Succession of Fouling Communities on an Artificial Substrate of a Mussel Culture in the White Sea

The pattern of development of a fouling community, based on research data on their formation in the mussel culture in the White Sea is presented. The succession may come to an end either by the mussel community or by the community of Styela rustica, which is analogous to the natural epitbenthic biocenosis of Laminaria saccharina. It also might develop with the deletion of some stages. The presence of the mussel phase is not necessary in the development of fouling. The final stage of the fouling development is determined, apparently, by hydrological conditions.     

Fouling Communities of Mussel Aquaculture Installations in the White Sea

We have classified fouling communities developing on artificial substrates of mussel mariculture plants in Kandalakshskii Bay of the White Sea. Several major types of fouling have been distinguished: mussel fouling; fouling similar to the epifauna of the benthos biocenosis of Laminaria saccharinadominated by the ascidian Styela rusticaor sponges Halichondria paniceaand Halisarca dujardini; young fouling developed by algae and the ascidians of the genus Molgula; and ecotone communities in-between these types of fouling. The characteristics of the distinguished communities are provided.     

Mutual effects of several fouling organisms of the White Sea (<Emphasis Type="Italic">Mytilus edulis,Styela rustica</Emphasis>, and <Emphasis Type="Italic">Hiatella arctica</Emphasis>) on their growth rate and survival

The mutual effects of several fouling species (the bivalves Mytilus edulis and Hiatella arctica and a solitary ascidian Styela rustica) on their growth rate and mortality were studied through field experiments. The interactions between S. rustica and H. arctica appeared to be the least antagonistic. In contrast, the mussel was the most “aggressive” species with regard to both competitors. It was observed that the ascidians died, because they were intensively braided and gummed up with the byssus threads of the mussels. However, in some cases the intraspecific competition was stronger than the interspecific one.     

Byssus production rate of the White Sea blue mussel <Emphasis Type="Italic">Mytilus edulis</Emphasis> (Linnaeus, 1758) in the presence of metabolites of some hydrobionts

The byssus production of the blue mussel Mytilus edulis L. was studied in the laboratory in the presence of the metabolites of the following animals: a predator (a starfish Asterias rubens L.) and several species competing with the mussel in White Sea fouling communities (a bivalve Hiatella arctica L. and a solitary ascidian Styela rustica L.). The byssus threads and attachment plaques produced by each mussel per day were counted. The number of byssus threads and plaques was smallest in pure sea water and in the presence of metabolites produced by conspecific individuals.     

Fouling Communities of Hydrotechnical Constructions in Nakhodka Bay (Sea of Japan)

The fouling of mooring facilities in Nakhodka Bay, Sea of Japan, has been studied. The main fouling communities have been distinguished dominated by green algae Enteromorpha linza and Ulva fenestrata, brown algae Laminaria japonica and Costaria costata, a hydroid Obelia longissima, a polychaete Pseudopotamilla occelata, cirripede barnacles Balanus crenatus and Semibalanus cariosus, a bivalve mollusk Mytilus trossulus, and an ascidian Halocynthia aurantium. The naturalization of some species-invaders in the fouling of mooring facilities in Nakhodka Bay has been registered, namely hydroids Laomedea flexuosa and Clytia languida, a polychaete Pseudopotamilla occelata, and a bryozoan Bowerbankia gracilis.     

A comparative study of the pumping activity of some fouling animals from the White Sea

The pumping activity of the bivalve Mytulus edulus, Hiatella arctica, and solitary ascidian Styela rustica from the White Sea has been studied. The dependence of the pumping activity on the mass and size of animals has been analyzed. It was shown that the mussel is the most active filter-feeder, that the least value of the pumping activity has been registered for H. arctica, and that S. rustica occupies an intermediate position according this index. The results obtained are examined in connection with the competitive relationship of the animals in the fouling community.     

Effect of excretory-secretory products of some fouling organisms on settling and metamorphosis of the larvae of <Emphasis Type="Italic">Styela rustica</Emphasis> (Ascidiae)

The effect of the excretory-secretory products of some fouling animals on the settling and metamorphosis of larvae of the solitary ascidian Styela rustica was assessed. The substances secreted by the sponge Halichondria panicea stimulated settling of larvae, but concurrently blocked their metamorphosis. The excretory-secretory products of the mussel Mytilus edulis and the ascidian Molgula citrine did not affect settling of the S. rustica larvae but impeded their subsequent development. Water conditioned by the bivalve Hiatella arctica, stimulated settling and, apparently, metamorphosis of the larvae of S. rustica. The chemical substances produced by adult individuals of S. rustica facilitated settling of conspecific larvae but slightly delayed their metamorphosis.     

Subtidal Underwater Rock Communities of the White Sea: Structure and Interaction with Bottom Flow

The taxonomic composition and quantitative characteristics of typical subtidal rock communities of the White Sea (biocenosis of Laminaria and biocenosis of Lithothamnion and the sponges Polymastia arctica) were studied. The hydrodynamic activity of the near-bottom water layer within and beyond the communities was estimated using the plaster structures technique. A comparative analysis of our results and the literature data revealed three main types of interactions between the benthic community and the water flow: turbulation of the water flow (mussel banks), smoothing of turbulent pulsations (Laminaria community), and an insignificant effect on the water flow (community of Lithothamnion and sponges).     

