the time of deposition of the Serripes Zone the basin apparently deepened and the fauna greatly diversified with immigration of arctic elements. It went back finally to beach-deposit conditions. The characteristic species —Serripes groenlandicus, Macoma praetenius, and Beta borealis — are also characteristic of the Early Pleistocene of the North Sea Basin with which the Serripes Zone is provisionally correlated. The climate is interpreted as having been colder than that of the Tapes and Mactra zones. A preliminary list of the Mollusca is given.

The data from the expedition of the program RUSALCA conducted in 2004 showed unexpectedly high quantitative indices of macrobenthos in the southeastern Chukchi Sea. Extensive areas of the bottom north- west of the Bering Strait were dominated by the bivalve Macoma calcarea . The greatest biomass of benthos in Macoma -dominated areas was 4232 g/m 2 with an average of 1382 g/m 2 for the investigated region. Such a high biomass of soft-bottom communities, which is extremely uncommon even in the temperate regions of the oceans, is reported for the Arctic for the first time. The long-term existence (more than 70 years) of highly pro- ductive benthic communities dominated by Macoma calcarea in one and the same area of the Chukchi Sea can most likely be attributed to gyres, which constantly arise in the region northwest of the Bering Strait. These cyclonic gyres carry nutrient-rich bottom water to the surface and hinder larval transport away from mother populations. They also keep and concentrate major food sources of benthos (live and dead phyto- and zooplank- ton and fecal pellets) over the benthic community locations. Most likely, a significant proportion of the primary production in the southeastern Chukchi Sea is used by benthos within the investigated Macoma community. Findings of three relatively large warm-water Pacific species near Point Hope in the Chukchi Sea are probably indicative of the progressive climate warming during the last century.

A main molecular subdivision in the circumpolar Macoma balthica complex has been described between Atlantic and Pacific taxa. In NE Europe, the clams of the White and Barents Seas, however, show deviant genetic structures. Using allozyme and mitochondrial DNA data, we explore the hypothesis that these deviations result from hybridization between an Atlantic (M. b. rubra) and an invading Pacific (M. b. balthica) lineage. A practically pure Atlantic Macoma extends from France north to the Varanger Peninsula (NE Norway), whereas populations farther east have genetic compositions intermediate between true Atlantic and true Pacific. Admixture estimates range from 32 to 90% Pacific contribution, with a notable deviation in a nearly pure Atlantic outpost in the Mezen Bay (NE White Sea). The pattern of variation is not one of a simple collinear mixing however. Different characters exhibit different degrees of introgression, and the relative introgression varies regionally. Yet, there are practically no interlocus genotypic disequilibria between the diverged loci, which brings out the White Sea-Barents Sea M. balthica as the best-documented marine animal hybrid swarms so far, arisen through amalgamation of genomes previously isolated since pre-Pleistocene times. On top of the main admixture pattern, strong geographical structuring is also seen in characters unrelated to the principal systematic distinction. The persistence of the regional patterns indicates restricted gene flow at the present time, despite the high dispersal potential of the species. The causes of this structuring could be in a complex history of colonization events and features of local hydrography enhancing isolation and divergence of populations.

The lady crab (Ovalipes ocellatus) is one of the most common native species of swimming crab (Portunidae) of the Atlantic Coast of North America but most populations occur south of Cape Cod, Massachusetts. There is a disjunct population in Northumberland Strait (southern Gulf of St Lawrence), which was the focus of this study. Adult lady crabs were collected by trawling in water >4 m deep from May to October 1999 to 2005 to determine abundance, distribution, and diet. Lady crab occurred only in a small area (about 2,500 km2) in the central part of Northumberland Strait where bottom water temperature was >18°C during summer, and the substrate was mainly sand or sandy gravel. Male lady crab attained a maximum carapace width (CW) of 112 mm compared to 92 mm CW for females. The summer and autumn diet consisted mainly of infauna. The principal prey (each >5% of diet by weight) were: small bivalve molluscs (primarily Atlantic razor clam Siliqua costata and Macoma sp.; 43%), small rock crab (Cancer irroratus; 13%), polychaetes (11%), fish remains (9%), and small lady crab (9%). All stomachs collected during May (water temperature ≤10°C) were empty. There was little evidence of any difference in feeding intensity between 0700 h and 1900 h.

Ecosystem engineers can influence community and ecosystem dynamics by controlling resources, modifying the flow of energy or biomass, or changing physical characteristics of the habitat. Invasive hybrid cordgrass ( Spartina alterniflora × Spartina foliosa ) is an ecosystem engineer in salt marshes in San Francisco Bay, California, U.S.A. that raises intertidal elevations and may be either increasing C 4 plant carbon input into food webs or tying up carbon in a form that is not usable by consumers. A manipulative experiment compared abundance, growth, and stable isotope (δ 13 C and δ 15 N) composition of the clam Macoma petalum (= M. balthica ) among native marsh, hybrid Spartina , and mudflats in central San Francisco Bay. We found higher densities (individuals m −2 ) of M. petalum on mudflats compared to either native or hybrid Spartina ( p Macoma petalum shell growth was significantly greater in mudflats than in either vegetation type in 2002 ( p = 0.005) but not 2003. Differences in shell growth between native and hybrid Spartina were not significant. Stable isotope results showed differences between habitats in δ 13 C but not δ 15 N. Carbon signatures of M. petalum placed in Spartina were much more depleted than the isotopic signature of Spartina . Neither native nor hybrid Spartina appears to be a significant carbon source for M. petalum in San Francisco Bay, and we found no evidence that hybrid Spartina contributes carbon to M. petalum beyond what is provided by S. foliosa , despite the hybrid's much greater biomass. Our results show that loss of mudflat habitat, rather than increased input of C 4 carbon, is the greatest effect of the invasion of hybrid Spartina on M. petalum .

