Variable responses of native eelgrass Zostera marina to a non-indigenous bivalve Musculista senhousia

The transport and establishment of non-indigenous species in coastal marine environments are increasing worldwide, yet few studies have experimentally addressed the interactions between potentially dominant non-native species and native organisms. We studied the effects of the introduced mussel Musculista senhousia on leaf and rhizome growth and shoot density of eelgrass Zostera marina in San Diego Bay, California. We added M. senhousia over a natural range in biomass (0–1200 g dry mass/m²) to eelgrass in transplanted and established beds. The effects of the non-indigenous mussel varied from facilitation to interference depending on time, the abundance of M. senhousia, and the response variable considered. Consistent results were that mussel additions linearly inhibited eelgrass rhizome elongation rates. With 800 g dry mass/m² of M. senhousia, eelgrass rhizomes grew 40% less than controls in two eelgrass transplantations and in one established eelgrass bed. These results indicate that M. senhousia, could both impair the success of transplantations of eelgrass, which spread vegetatively by rhizomes, and the spread of established Z. marina beds to areas inhabited by M. senhousia. Although effects on leaf growth were not always significant, in August in both eelgrass transplantations and established meadows leaf growth was fertilized by mussels, and showed a saturation-type relationship to sediment ammonium concentrations. Ammonium concentrations and sediment organic content were linear functions of mussel biomass. We found only small, non-consistent effects of M. senhousia on shoot density of eelgrass over 6-month periods. In established eelgrass beds, but not in transplanted eelgrass patches (≈0.8 m in diameter), added mussels suffered large declines. Hence, eelgrass is likely to be affected by M. senhousia primarily where Z. marina beds are patchy and sparse. Our study has management and conservation implications for eelgrass because many beds are already seriously degraded and limited in southern California where the mussel is very abundant. Received: 31 May 1997 / Accepted: 4 September 1997     

Subtidal populations of the blue mussel <Emphasis Type="Italic">Mytilus edulis</Emphasis> as key determinants of waterfowl flocks in the southeastern Barents Sea

Ornithological surveys conducted over the Pechora Sea (the southeastern part of the Barents Sea) in the 1990 s revealed huge non-nesting flocks of marine ducks, the largest in the European North. Especially dense waterfowl aggregations are constantly observed at the shallows near Dolgij Island during molting period and migration to wintering places. All the marine ducks flocking there are specialized benthos feeders predominantly consuming mussels Mytilus edulis. At the same time, numerous previous benthic studies in the Pechora Sea did not reveal mussels near Dolgij Island where benthic biomass was somewhat lower than in the adjacent areas (Denisenko in Mar Ecol Prog Ser 258:109–123. 2003) which left the food source for these abundant bird flocks enigmatic. In the course of an expedition in summer 2007 we found subtidal populations of M. edulis in shallows to the southwest of Dolgij Island. These populations were confined to a coastal zone and were characterized by a highly disjunct distribution with the biomass reaching up to 4 kg m⁻². We describe these subtidal populations as well as an intertidal mussel population on the western shore of Dolgij Island.     

Habitat engineering by the invasive zebra mussel <Emphasis Type="Italic">Dreissena polymorpha</Emphasis> (Pallas) in a boreal coastal lagoon: impact on biodiversity

Habitat engineering role of the invasive zebra mussel Dreissena polymorpha (Pallas) was studied in the Curonian lagoon, a shallow water body in the SE Baltic. Impacts of live zebra mussel clumps and its shell deposits on benthic biodiversity were differentiated and referred to unmodified (bare) sediments. Zebra mussel bed was distinguished from other habitat types by higher benthic invertebrate biomass, abundance, and species richness. The impact of live mussels on biodiversity was more pronounced than the effect of shell deposits. The structure of macrofaunal community in the habitats with >10³ g/m² of shell deposits devoid of live mussels was similar to that found within the zebra mussel bed. There was a continuous shift in species composition and abundance along the gradient ‘bare sediments—shell deposits—zebra mussel bed’. The engineering impact of zebra mussel on the benthic community became apparent both in individual patches and landscape-level analyses.     

Impacts of mussel (<Emphasis Type="Italic">Mytilus galloprovincialis</Emphasis>) farming on oxygen consumption and nutrient recycling in a eutrophic coastal lagoon

Fluxes of oxygen, nitrogen and phosphorus were determined in two areas of the Sacca di Goro lagoon, at a site influenced by the farming of the mussel Mytilus galloprovincialis and a control site. Mussel farming induced intense biodeposition of organic matter to the underlying sediments, which stimulated sediment oxygen demand, and inorganic nitrogen and phosphorus regeneration rates compared to the nearby control station. Overall benthic fluxes (–11.4 ± 6.5 mmol O₂ m⁻² h⁻¹; 1.59 ± 0.47 mmol NH₄⁺ m⁻² h⁻¹ and 94 ± 42 μmol PO₄³⁻ m⁻² h⁻¹) at the mussel farm are amongst the highest ever recorded for an aquaculture impacted area and question the belief that farming of filter-feeding bivalves has inherently lower impacts than finfish farming. In situ incubations of intact mussel ropes demonstrated that the mussel rope community was an enormous sink for oxygen and particulate organic matter, and an equally large source of dissolved inorganic nitrogen and phosphate to the water column. Overall, a one meter square area of␣mussel farm (mussel ropes and underlying sediment) was estimated to have an oxygen demand of 46.8 mmol m² h⁻¹ and to regenerate inorganic nitrogen and phosphorus at rates of 8.5 and 0.3 mmol m² h⁻¹, with the mussel ropes accounting for between 70 and more than 90% of the overall oxygen and nutrient fluxes. Even taking into account that within the farmed area of the Sacca di Goro lagoon, there are 15–20 m⁻² of open water for each one covered with mussel ropes, the mussel ropes would account for a large and often dominant part of overall oxygen and nutrient fluxes. These results demonstrate that it is essential to take into account the activity of the cultivated organisms and their epiphytic community when assessing the impacts of shellfish farming. Overall, whilst grazing by the mussel rope community could act as a top-down control on the phytoplankton, most of the ingested organic matter is rapidly recycled to the water column as inorganic nutrients, which would be expected to stimulate phytoplankton growth. Consequently, the net effect of the mussel farming on phytoplankton dynamics, may be to increase phytoplankton turnover and overall production, rather than to limit phytoplankton biomass.     

Variability in annual recruitment success as a determinant of long-term and large-scale variation in annual production of intertidal Wadden Sea mussels (<Emphasis Type="Italic">Mytilus edulis</Emphasis>)

To understand the background of the strong variation and recent decline of stocks and production of mussels (Mytilus edulis) on tidal flats of the Wadden Sea, we analysed long-term (twice-annual for 26 years) and multi-station (15 sites) estimates of numbers, mean individual weights, biomass, and annual production on Balgzand, a 50-km² tidal-flat area in the westernmost part of the Wadden Sea (The Netherlands). Somatic production was estimated from summed growth increments of soft tissues per half-year period and expressed in ash-free dry mass (AFDM). In adults, positive values in spring/summer regularly alternated with negative values in autumn/winter, when up to ∼25% (mean: 14%) of individual weight gains in the preceding season were lost. No weight losses were observed during the first winter of the life of mussels. The 26-year mean of net somatic tissue production P amounted to 5.5 g AFDM m⁻² a⁻¹ at a mean biomass B of 3.2 g AFDM m⁻²; the ratio P/B varied strongly with age composition of the mussel population and ranged between 0.5 and 3.0 a⁻¹ (mean: 1.7). Within the restricted areas of mussel beds, mean biomass and annual production values were two orders of magnitude higher. In the Wadden Sea, mussel beds cover a typical 1% of extensive tidal flat areas. Numerical densities of recruits showed straight-line relationships with subsequent life-time year-class production. Once recruits had reached an age of ∼10 months, their numbers predicted subsequent production within narrow limits. Production per recruit averaged 0.21 g AFDM for 10-mo recruits and was not related to recruit density. Local variation in annual production varied strongly, with maximal values between mid-tide and low-tide level, where recruitment was also maximal. Production per recruit was higher at low than at high intertidal levels. Frequently failing recruitment is indicated as the main cause of declining mussel stocks in the Wadden Sea. As in other bivalve species, a declining frequency of the occurrence of cold winters appears to govern declining recruitment success and consequently declining production.     