Recent studies have documented strong, top-down predation effects of sea ducks on mussel populations in rocky intertidal communities. However, the impact of these gregarious preda- tors in soft-bottom communities has been largely unexplored. We evaluated effects of predation by wintering surf scoters Melanitta perspicillata and white-winged scoters M. fusca on clam populations in soft-bottom intertidal habitats of the Strait of Georgia, British Columbia. Specifically, we docu- mented spatial and temporal variation in clam density (clams m -2 ), scoter diet composition, and the consequences of scoter predation on clam abundance. Of the 3 most numerous clams, Manila clams Venerupis philippinarum and varnish clams Nuttallia obscurata were the primary prey items of both scoter species, while clams of the genus Macoma were rarely consumed by scoters. Between scoter arrival in the fall and departure in the spring, Manila clams decreased in density at most sample sites, while varnish clam densities did not change or declined slightly. Our estimates of numbers of clams consumed by scoters accounted for most of the observed declines in combined abundance of Manila and varnish clams, despite the presence of numerous other vertebrate and invertebrate species known to consume clams. For Macoma spp., we detected an over-winter increase in density, presum- ably due to growth of clams too small to be retained by our sieve (

We report the application of a recently developed molecular method, single step nested multiplex PCR (SSVM-PCR) assay and microscopy to identify and investigate temporal patterns of bivalve larvae in a Danish estuary, Isefjord. All samples were collected during the SUSTAINEX program from June to November 2001. Using the molecular assay, larvae could be categorized into six groups: the blue mussel, Mytilus edulis, Ensis spp., species of the Myoidae superfamily (Mya spp.), the common cockle (Cardiidae family), members of the Abra and Macoma genera of the Tellinoidae superfamily and members of the surf clam genera, Spisula spp. A seventh group was composed of unknown larvae. Greater resolution was possible by microscopy, but only for relatively large and intact individuals (>150-200 μm). The molecular approach was capable of differentiating between larvae regardless of shell size. Where it was possible to directly compare identifications based on both methods, concordance was high for M. edulis, Macoma balthica/Abra alba and E. americanus, whereas identification of Myoidae spp. and Cardiids was less consistent. Over the course of the study, two patterns of larval occurrence were observed. Larvae from species known to exhibit a protracted annual spawning period (M. edulis, Myoidae spp., Mysella bidentata and Cardiids) were present in the water column throughout the sampling period, whereas larvae of Abra alba, Barnea candida, E. americanus, Macoma balthica, Musculus marmoratus Scrobicularia plana and Tapes pullastra appeared at clearly defined periods.

Colonization of formulated artificial substrate was investigated in a tropical estuarine ecosystem for six months to determine possible response of benthic macroinvertebrates to disturbance arising from substrate alteration. Five different formulated substrates which served as content for ten cages were used for this experiment. A total of 406 individuals comprising 18 macroinvertebrate species colonized the cages. In overall, cage 1 which is the control recorded the highest number of individuals and average number (8) of colonizing species at a time, while the least number (4) was observed in cage 3. Cages 2, 4, and 5 recorded average of five species each. In terms of number of individuals, cage 1 which is the control, had the highest number (153), followed by cage 2 which recorded 122 individuals. Cages 3, 4 and 5, recorded 41, 61 and 80 individuals respectively. Analysis of temporal variability in the number of species observed in the experimental cages shows that there was a progressive decrease in the number of colonizing species after the second month. Colonizing macroinvertebrate groups were not different from the known resident species in the estuarine lagoon. However, pattern of variation in the number of individuals of colonizing species were remarkably different. The assemblage recorded in this experiment was defined by the populations of two bivalve mollusc; Macoma cumana and Tellina nymphalis , two gastropod mollusc; Pachymelina aurita and Tympanotonus fuscatus and two crustacean arthropods; Clibanarius africanus and Penaeus notialis . Among the FFGs recorded, collector-filterers dominated in number of individuals and species. Nutrient concentrations of formulated substrate had strong effects on number of colonizing individuals and species.

Nutrient and organic loading associated with escalating human activities increases biological oxygen demand from microbial decomposition. In the Neuse River estuary, North Carolina, recurrent nuisance algal blooms, bottom-water hypoxic events, and fish kills during summers of the 1990s suggest that uncapped nutrient loading may have increased the frequency, duration, and/or spatial scope of important biological effects of hypoxia during summer, when persistent water column stratification can occur and microbial metabolism is greatest. We test the hypothesis that the severity of benthic biological effects of hypoxia in this estuary has increased over a 30-year period of dramatic human population growth in eastern North Carolina by comparing survival over summer of the benthic bivalve Macoma spp. between historical (1968–1970) and recent (1997–1998) years. Macoma is a demonstrated indicator of oxygen availability, the benthic biomass dominant in the Neuse and other temperate estuaries and the major prey link to higher trophic levels. All three historical summers exhibited patterns of collapse in Macoma populations indistinguishable from the recent summer of severe hypoxia (1997) but distinct from the modest changes documented during the mildly hypoxic summer of 1998. The only Macoma to survive any severely hypoxic summer were those in shallows where oxygen could be renewed by surface mixing. Thus, the biological effects of hypoxia observed in the Neuse River estuary in the late 1990s appear no more severe than 30 years before. Historic rates of organic loading to the Neuse River estuary may have been sufficient to induce widespread and intense hypoxia beneath the surface mixed layer, implying that even if algal blooms are diminished through nutrient reductions, the severity of biological effects of bottom-water hypoxia may not change detectably.

The 87Sr/86Sr isotope ratio in subfossil carbonate mollusc shells from raised-beach sediments is used as a palaeosalinity indicator for the Baltic Sea. The main purpose is to demonstrate the applicability of the method in a formerly glaciated area, using 87Sr/86Sr ratios found in modern shells (Mytilus edulis, Macoma baltica, and Radix balthica) to elucidate the parameters and conditions to be used for palaeosalinity determinations based on subfossil shells. Proxy salinity data are calculated assuming that the Sr concentration in Baltic Sea water is controlled by a two-component, conservative mixing between seawater and river water. Based on replicate determinations of the 87Sr/86Sr ratio in modern shells, proxy salinity data can be quantified with a relative precision of better than ± 5% for salinities up to ∼ 10‰. Comparison with measured, present-day salinities indicates that the accuracy is generally better than ± 5% for the proxy data. With this precision and accuracy, the present-day salinity differences between the major Baltic sub-basins (Bothnian Bay, Bothnian Sea, and Baltic Proper) can be detected. Palaeosalinities were determined using subfossil shells (M. edulis and M. baltica) with calibrated 14C ages of 6770–3070 cal BP. The shell deposits occur from 65 m a.s.l. down to the present-day sea level. The palaeosalinities determined for the Bothnian Bay (4.8–10.3‰) and the Bothnian Sea (7.3–10.3‰) are in good agreement with earlier estimates of the maximum Littorina Sea stage palaeosalinity in these sub-basins (8–10‰). At one locality, two different shell species from the same shell deposit were dated (M. edulis and M. baltica). The shells differed in age by 460 ± 70 14C years, a possible indication that the ecological conditions at a locality may have been favourable to different faunas at different times.