Documentation of sites of intertidal blue mussel (Mytilus edulis L.) beds of the Lower Saxonian Wadden Sea, southern North Sea (as of 2003) and the role of their structure for spatfall settlement

Field surveys (dating back to 1950) and aerial photograph series (dating back to 1966) were evaluated to determine sites of intertidal blue mussel (Mytilus edulis) beds at the Wadden Sea coast of Lower Saxony. Maps were prepared indicating sites of blue mussel beds during the last decades. A table gives additional information on the presence (or absence) of blue mussel beds at each site at the time of large-scale surveys. Altogether 187 sites of M. edulis beds were recorded in the investigation area. In spring 1996, there were still only 19 sites where mussel beds still occurred, although at 51 sites residual mussel-bed structures were present, e.g. shell bases of former beds or protruding patches (which had been occupied by M. edulis before the beds vanished) and open spaces. At that time, the majority of the sites contained neither mussel beds nor mussel-bed structures. The analysis of recent data confirmed that mussel larvae have preferred to settle in sites of present mussel beds and sites with bases of former mussel beds. There was no preferential selection of one of these categories (settled beds vs. shell bases). On the other hand, the presence of mussel beds or mussel bed structures is not obligatory for settlement, because sites without those structures were also re-settled by the spatfall in 1996, even though on a smaller scale.     

Subtidal and intertidal mussel beds (Mytilus edulis L.) in the Wadden Sea: diversity differences of associated epifauna

In 1997 and 1998, surveys were performed to compare species composition, abundance and diversity of non-attached epifauna (>1 mm) in low intertidal and adjacent shallow subtidal zones of three mussel beds (Mytilus edulis L.) near the island of Sylt in the North Sea. The community structure was similar when compared within tidal zones: no significant differences in species numbers and abundances were recorded between locations and between years. A comparison between tidal zones, however, revealed higher diversity, species densities and total species numbers in the subtidal (per 1,000 cm²: H ′=2.0±0.16; 12 ±1 species density; 22 species) than the intertidal zone (per 1,000 cm²: H ′=0.7±0.27; 6±2 species density; 19 species). Abundances significantly dropped with increasing submergence from 2,052 (±468) m^–2 to 1,184 (±475) m^–2. This was mainly due to significantly higher densities of both juvenile periwinkles, Littorina littorea, and crabs, Carcinus maenas, in intertidal mussel beds. However, many less dominant species were significantly more abundant in subtidal mussel beds. This study revealed that in the non-attached epifaunal community of mussel beds the tidal level effect within metres was strong, whilst the spatial variability in a much wider (kilometre) range but the same tidal level was negligible. The high epifaunal diversity in the subtidal zone suggests that the protective measures for mussel beds against the effects of mussel fishery should be extended from the intertidal to the subtidal zone, if the integrity of the mussel bed community in the Wadden Sea National Park is to be maintained. Electronic Publication     

Population dynamics of the invasive Pacific oyster Crassostrea gigas during the early stages of an outbreak in the Wadden Sea (Germany)