Patterns of nine intracellular free amino acids (FAA), which are utilized as organic osmolytes for salinity-induced cell volume regulation in marine osmoconformers, were compared in five Macoma balthica populations and seven Mytilus spp. populations along their European distribution. Three types of FAA patterns were classified within both taxa: a northern Baltic type, a southern Baltic type and an Atlantic/Mediterranean type which mainly differ regarding the share of alanine and taurine. Differences are discussed in relation to habitat salinity and population genetics. Along a salinity gradient, the total size of the intracellular FAA pool did not differ between sympatric M. balthica and Mytilus spp., and was significantly correlated with habitat osmolality in a range from 70 to 600 mmol kg−1 H2O (oligohaline to mesohaline) in both bivalves. In M. balthica, this correlation was mainly based on significant correlations of alanine (15–100 mmol kg−1 DW), glycine (30–100 mmol kg−1 DW) and taurine (0–70 mmol kg−1 DW) with habitat osmolality. In Mytilus spp., only glycine (25–100 mmol kg−1 DW) and taurine (4–180 mmol kg−1 DW) were significantly correlated with habitat osmolality. The concentration of alanine was three times lower in Mytilus spp. than in M. balthica and did not correlate with habitat osmolality. Within a habitat osmolality range from 600 to 1,100 mmol kg−1 H2O (mesohaline to marine) the concentration of FAA remained constant in both taxa. It is suggested that under marine conditions additional organic osmolytes must become more important for cell volume regulation in Macoma and Mytilus.

Based on a bottom trawl survey, it is shown that in Karaginskii and Olyutorskii bays of the western part of the Bering Sea, Lycodes raridens is a typically elittoral species, since it was recorded at depths of 52–120 m, and the majority of individuals of this species were found in the range of depths 50–100 m. The maximum concentrations of Lycodes raridens were 139 ind./km2 or 98 kg/km2, whereas the average values of the distribution density of this species were 7.8 ind.km2 or 5.4 kg/km2. In Karaginskii and Olyutorskii bays, Lycodes raridens is represented by individuals with a length of 16–86 cm, weight of 34–4510 g, at the age of 1+ to 7+, although fish with a length of 30–50 cm, weight of 100–1000 g, and age of 3+–4+ dominate. The data obtained suggest that in the autumn or autumn-winter period, the study species spawns in the western part of the Bering Sea. The dominant food of Lycodes raridens here are amphipods, polychaetes, bivalves, and decapods; by occurrence and the relative content in the stomach (% of weight), amphipods (as a rule, Melita spp.) and bivalves (Yoldia spp. and Macoma sp.) dominate. Lycodes raridens is a benthophage in which, at the early stage of the life cycle, gammarids prevail in food; with growth, its diet becomes more diverse due to bivalves, polychaetes, and decapods.

Large areas of the bottom sediments of the Baltic Sea are temporarily or permanently anoxic. These sediments are also an important sink for a variety of contaminants. Reoxygenation of bottom waters allows recolonisation by benthic infauna, which may have important implications for the fate of buried contaminants. This study used tracers to experimentally examine the role of bioturbation by benthic infauna in transporting sediment-associated contaminants in the Baltic Sea. Three different tracer methods were used in two experiments, using three key Baltic macrofaunal species: the amphipod crustacean Monoporeia affinis ; the Baltic clam Macoma baltica ; and the priapulid worm Halicryptus spinulosus . In the first experiment, a reoxygenation–recolonisation scenario was recreated in the laboratory, using hypoxic sediment cores collected in the field, to determine if there was remobilisation of buried 137 Cs from the Chernobyl nuclear accident in 1986. The potential for the infauna to bury newly settled surface contamination was also investigated, using a fluorescent particle tracer. In the second experiment, artificially-created radiolabelled tracer layers ( 14 C and 51 Cr) were used to quantify both upward and downward movements of organic matter and sediment-associated contaminants by bioturbation. In both experiments there were clear visual differences between the sediment effects of the three species. Halicryptus spinulosus buried deepest into the sediment, creating a network of burrows, Monoporeia affinis burrowed actively in the upper few centimeters of the sediment, and Macoma baltica was quite stationary, but appeared to filter- and deposit feed at the sediment surface. Mixing depths in the hypoxic sediment varied from 4.0 ± 3.5 cm for M. baltica to 7.8 ± 2.1 cm for H. spinulosus . Biodiffusion rates ( D b ) were similar for all treatments but biotransport rates ( r ) were significantly different between treatments, mainly due to a high r value for H. spinulosus . In the experiment with radiolabelled tracer layers, 51 Cr was transported more than 14 C, and tracer originally at the surface transported more than tracer buried 4 cm below the surface. There was also transport of all tracers in treatments without added macrofauna. The most likely explanation is bioturbation by the meiofauna that were undoubtedly present in both experiments. Bioturbation by macrofauna both buries surface contaminants and remobilises those that are buried, but the effects are small and on a similar scale to transport caused by meiofauna. In addition, 137 Cs profiles at the hypoxic site indicated that resuspension and redeposition of sediment by physical processes had occurred, and also showed that contaminants from the last 40 years were still present in the top 5–10 cm of the sediment, well within active mixing depths. At this site, as at many others in the Baltic, physical processes are likely to be far more important than biological processes in the redistribution of contaminants on a decadal timescale.

Blue crabs (Callinectes sapidus) and deposit- and suspension-feeding bivalves (Macoma balthicaand M. mitchelli) play important roles in the food web of Chesapeake Bay and may serve as indicators of ecological health, particularly in small subestuaries where conditions may be strongly linked to watershed and local factors. We sampled 19 subestuaries of Chesapeake Bay to test hypothesized relationships of blue crabs and bivalves with salinity (a regional indicator), watershed- and local-scale land use, and local habitat and water quality. We divided the subestuaries into five land-use categories: forested, developed, agricultural, mixed agricultural and mixed-developed. We measured water quality, sediment class, physical habitat and adjacent land use/land cover at each of six stations within each subestuary. Fyke nets were employed to estimate blue crab abundance and size-structure, while cores were used to estimate bivalve densities and biomass. Classification and regression tree (CART) analysis indicated that 51% of the variance in blue crab abundance was explained by salinity, watershed land use and shoreline marsh habitat. Crab abundance was greatest at salinities >16 ppt, but in lower salinities crabs were most abundant along marsh shorelines in forested and mixed land-use watersheds. Juvenile crabs