Since the late 1990s, the Pacific oyster (Crassostrea gigas) has spread into the East Frisian Wadden Sea (Germany). This invasion provided an opportunity to study the population dynamics and the patterns of spread during the initial bioinvasion process. With its source area in The Netherlands, the bioinvasion continues in an eastward direction, as documented by a gradient of high abundances in the west and low abundances in the east during the first study year. One year later, abundances of the Pacific oyster were more heterogenic and differed between adjacent tidal basins. The increase in population sizes at all study sites was very high, reaching levels similar to native occurrence populations. The growth constant (K) varied between 0.300 and 0.990 year⁻¹. The mussel bed with the highest densities had a mean abundance of >300 ind. m⁻², and a maximum of 1,460 ind. m⁻². Furthermore, the bioinvasion was facilitated by a low mortality (Z) found for populations between 0.5 and 1.5 years old (Z = 0.03–0.13 year⁻¹). At present, Pacific oysters are well established at several locations in the East Frisian Wadden Sea and may become with these reproductive potential self-sustaining populations.     

Spring-to-summer changes and regional variability of benthic processes in the western Canadian Arctic

Seasonal dynamics in the activity of Arctic shelf benthos have been the subject of few local studies, and the pronounced among-site variability characterizing their results makes it difficult to upscale and generalize their conclusions. In a regional study encompassing five sites at 100–595 m water depth in the southeastern Beaufort Sea, we found that total pigment concentrations in surficial sediments, used as proxies of general food supply to the benthos, rose significantly after the transition from ice-covered conditions in spring (March–June 2008) to open-water conditions in summer (June–August 2008), whereas sediment Chl a concentrations, typical markers of fresh food input, did not. Macrobenthic biomass (including agglutinated foraminifera >500 μm) varied significantly among sites (1.2–6.4 g C m⁻² in spring, 1.1–12.6 g C m⁻² in summer), whereas a general spring-to-summer increase was not detected. Benthic carbon remineralisation also ranged significantly among sites (11.9–33.2 mg C m⁻² day⁻¹ in spring, 11.6–44.4 mg C m⁻² day⁻¹ in summer) and did in addition exhibit a general significant increase from spring-to-summer. Multiple regression analysis suggests that in both spring and summer, sediment Chl a concentration is the prime determinant of benthic carbon remineralisation, but other factors have a significant secondary influence, such as foraminiferan biomass (negative in both seasons), water depth (in spring) and infaunal biomass (in summer). Our findings indicate the importance of the combined and dynamic effects of food supply and benthic community patterns on the carbon remineralisation of the polar shelf benthos in seasonally ice-covered seas.     

Biotic resistance to invasion along an estuarine gradient

Biotic resistance is the ability of native communities to repel the establishment of invasive species. Predation by native species may confer biotic resistance to communities, but the environmental context under which this form of biotic resistance occurs is not well understood. We evaluated several factors that influence the distribution of invasive Asian mussels (Musculista senhousia) in Mission Bay, a southern California estuary containing an extensive eelgrass (Zostera marina) habitat. Asian mussels exhibit a distinct spatial pattern of invasion, with extremely high densities towards the back of Mission Bay (up to 4,000 m⁻²) in contrast with near-complete absence at sites towards the front of the bay. We established that recruits arrived at sites where adult mussels were absent and found that dense eelgrass does not appear to preclude Asian mussel growth and survival. Mussel survival and growth were high in predator-exclusion plots throughout the bay, but mussel survival was low in the front of the bay when plots were open to predators. Additional experiments revealed that consumption by spiny lobsters (Panulirus interruptus) and a gastropod (Pteropurpura festiva) likely are the primary factors responsible for resistance to Asian mussel invasion. However, biotic resistance was dependent on location within the estuary (for both species) and also on the availability of a hard substratum (for P. festiva). Our findings indicate that biotic resistance in the form of predation may be conferred by higher order predators, but that the strength of resistance may strongly vary across estuarine gradients and depend on the nature of the locally available habitat.     

Mytilid mussels: global habitat engineers in coastal sediments

Dense beds of mussels of the family Mytilidae occur worldwide on soft-bottoms in cold and warm temperate coastal waters and have usually been considered hot spots of biodiversity. We examined intertidal mussel beds at four distant locations around the globe with the same sampling method, to find out whether this “hot spot” designation holds universally. We studied species assemblages within the matrices of byssally interconnected mussels engineered by Mytilus edulis in the North Sea, by mixed Perumytilus purpuratus and Mytilus chilensis at the southern Chilean coast, by Musculista senhousia in the Yellow Sea and by Xenostrobus inconstans at the coast of southern Australia. In all cases, species assemblages inside mussel beds were significantly different from those outside with many species being restricted to one habitat type. However, species richness and diversity were not generally higher in mussel beds than in ambient sediments without mussels. In the North Sea (M. edulis) and at the Chilean coast (P. purpuratus, M. chilensis), mussel beds have markedly higher species numbers and diversities than surrounding sediments, but this was not the case for mussel beds in Australia (X. inconstans) and the Yellow Sea (M. senhousia) where numbers of associated species were only slightly higher and somewhat lower than in adjacent sediments, respectively. In conclusion, although soft bottom mytilid mussels generally enhance habitat heterogeneity and species diversity at the ecosystem level, mussel beds themselves are not universal centres of biodiversity, but the effects on associated species are site specific.     

Oxygen and ammonium dynamics during a farming cycle of the bivalve <Emphasis Type="Italic">Tapes philippinarum</Emphasis>

Fluxes of dissolved oxygen and ammonium across the water sediment interface were measured in a control and in an experimental area farmed with the clam Tapes philippinarum. Young clams were seeded in March 2003 at mean (~500 ind m⁻²) and high (~1500 ind m⁻²) densities in a sandy area (2100 m²) of the Sacca di Goro Lagoon, Italy. Approximately every two months, until October 2003, intact sediment cores were collected and incubated in the light and in the dark and surface sediments (0–2 cm) were analysed for organic matter and nitrogen content. Clams farming induced pronounced changes in sediment characteristics and metabolism. Oxygen consumption and ammonium production at the high density area were, on average, 3 to 4 and 1.9 to 4.9 folds higher than those measured in the control field respectively; rates were positively correlated with clams biomass. Experimental fields resulted “Net and Total Heterotrophyc” in 3 out of 4 sampling dates and clams were the major factor shifting the benthic system towards this status. In only one occasion the appearance of the macroalgae Ulva spp. pushed the system rapidly towards hyperautotrophic conditions. Our results indicated that clams have the potential to drive benthic metabolism in farmed areas and to sustain macroalgal growth through regeneration of a limiting nutrient for seawater as inorganic N.     

Composition, abundance and stratification of soft-bottom macrobenthos from selected areas of the Ross Sea shelf (Antarctica)

Composition, abundance and stratification of soft-bottom macrobenthos were studied at three selected sites on the Ross Sea shelf (Antarctica) with different geomorphology and sedimentation regimes. Sites A (southwest Ross Sea, 810 m depth) and B (Joides basin, 580 m depth) were characterized by biogenic mud and clay sediments, whereas site C (Mawson bank, 450 m depth) featured sandy sediments mixed with a conspicuous biogenic component characterized by shells and tests of calcareous invertebrates (mainly barnacles of the genus Bathylasma). The macrofauna of sites A and B was mainly composed of infaunal polychaetes and bivalves. The assemblages comprised both surface and sub-surface deposit feeders, including some conveyor-belt polychaetes (Maldanidae and Capitellidae) that are responsible for high sediment mixing and bioturbation. The macrobenthos of site C was dominated by crustaceans, polychaetes and echinoderms (ophiuroids), and mainly by filter feeders and epifaunal or interstitial forms. Abundances were higher (up to 1040 ind. m⁻²) at site B than at sites A and C (430 and 516 ind. m⁻², respectively). At sites A and B the benthos was mainly concentrated in the upper 5 cm of the sediment, and abundances declined sharply in the deeper sediment layers. These results indicate a high degree of consistency between sediment features and benthic community structure, which are both strongly related to local hydrography and bottom dynamics. Sites A and B represent areas where the organic input to the seafloor by vertical sedimentation from the upper water column is high. Site C, however, is characterized by high sediment instability and food particles advecting mainly horizontally. The community is more physically controlled by unpredictable, and probably frequent, disturbance events (e.g., bottom turbid currents, sediment reworking and displacement). Individuals were relatively small, indicating that probably they are not able to grow up to the adult size and reproduce. The community may be represented by “pseudopopulations” depending on the settlement of larvae invading from neighbouring areas. Accepted: 23 October 1998     