Variation in habitat quality and resource availability can affect the distribution and growth of animals. Thin-shelled clams dominate many benthic communities in Chesapeake Bay, both in numbers and in biomass, and they can comprise up to 50% of the blue crab (Callinectes sapidus) diet. Our objective was to determine which habitats were optimal for juvenile crab growth and how growth related to food availability. We experimentally examined benthic infaunal food availability (primarily bivalves) and concurrent growth of juvenile blue crabs at 30–40 sites along 50 km of the York River during fall 2000 and spring 2001. Each year, 4–10 replicate sites along the York River were established in each of five habitats: (1) Seagrass, (2) Mud at the river mouth, (3) Sand at the river mouth, (4) Mud upriver, and (5) Sand upriver. Food availability inside and outside of 0.43-m2 crab growth cages was examined, along with crab growth after 3–6 months inside cages. In both years, after 3–6 months, the Baltic clam, Macoma balthica, was abundant inside and outside the cages, whereas the soft-shell clam, Mya arenaria, was only abundant inside cages. Densities of Macoma were greatest in upriver mud and sand, while those of Mya were greatest in upriver sand. Crab growth was significantly greater in spring-summer than fall-winter and was significantly higher in upriver mud and sand, where clam densities were highest, than at the river mouth. The upriver region was near the turbidity maximum, which may enhance pelagic and benthic productivity and thereby provide more food for clams and therefore for blue crabs. Crab growth in seagrass was intermediate between that upriver and at the mouth, suggesting that upriver, unvegetated, subtidal habitats adjacent to salt marshes serve as valuable nursery habitats rivaling seagrass beds.

Question: Do clams from populations not exposed to a predator retain the ability to respond to that predator??Motivation: If maintaining the potential for phenotypic plasticity involves a significant cost. plasticity should be selected against in constant environments.Background: Clams of the species Macoma balthica (a burrowing bivalve) respond to shore crabs by burrowing deeper in the sediment. Norwegian M. balthica are not exposed to crabs naturally whereas Dutch M. balthica are naturally exposed to variable crab densities.Sites: Collection: the Balsfjord near Tromso, Norway, and the Wadden Sea near Harlingen, The Netherlands. Holding tanks: outdoor basins with a continuous flow of unfiltered water from the Wadden Sea.Method: We introduced a mixture of clams from both sites into experimental aquaria with a thick layer of sandy sediment. Twelve aquaria contained one shore crab; twelve had none. We measured burrowing depth 7 days after the start of each experiment.Result: Clams from the two sites show similar burrowing responses after exposure to predatory crabs, supporting the hypothesis that maintaining the potential for plasticity costs very little.

Kraternaya Bight (Yankich Island, the Kuril Islands) is a submerged crater of the active Ushishir Volcano characterized by terrestrial and underwater gasohydrothermal activity. The gasohydrothermal activity of the volcano produces a great influence on the physico-chemical characteristics of water column, as well as planktonic and bottom communities of marine organisms of the bight. The bivalve mollusks Macoma golikovi and Macoma calcarea are dominant macrobenthos species of Kraternaya Bight, which inhabit all the zones of underwater volcanic activity. Macoma golikovi occurs at a depth of 1–45 m. This species forms the densest aggregations (with an average biomass of 500–800 g m −2 and a population density of 500–700 ind. m −2 ) at a depth of 3–15 m, on steep underwater crater slopes (40–45°), on slightly silted gravel. The majority of gasohydrothermal vents are located at that depth. Macoma calcarea occurs at a depth of 5–55 m. In contrast to Macoma golikovi, this species is dominant at a depth of 25–40 m, on gentle slopes of the crater (to 20°), on silt and silty sand. That depth is less subjected to the influence of volcanic products. Its average biomass is 450–750 g m −2 with a population density of 200–450 ind. m −2 there. It is assumed that the differences in the distribution are due the fact that Macoma golikovi is mainly a suspension-feeder in Kraternaya Bight, and Macoma calcarea is a deposit-feeder. The studies conducted between 1987 and 1997 on the quantitative distribution of Macoma golikovi and Macoma calcarea in four areas of the bight, which had different intensity, temperature and chemical composition of gasohydrothermal vents and volcanic water seeping showed that both species formed settlements with lesser density in the area of the gasohydrothermal vents containing hydrogen sulfide. Beyond the area of volcanic activity, these species were most abundant. In 1993 and 1995, the mortality of the populations of the bivalve mollusks was observed in the areas of hydrogen sulfide gasohydrothermal vents and volcanic water seeping. In other areas of the bight, their abundance decreased sharply suggesting a relationship to the gasohydrothermal activity of the volcano in those years. Circumstantial evidence for this suggestion was that in 1993, the maximal concentrations of Fe, Zn, Cd, and Mn were observed in the tissues of both species. Macoma golikovi accumulated 5–10 times more Zn than Macoma calcarea. The concentration of Zn in the tissues of Macoma golikovi reached 2000 μg g −1 .

This article synthesises a series of studies concerned with physical, chemical and biological processes involved in sediment dynamics (sedimentation, erosion and mixing) of the Molenplaat tidal flat in the Westerschelde (SW Netherlands). Total sediment accretion rate on the flat (sand to muddy sand) was estimated to be ~2 cm yr-1, based on 210Pb and 137Cs profiles. 7Be showed maximum activity in the surface sediments during summer, reflecting accretion of fine silt at this time of year, and total vertical mixing of sediment to be in the order of 50 cm2 yr-1. The extent to which different physical and biological processes (tidal currents, air exposure, bio-stabilisation, biodeposition and bioturbation) contributed towards sediment dynamics was estimated. A sediment transport model based on physical factors estimated sedimentation rates of 1.2 cm yr-1, but did not account for tidal or seasonal variation in suspended particulate matter (SPM), wind or effects of spring-neap tidal cycles. When the model was run with an increased critical bed shear stress due to the microphytobenthos, net sedimentation rates increased 2-fold. These higher rates were in closer agreement with the rates derived from the depth profiles of radionuclides for the central region of the tidal flat (2.0 to 2.4 cm yr-1). Therefore a significant part of the sedimentation rate (~50%) may be explained by spatial-temporal changes in biological processes, including 'bio-stabilisation' by microphytobenthos, together with the enhanced biodeposition of silt by suspension feeders, and offset by processes of 'bio-destabilisation' by grazers and bioturbators. In the centre of the tidal flat there was a shift from high sediment stability in spring-summer 1996 to low sediment stability in autumn 1997, quantified by a significant reduction in critical erosion velocity of 0.12 to 0.15 m s-1, and accompanied by a 30- to 50-fold increase in sediment erosion rate. The change was associated with a shift from a tidal flat dominated by benthic diatoms and a low biomass of bioturbating clams (Macoma balthica), to a more erodable sediment with a lower microphytobenthos density and a higher biomass of M. balthica. Vertical mixing of sediment and organic matter, studied using a variety of tracers, was rapid and enhanced by advective water flow at sandy sites and by burrowing polychaetes and bivalves at silty sites. [KEYWORDS: Sediment dynamics · Erosion · Sedimentation · Mixing · Microphytobenthos · Biostabilisation · Bioturbation · Modelling]