Epiphytic Refugium: Are Two Species of Invading Freshwater Bivalves Partitioning Spatial Resources?

Enumeration of benthic (bottom dwelling) and epiphytic (attached to plants) zebra and quagga mussels (Dreissena polymorpha and D. bugensis, respectively) at Lake Erie near-shore sites in fall of 2000 revealed an unexpected prevalence of the zebra mussel on submerged plants. Even at Buffalo, New York, USA, where benthic dreissenids have been 92–100% quagga mussel since 1996, zebra mussels constituted 30–61% of epiphytes numerically. This may reflect a partitioning of settling space consistent with interspecific competition. A seasonal epiphytic refugium might allow the zebra mussel to persist even where the benthos is almost exclusively quagga mussel. This revised version was published online in July 2006 with corrections to the Cover Date.     

A rapid assessment of the sedimentary buffering capacity towards free sulphides

Combined field surveys and laboratory studies were conducted in two Italian coastal lagoons, which differ for geomorphology, hydrodynamics and eutrophication degree (Sacca di Goro and Lesina lagoons, Adriatic Sea). Research aimed at assessing with a rapid technique the potential buffering capacity of sedimentary iron towards sulphides. In Spring and Summer 2004, the main pools of iron and sulphides were analysed in the uppermost sediment horizon (0–5 cm) at four stations in each lagoon. In parallel, experiments with laboratory incubations of sediment slurries were carried out at two sites in each lagoon in order to assess the sediment capacity of binding and retaining sulphides. Sediment slurries were kept stirred and anoxic with N₂ purging. Aliquots of dissolved sulphides (DS) were then added and DS concentrations were monitored until they were undetectable. On average, the total reactive iron (RFe), extracted with 6 N HCl, ranged from 170 to 400 μmol cm⁻³ in the Sacca di Goro stations, and comprised between 40 and 150 μmol cm⁻³ in the Lesina sites. The labile iron ferric quota (LFe: extractable with 0.5 N HCl) is considered representative of the microbially reducible iron fraction and was highest in spring in Sacca di Goro (up to 20 μmol cm⁻³). Differences among stations evidenced by PCA analysis, can be inferred from RFe, LFe and AVS, which represent the iron buffer and its saturation status, respectively. The sedimentary DS uptake was 6 μmol cm⁻³ of fresh sediment in Lesina and 8–12 μmol cm⁻³ in Sacca di Goro, indicating a direct relationship between DS removal and iron availability. Guest editors: A. Razinkovas, Z. R. Gasiūnaitė, J. M. Zaldivar & P. Viaroli European Lagoons and their Watersheds: Function and Biodiversity     

Carbonate standing stock and carbonate production of the bryozoanPentapora fascialis in the North-Western Mediterranean

Summary Pentapora fascialis, one of the largest living bryozoan, is often a predominant part of the benthos on hard subtidal bottoms in the Mediterranean Sea. Conversion factors calculated from laboratory measurements of colony size, biomass and skeleton weight, combined with density of colonies and mean annual growth rate allowed the estimation of carbonate standing stock, biomass and carbonate production ofPentapora fascialis in five sites in the Ligurian Sea. Carbonate standing stock ranged from 281 to 2490 g·m⁻², colony biomass varied from 8.82 to 78.01 g·m⁻², with a ratio biomass to carbonate standing stock of about 3%. Carbonate production of the bryozoan ranged in the five sites from 358 to 1214 g·m⁻²·y⁻¹. If compared with the few data available on carbonate production of bryozoans and other sublittoral benthic bioconstructors in the temperate regions,Pentapora fascialis has to be considered one of the major contributors to the carbonatebudget.     

Assessment of nutritional subsidies to freshwater mussels using a multiple natural abundance isotope approach

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Impact of the infaunal Manila clam, <Emphasis Type="Italic">Ruditapes philippinarum</Emphasis>, on sediment stability

The aim of this study was to examine the impact of bioturbation by the Manila clam, Ruditapes philippinarum, on sediment stability. A laboratory benthic annular flume system (AFS) was deployed to evaluate the relationship between sediment stability of a subtidal mudflat and density of the infaunal clam under the influence of different current velocities. There was a significant correlation between mean erosion rate and current velocities in all treatments with clams (p < 0.001). There was also a significant correlation between mean erosion rate and R. philippinarum density (p < 0.001), reflecting bioturbation-enhanced sediment erosion. The effects of clam density on sediment erodability were more marked at the lower current velocities. In the control, the critical erosion velocity (Ū_crit) was about 32 cm s⁻¹. With increasing R. philippinarum density, Ū_crit decreased down to the minimum value of about 20 cm s⁻¹ at a density of 206 clams m⁻². This study demonstrated that the burrowing activity of R. philippinarum reduces sediment stability, particularly at relatively low current velocities (25 cm s⁻¹) and at densities below those found in the clam cultivation areas within the Sacca di Goro lagoon.     

Benthic oxygen flux in the highly productive subarctic Lake Myvatn, Iceland: In situ benthic flux chamber study

In situ paired light and dark-stirred benthic flux chambers were used to estimate dissolved oxygen flux across the sediment–water interface in Lake Mývatn, Iceland. Three sampling stations were selected, each station reflecting a specific sedimentary environment, benthic communities, and water depth. During this study the phytoplankton density was low. Spatial and seasonal variations of bottom DO concentration and DO flux have been observed during this study. The oxygen consumption rate at all study sites had a mean of –89 (±44) mmol m^–2 d^–1 while the oxygen production rate due to benthic algae had a mean of 131 (±103) mmol m^–2 d^–1. There was a strong correlation (r=0.91) between oxygen consumption rate and temperature. This was presumably because of the temperature influence on rates of microbial and macrobenthic processes. The mean benthic primary production rate at all study sites was 1216 (±957) mg C m^–2 d^–1 between June 2000 and February 2001. Annual gross benthic primary production was estimated from the gross mean daily benthic DO production (P) and Redfield's C:O₂ ratio of 106:138 to be 420 g C m^–2 y^–1 at station HO, 250 g C m^–2 y^–1 at B2 and 340 g C m^–2 y^–1 at station 95. Thus, the mean gross benthic primary production was estimated as 1151 mg C m^–2 d^–1 at station HO, 685 mg C m^–2 d^–1 at station B2, and 932 mg C m^–2 d^–1 at station 95.     

Phytoplankton composition of the Kandalaksha Gulf, Russian White Sea: Dinophysis and lipophilic toxins in the blue mussel (Mytilus edulis)

Dinophysis acuminata and D. norvegica were observed in plankton net samples during the summer of 2002 from the Kandalaksha Gulf in the White Sea (North European Russia). Prorocentrum lima was found as an epiphyte on subtidal macroalgae in August, but not observed in plankton net samples. Protein phosphatase 2A (PP2A) inhibition measured 127.8 ng OA-equivalent/g of mussel (Mytilus edulis) hepatopancreas from samples collected a few days after when Dinophysis was recorded at a density of 1550 cells L⁻¹. Liquid chromatography–mass spectrometry confirmed presence of several classes of lipophilic shellfish toxins associated with Dinophysis spp. in the mussels including okadaic acid, dinophysistoxin-1, pectenotoxins and yessotoxins. No azaspiracid was detected. This represents the first identification of phycotoxicity in the White Sea.