The feeding guilds of four communities of molluscs living in the intertidal zone of Enseada in Caraguatatuba Bay and Barra Velha and Araca beaches along the Sao Sebastiao Channel were identified. Three-letter codes were assigned to each feeding guild, identifying the feeding habit, mobility patterns related to feeding, and food-capturing structures. Data were obtained from qualitative analyses of the digestive tract contents and from observations on the feeding behaviour of the gastropod Olivella minuta and the bivalves Tagelus plebeius, Macoma constricta, Tellina lineata, and T. versicolor. These data were complemented by published information on the feeding of congeneric or confamilial species. Six feeding guilds were identified. The SDS (suspension-feeder, discreetly motile, inhalant siphon and ctenidia) and DDS (deposit-feeder, discreetly motile, inhalant siphon and ctenidia) groups were more abundant. Suspension-feeding bivalves showed higher Trophic Importance Index (TI) values in all the study areas. Deposit-feeders showed high TI values only at Barra Velha beach. The species in the SDS feeding guild tended to occur in areas with lower salinity and moderately to well-sorted sediments with very fine sand predominating. The distribution of the feeding guilds correlated best with salinity, particle size distribution, and the contents of silt-clay and organic matter in the sediment.

We report results of a laboratory experiment that examines the effect of density-dependent interactions among infauna on organism–sediment–seawater relationships. The experiment includes a time series documentation of feeding behavior of the deposit- and suspension-feeding bivalve Macoma balthica , concurrent with measures of sediment biogeochemical processes that are affected by different feeding modes. We hypothesized that feeding behavior and emigration rates might shift with increasing density, and that these shifts would have cascading effects on benthic primary productivity, sediment–seawater exchanges, and porewater concentrations of ammonium and silicate. Macoma individuals were maintained in aquaria at three different densities (46, 230, and 460 individuals m −2 ) that fall within their natural abundances in the Chesapeake Bay. Individuals fed mostly on suspended material throughout the experiment, resorting to deposit feeding behavior only at the highest densities. Disturbances on the sediment surface during deposit feeding periods were not sufficient to impact benthic primary productivity and the associated interception effect of microalgae on sediment–seawater exchanges, as seen in previous studies. However, the bivalves impacted sediment fluxes directly through bioturbational activity, and these effects showed significant density interactions. For ammonium, fluxes ranged from 1.3 to 3.7 mmol m −2 d −1 and generally did not increase as a function of increasing Macoma density. Rather, highest fluxes generally were observed in intermediate density treatments. For silicate, a different trend was observed. Fluxes tended to parallel density, and ranged from 6.4 to 13.5 mmol m −2 d −1 , however, the relationship was not linear. These observations suggest that as infaunal density shifts, so does the balance of ammonium to silicate efflux. Thus, benthic population structure may impact water column processes not only through enhancing the flux, but also through alteration of elemental ratios. Experiments that uncover the strength of these interactions, and models that blend the actions of individuals, populations, and material components, would greatly enhance our understanding of the geochemistry of organism–sediment relations.

Pore water concentrations and benthic fluxes of dissolved Fe, P and N were measured at two coastal basins in the Gulf of Finland, northern Baltic Sea, during a seasonal cycle. The bioturbated inner coastal basin, where exchange of near-bottom water is efficient, had a better ability to retain P in sediments than the outer basin, where near-bottom water O2 concentration decreases during summer. Under the presence of O2 high pore water dissolved Fe:P ratio (>3.6 w:w) in surface layer of the sediment, measured especially in winter, indicated negligible or low P-release and high N:P ratio in the efflux. On the contrary, low Fe:P ratio (

The article deals with the structure of zoobenthocenoses in the Lithuanian economic zone of the Baltic Sea, productivity and distribution of certain typical cenoses depending on the prevailing type of ground and depth. Zoobenthocenoses are identified according to the dominant organism species (Mytilus edulis–Balanus improvisus, Macoma baltica–Nereis diversicolor). The economic zone under consideration is divided into six regions each of them being discussed separately. Nutrition of some commercial fishes (cod, flounder, vimba, Baltic herring, Baltic sprat, smelt), in whose diet benthos turns to be an important component, is also discussed.

Distribution of sediment in the sea bottom of Nida–Klaipėda water area is determined by peculiarities of the bottom relief, hydrodynamic conditions, geological structure of shallow and nearshore zones. Sediment reflects the main features of sedimentation and biocenoses that are related to this process. While studying regularities of the distribution of the biocenoses of the sea ground fauna, abundance of animals and amount of biomass, a direct connection can be noticed with the granulometric composition of the bottom sediment and the depth. Coarser sediment of the bottom make optimum conditions for the development of zoobenthos. With the sediment getting finer and the depth decreasing, the biomass of some zoobenthic organisms also decreases (e.g. Macoma baltica), but the abundance of polychaetes (Pygospio elegans) notably increases. In the water area of the ground dumping, zoobenthic populations resistant to dumping of grounds are forming. Intensive bottom sedimentation with greater amounts of substrata m...

During the first winter of their life, large numbers of the tellinid bivalve Macoma bal- thica (L.) migrate from their first-summer coastal nurseries on high tidal flats in the Wadden Sea to areas further offshore, including low tidal flats and subtidal areas in the Wadden Sea and adjacent parts of the North Sea. In the western part of the Dutch Wadden Sea, we compared M. balthica redis- tribution in cold and mild winters during the 29 years from 1973 to 2002 to find out whether migra- tion activity was related to winter character. Migration occurred on a larger scale in cold winters than in mild winters, with larger decreases in density of coastal populations and larger increases in den- sity of offshore populations following a cold winter than following a mild winter. In subtidal areas, such secondary recruitment, at an age of almost 1 yr, was higher than primary recruitment through post-larval settlement. In subtidal areas, M. balthica year-classes were particularly strong when their first winter had been cold and when primary first-summer recruitment had been abundant in nearby coastal tidal-flat areas.

Sediments in Sydney Harbour, Nova Scotia, are highly contaminated by polynuclear aromatic hydrocarbons (PAHs), polychlorinated biphenyls (PCBs), and heavy metals. Histopathologic and histochemical evaluations were made on the Baltic clam, Macoma balthica, exposed to 11 Sydney Harbour sediment samples. Histologic lesions in digestive gland (tubular dilation or atrophy, macrophage aggregates, tubular cell necrosis, and tissue inflammation) and gonads (macrophage aggregates, supporting cell, germ cell, and ovarian cell necroses) were frequently detected in clams exposed to the most contaminated sediments from the harbor. Clams exposed to these contaminated sediments also had the highest acid phosphatase activity. The average scores of tubular dilation or atrophy, ovarian cell necrosis, and the sums of mean digestive gland lesions correlated significantly with sediment PCBs, and the activities of acid phosphatase correlated significantly with sediment heavy metals, PAHs, and PCBs. Among the lesions, digestive gland tubular dilation or atrophy, tubular cell, germ cell, and ovarian cell necroses, and the activity of acid phosphatase are the best sublethal effect indicators in Macoma exposed to Sydney Harbour sediments. Key words: biomarkers, chronic biologic effects, clams, histology, histochemistry, Macoma balthica, marine sediment, polynuclear aromatic hydrocarbons, polychlorinated biphenyls.

The coastal environment of West Bengal is recognized as the most diversified and productive ecosystem among all the maritime states of India. This area faces organic pollution from domestic sewage and toxic pollution from industrial effluents leading to serious impacts on water and sediment quality as well as on biodiversity. This article aims at providing information on concentration levels of heavy metals (iron, zinc, copper, manganese, cadmium, and mercury) in the tissues of a suspension feeder bivalve Macoma birmanica, whole body tissues of five gastropods (Cerithidea cingulata, C. obtusa, Telescopium telescopium, Thais lacera and Nerita articulata) and four finfishes (Liza tade, Harpodon nehereus, Trichiurus sp., and Boleopthalmus sp.) collected from water bodies of Sagar Island (22° 19′ N; 80° 03′ E), the largest delta of Sundarbans. A general trend with the following in decreasing order occurred: Fe>Zn>Mn>Cu>Cd>Hg. Both species-dependent variability and temporal variations were pronounced. A high degree of organ specificity was evident in M. birmanica where gill and mantle exhibited higher metal accumulation due to the ion exchange property of the mucous layer covering these organs. Gastropods and finfishes were differentially selective for a range of metals, reflecting the availability of the elements in the environment and the diet and digestive physiology of these organisms. An overall elevated concentration of these metals was observed during monsoon months when low pH and salinity prevailed in the ambient medium. A continuous monitoring program is recommended to establish the studied organisms as bioindicators and to identify future changes to conserve the “health” of this fragile ecosystem.

Bioavailability of sediment-associated contaminants was assessed in parallel tests by means of a bivalve (Macoma nasuta) bioaccumulation assay and a novel in vitro digestive fluid extraction procedure. Digestive fluid was obtained from the deposit-feeding polychaete Arenicola brasiliensis and used to extract sediments from a U.S. Navy facility. Both the digestive fluid extraction and the bivalve bioaccumulation test identified Cd, Pb, high molecular weight polycyclic aromatic hydrocarbons (HPAHs), and polychlorinated biphenyls (PCBs) as the contaminants of concern; both procedures indicated that As, Cu, Hg, Ni, Zn, and low molecular weight polycyclic aromatic hydrocarbons were rarely, if ever, of concern. The only contaminant for which the techniques consistently differed was Cr, a result attributable to constraints on intestinal absorption of the metal by the bivalves. For Cd and Pb, the concentration attained in digestive fluid during a brief extraction was highly correlated with concentration attained in the bivalve after a 28-d exposure; correlation was marginal for HPAHs and was nonsignificant for PCBs. However, bulk sediment concentrations were equally good predictors of bioaccumulation because of minimal differences in bioavailability from the most contaminated sediments. In vitro contaminant extraction with the digestive fluid assay has potential as a screening tool to predict relative bioaccumulation risk, and has several advantages over traditional tests.

Juveniles of the bivalve Macoma balthica live on tidal flats in the Wadden Sea. This study examined the interaction of Macoma with the infaunal polychaetes Arenicola marina and Nereis diversicolor and the gastropod Retusa obtusa. The distribution of M. balthica spat on the flats, shortly after settlement in April, showed a positive correlation with the Arenicola distribution and a negative correlation with Nereis distribution. There were no locations where Macoma spat and Retusa occurred together. In August, Macoma spat had grown too large for predation by intertidal infauna. Small individuals of Macoma spat were found in stomachs of Arenicola (0.14 worm 1 ) and Nereis (0.05 worm 1 ). Laboratory experiments showed that Nereis and Retusa could reduce Macoma spat abundance, both in the absence and presence of sediment and alternative prey. Arenicola reduced the abundance of small Macoma (

As a characteristic buried tellinid bivalve, Macoma balthica has a long inhalent siphon that enables it to feed in two different ways: deposit and suspension feeding. To deposit feed efficiently on benthic microalgae, Macoma has to live close to the sediment surface, where it can graze an extensive surface area, but is within reach of many predators. Individuals that are more safely buried at a greater depth can only suspension feed, or deposit feed from a small surface area. We expected local differences in burying depth on intertidal mudflats to be caused by differences in immersion time (i.e. time available for feeding, particularly suspension feeding), since immersion time has been shown experimentally to affect body condition positively, and since body condition and burying depth in Macoma are postively related in the field. To test this we experimentally manipulated immersion time, and followed changes in burying depth and body condition. In the experiments, longer immersion time went consistently with greater burying depth of Macoma and higher body condition. On a transect in the western Wadden Sea, the deepest Macoma were indeed found at the intertidal level with the longest immersion time, but these were at that time not the animals with the highest body condition. Within each locality, however, body condition was positively correlated with burying depth. The experimental data and the within-locality data support the hypothesis that longer immersion time may influence burying depth through body condition. However, the fact that between-locality differences in burying depth seemed to be consistently related to immersion time, but not to body condition, indicates that body condition alone does not explain place-to-place variation in burying depth.

Bottom-water hypoxia (≤ 2 mg l -1 dissolved oxygen (DO)) greatly modifies the benthic habitat of estuaries, depending upon spatial extent, duration, and frequency. Bottom-water hypoxia often develops under conditions of density stratification, which inhibits vertical mixing, and warm temperatures, which enhance biological oxygen demand. Long-term, mid-channel data from the Neuse River Estuary in North Carolina permitted evaluation of how stratification and temperature combined to affect DO concentrations at the bottom. Salinity stratification (∆S) and water tempera- ture (T ) explained respectively 30 and 23% of the variance in bottom-water DO concentrations. The amount of salinity stratification required to induce bottom-water hypoxia declined with increasing water temperature. About 80% of observed hydrographic profiles exhibited bottom hypoxia when ∆S exceeded 5 psu and T exceeded 20°C. Using cross-channel hydrographic surveys as verification, we derived a general set of methods to estimate the lateral extent of low-DO bottom water from mid- channel hydrographic profiles. The method involves cross-estuary and along-estuary extrapolation based on assumption of a flat oxycline. Occasional violation of this assumption resulted in modest overestimation in cross-channel extent of low DO. Application of this method produced estimates ranging from 0 to 116 km 2 of bottom area (0 to 42% of the estuarine study area) exposed to hypoxia over all sample dates in summer 1997. The maximal bottom area exposed to hypoxia corresponded closely with an independent estimate of the area (100 km 2 ) that experienced almost complete mor- tality of Macoma spp. clams, the key benthic resource for demersal fishes and crabs. Consequently, mid-channel hydrographic profiles taken along the mid-channel of the estuary can be employed to assess the spatial scale of bottom habitat degradation due to hypoxia.

The bivalve Macoma balthica is a common species in the Wadden Sea and North Sea. Juveniles temporarily use nurseries in the high intertidal. To explain this nursery use, predation pressure was examined for both juvenile and adult Macoma at low and high tidal flats. The study was carried out in the eastern Dutch Wadden Sea.Shrimps Crangon crangon, adult crabs Carcinus maenas, gobies Pomatoschistus and juvenile flatfish were more abundant and larger on low than on high tidal flats, but 0-group Carcinus was more abundant on the high tidal flats. Crangon and 0-group Carcinus stomachs frequently contained Macoma remains. These predators selectively preyed on small 0-group Macoma, both in the field and in laboratory experiments. The effect of predation by epibenthic animals and birds, on the low and high tidal flats, was examined in exclosure experiments (2 mm mesh). There was no effect of epibenthos exclosure on adult Macoma. For 0-group Macoma, densities were higher in exclosures than in the controls where predators had normal access. The density reduction by epibenthic predators was much larger in the low than in the high intertidal. We found no effect of bird predation on densities of 0- and 1+group Macoma.Thus, 0-group Macoma is under high predation pressure by epibenthos in the low intertidal, especially by shrimps, while they are relatively safe in the high intertidal. However, most of the shellfish outgrow their epibenthic predators during their first summer. Therefore, it becomes safe for the bivalves to redistribute to locations where epibenthic predators are abundant, during their first winter. On the other hand, it did not become clear from this study why many of the larger Macoma leave the high intertidal. Concluding, the nursery use of Macoma-spat in the high intertidal is probably, at least partly, an adaptation to avoid epibenthic predation.

Twenty-six shell gravel sites at elevations between 3 and 35 m a.s.l. were found on the Aland Islands. The faunal composition was determined on 14 of these locations and Mytilus edulis and Macoma bulthic a were found everywhere. Another common species was Hydrobia ulvae. while H. ventrosa, Littorina saxatilis and Cerasto-derma glaucum were more sporadic. This species assemblage implies that the salinity of the area as about 10parts per thousand. when the animals lived there. The deposits on the N-NE hilltops generally have southerly to westerly aspect, while those of the south-central hilltops are easterly. On the basis of a C-14-dated stratified deposit it is suggested that each deposit was formed during a relatively short period of time. As a hilltop emerged from the sea. glacial debris was flushed down. Shell-bearing molluscs colonized the bare rock surface and were flushed down and formed the shell gravel deposit as the shoreline displacement continued. At a later stage. as the more concave part of the hilltop was subject to wave action. erosion was weaker and the shell gravel deposit was covered by beach sand. Since the hills may be of different elevation, it is concluded that trying to date ancient shorelines from shell gravel deposits is not justified. as each deposit originates from the hill with which it is associated.

The population development of the 1998 year class of the bivalve Macoma balthicd was studied by repeated sampling of a tidal flat area in the eastern Dutch Wadden Sea from May 1998 to August 2000. The juveniles migrated twice, once in mid-1998 from their primary settlement locations in the low sandy intertidal to the nursery in the high intertidal (spring migration, 25 % of all juvenile Mbalthica relocated from low to high intertidal) and once in late 1998/early 1999 from the nursery to the low intertidal and the subtidal (winter migration, between 8 and 15 % of all 0-group (less than 1 yr old) M. balthica relocated from high to low intertidal). During winter, M. balthica migration was most intensive at the lowest temperatures, Relative to the abundance on the tidal flats, the 1+ group (older than 1 yr) M. balthica (i.e. from the second summer onwards) were rare in the tidal channels of the Wadden Sea and were only slightly more common in the North Sea adjacent to the tidal inlet. During both the spring and winter migration, many bivalves disappeared from the tidal flat population. This could partly be explained by normal mortality and by emigration to the subtidal channels and the North Sea. The remaining mortality was probably due to the risks inherent in migration: predation during pelagic floating or not reaching the right locations, The number of bivalves that disappeared was very high. In spring, slightly more disappeared than relocated. In winter, > 4 times more bivalves disappeared than relocated, Migration is obviously a very dangerous activity, therefore great advantages must be associated with the nursery use that makes such migrations necessary. An analysis of literature data on the density of M. balthica shows that in the eastern Dutch Wadden Sea, 90 % of the population lives in the intertidal, about 10% in the adjacent North Sea, and a negligible fraction in the subtidal channels of the Wadden Sea.

Trace-metal concentrations in sediment and in the clam Macoma petalum (formerly reported as Macoma balthica), clam reproductive activity, and benthic macroinvertebrate community structure were 3 investigated in a mudflat 1 kilometer south of the discharge of the Palo Alto Regional Water Quality Control Plant (PARWQCP) in South San Francisco Bay, Calif. This report includes the data collected by U.S. Geological Survey (USGS) scientists for the period January 2013 to December 2013. These data serve as the basis for the City of Palo Alto’s Near-Field Receiving Water Monitoring Program, initiated in 1994. Following significant reductions in the late 1980s, silver (Ag) and copper (Cu) concentrations in sediment and M. petalum appear to have stabilized. Data for other metals, including chromium (Cr), mercury (Hg), nickel (Ni), selenium (Se), and zinc (Zn), have been collected since 1994. Over this period, concentrations of these elements have remained relatively constant, aside from seasonal variation that is common to all elements. In 2013, concentrations of Ag and Cu in M. petalum varied seasonally in response to a combination of site-specific metal exposures and annual growth and reproduction, as reported previously. Seasonal patterns for other elements, including Cr, Ni, Zn, Hg, and Se, were generally similar in timing and magnitude as those for Ag and Cu. In M. petalum, all observed elements showed annual maxima in January–February and minima in April, except for Zn, which was lowest in December. In sediments, annual maxima also occurred in January–February, and minima were measured in June and September. In 2013, metal concentrations in both sediments and clam tissue were among the lowest concentrations on record. This record suggests that regional-scale factors now largely control sedimentary and bioavailable concentrations of Ag and Cu, as well as other elements of regulatory interest, at the Palo Alto site. Analyses of the benthic community structure of a mudflat in South San Francisco Bay over a 40-year period show that changes in the community have occurred concurrent with reduced concentrations of metals in the sediment and in the tissues of the biosentinel clam, M. petalum, from the same area. Analysis of the M. petalum community shows increases in reproductive activity concurrent with the decline in metal concentrations in the tissues of this organism. Reproductive activity is presently stable (2013), with almost all animals initiating reproduction in the fall and spawning the following spring. The community has shifted from being dominated by several opportunistic species to a community where the species are more similar in abundance, a pattern that indicates a more stable community that is subjected to fewer stressors. In addition, two of the opportunistic species (Ampelisca abdita and Streblospio benedicti) that brood their young and live on the surface of the sediment in tubes have shown a continual decline in dominance coincident with the decline in metals; both species had short-lived rebounds in abundance in 2008, 2009, and 2010. Heteromastus filiformis (a subsurface polychaete worm that lives in the sediment, consumes sediment and organic particles residing in the sediment, and reproduces by laying its eggs on or in the sediment) showed a concurrent increase in dominance and, in the last several years before 2008, showed a stable population. H. filiformis abundance increased slightly in 2011–2012 and returned to pre-2011 numbers in 2013. An unidentified disturbance occurred on the mudflat in early 2008 that resulted in the loss of the benthic animals, except for those deep-dwelling animals like Macoma petalum. Animals immediately returned to the mudflat in 2008, which was the first indication that the disturbance was not due to a persistent toxin or to anoxia. The reproductive mode of most species present in 2013 is reflective of the species that were available either as pelagic larvae or as mobile adults. Although oviparous species were lower in number in this group, the authors hypothesize that these species will return slowly as more species move back into the area. The use of functional ecology was highlighted in the 2013 benthic community data, which show that the animals that have now returned to the mudflat are those that can respond successfully to a physical, nontoxic disturbance. Today, community data show a mix of animals that consume the sediment, filter feed, have pelagic larvae that must survive landing on the sediment, and brood their young. USGS scientists continue to observe the community’s response to the 2008 defaunation event because it allows them to examine the response of the community to a natural disturbance (possible

The persistence of prey encountering intense predation varies by species, prey density, and habitat type; however, the collective impact of these factors has rarely been tested experimentally in natural marine systems. Using the thin-shelled clams Mya arenaria and Macoma balthica as prey, and the main epibenthic predator of whole adult clams, the blue crab Callinectes sapidus, we conducted a series of experiments in Chesapeake Bay tributaries that (1) links field abundance and distribution of bivalve prey species with habitat-specific mortality patterns; (2) represents the first comprehensive field test of species-specific, habitat-specific, and density-dependent mortality for subtidal, soft-bottom, deep-burrowing prey; and (3) thereby enables development of a conceptual model to be used as a heuristic tool linking predator–prey dynamics, habitat type, and evolutionary defense tactics for marine benthos. In 15 years of field monitoring, Mya was more common in sand than mud habitats, and Macoma was widely distributed and at higher densities than Mya in mud and sand. In field experiments, mortality of both Mya and Macoma was density dependent in those habitats where the clams are common. The blue crab population in the field exhibited a type III “guild functional response” on Mya in sand, and on Macoma in both mud and sand. Mortality was lower in sand than mud for Mya, and similar in mud and sand for Macoma, correlating with the high abundances of Mya in sand and Macoma in sand and mud. The persistence of large juvenile and adult bivalves when confronted with intense predation derived substantially from a low-density refuge from predation that varied in a species-specific manner with habitat type, demonstrating the species-specific importance of density and habitat to clam survival. We developed a conceptual model detailing the relative importance of behavior, morphology, habitat features, and the basic components of predator–prey interactions to the survival of bivalve molluscs. At one extreme are bivalve molluscs, such as oysters, that emphasize morphological refuges that increase the predator's handling time. At the other extreme are bivalves, such as Mya and Macoma, that reduce predator encounter rates. The model is intended to be used as a heuristic tool to develop testable hypotheses.

High recruitment of the bivalves Cerastoderma edule, Mytilus edulis, Macoma balthica and Mya arenaria in summer after severe winters is an often reported phenomenon in the Wadden Sea. After the severe winter of 1995/96 however, only Cerastoderma and Mytilus followed this pattern in the Sylt-Romo Bight. Repeated sampling of Cerastoderma, Macoma and Mya following a severe (1995/96), a moderate (1996/97), and a mild winter (1997/98) revealed that early recruitment was highest after the mild winter. In Cerastoderma the eventual high recruitment at the end of summer 1996 was caused by reduced benthic mortality. Low recruitment of Macoma and Mya after the severe winter may have been caused by a higher susceptibility to epibenthic predation and/or a higher susceptibility to passive re-suspension than in Cerastoderma and Mytilus. In all cases, post-settlement processes were decisive for reproductive success.

At large spatial scales (>1 km), the forces structuring marine communities are diverse and include oceanographic, geological, and human processes, as well as availability of regional species pools. Few studies in marine and estuarine systems have examined the joint effects of predation (top-down) and food availability (bottom-up) in controlling populations at these scales. We compared the relative influence of top-down and bottom-up factors to population abundance of the infaunal clam Macoma balthica in two ecosystems differing in spatial extent by an order of magnitude: the York River, 50 km long, and the Rhode River, 5 km long. In both rivers, two habitat types could be differentiated by either high or low density of Macoma. In the York, Macoma abundance was tenfold greater upriver than downriver, predator abundance was greater in the high clam density habitat (i.e. upriver), and predation was only evident where clams were dense (i.e. upriver, where food availability for predatory crabs was elevated). In this large-scale system, predators were apparently unable to travel between upriver and downriver habitats where food availability differed. Moreover, sedimentary carbon (i.e. food for deposit-feeding clams) was significantly greater in the high clam-density habitat; this is consistent with the hypothesis that bottom-up factors principally dictate clam density, and moreover, that clam density drives predator density. In the Rhode River, a smaller-scale system, predators could easily move among habitats differing in clam density. Macoma abundance and predator abundance were both threefold greater in mud than sand. In contrast, predation intensity was higher in sand than mud, consistent with a hypothesis of top-down control. As in the large-scale system, sedimentary carbon was significantly greater in the high clam-density habitat (i.e. mud). In this smaller scale system, both top-down and bottom-up factors contributed to habitat-specific patterns in clam population abundance. Hence, in this marine benthic ecosystem, bottom-up control was important at both large and small spatial scales, whereas top-down control was only important at the smaller spatial scale.