Effect of the excretory-secretory products of some marine invertebrates on byssus production of the blue mussel <Emphasis Type="Italic">Mytilus edulis</Emphasis> (Bivalvia: Mytilidae)

Under laboratory conditions, we investigated byssus production in the blue mussel Mytilus edulis, as affected by the excretory-secretory products (ESPs) of the mussel itself and some marine invertebrates: the predatory starfish Asterias rubens, and organisms competing with mussels in White Sea fouling communities—a bivalve Hiatella arctica, the solitary ascidians Styela rustica and Molgula citrine, and a sponge Halichondria panicea. The number of attachment disks produced by a mussel per day and the thickness of byssal threads were estimated. Excretory-secretory products of H. arctica and M. citrine had no effect on the number of attachment disks, while ESPs of S. rustica, H. panacea and A. rubens stimulated mussels to produce attachment plaques. The activity of the mussel was slightly increased at low levels of its own ESPs in seawater. The thickness of byssal threads decreased with an increase in the ESPs of mussels in seawater, but it increased in experiments with the ESPs of any other species tested.     

Dynamics of a biofouling community in finfish aquaculture: a case study from the South Adriatic Sea

Biofouling is one of the challenges that can strongly affect the finfish farm economy. Although several studies on biofouling in aquaculture have been conducted in the Mediterranean Sea, they focused on specific taxa or were limited to a particular period of sampling. The present study investigated for the first time the development, composition and variation in a biofouling community in a finfish farm with immersion time, season and depth. The results indicate that all these factors influence biofouling succession and recruitment. Moreover, the species that had a crucial role in structuring the community and in the farm cleaning activities were the ascidian Styela plicata and the bivalve Mytilus galloprovincialis. Compared with the literature data, the results highlight the heterogeneity in the composition of the biofouling present in the Mediterranean Sea. Moreover, such knowledge of the biofouling community could provide important information about management efforts and the costs that farmers will face when siting new fish farms.     

Modulating role of lipids and their fatty acids in adaptation of the White Sea mussels <Emphasis Type="Italic">Mytilus edulis</Emphasis> L. to environmental salinity change

Role of lipids and fatty acids (FA) in littoral and sublittoral White Sea mussels Mytilus edulis L. was studied at various stages of reproductive cycle in the phenotypic adaptation (acclimation) to changes of the sea water salinity. The obtained data indicate differences in the mussel lipid and fatty acid spectra, which are connected both with their location (littoral or sublittoral) and with the spawning period stage (3b—release of gametes or 3c—resorption of residual sex products). Lipids and FA of both mussel groups respond to the salinity changes to the greater degree at the 3b than at the 3c stage. In the littoral mussels at the 3b and 3c stages there were revealed differently directed changes in the content of membrane lipid—cholesterol—and in the cholesterol: phospholipids ratio. In the sublittoral mussels that are less adapted to extreme action of abiotic factors, more significant changes were found in the lipid and FA compositions.     

Salinity-stress-induced changes in the resistance of embryos of the White Sea herringClupea pallasi marisalbi to freshwater

The short-term effects of extremely low salinity (3‰) on the resistance of White Sea herring to freshwater were studied. An increasing resistance of the embryos against a sudden drop in salinity, and a decreasing resistance against gradual changes in salinity (to 0‰) was revealed.     

Recent changes in the contributions of river Rhine and North Sea to the eutrophication of the western Dutch Wadden Sea

From 1955 to the mid 1980s the loads of both nitrogen and phosphorus from the river Rhine to the Dutch coastal area, the Wadden Sea included, increased. Since 1985 the phosphorus loads has decreased significantly, while the nitrogen load remained about the same.Annual primary production in the western Dutch Wadden Sea has increased fromc. 40 g C m^–2 (1950) to 150 (mid 1960s) and over 500 g C m^–2 (1986). The biomass of macrozoobenthos has more than doubled since 1970. Simultaneously, the meat yield of cultured blue mussels (Mytilus edulis), has increased since the 1960s. Previously, it was indicated that the increase in primary production of the phytoplankton over the period 1950 to 1986 was stimulated by the load of dissolved inorganic phosphate from Lake IJssel, a reservoir supplied by Rhine water. Since 1990, however, primary production has been higher than was expected from decreased phosphate loads from Lake IJssel. It is argued that this lack of response may have been caused by increased concentrations of dissolved inorganic phosphate at sea originating from increased inflow from a.o. the Strait of Dover, which compensate for the decrease in phosphate from the rivers, possibly in combination with a significant improvement of the light conditions of the water in the Wadden Sea.     

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.     

Fauna associated with detached kelp in different types of subtidal habitats of the White Sea

The fauna, associated with Laminaria and other largebrown macroalgae was studied by using SCUBA anddredging in two different types of underwater habitatsof the White Sea.In shallow water fjords and bays, with a depth of nomore than 30–40 m, detached kelp (mainly Laminaria saccharina, L. digitata and Alaria esculenta) formed large accumulations. One ofthese benthic accumulations, which has existed morethan 20 years, was studied. It covers about2000 m², and is about 2 m thick. The upper layerof the accumulation of fronds is characterized by highturbulence and is well aerated. The lower layer ischaracterized by anoxic conditions. Mats of sulphurbacteria were not observed, although fronds in themiddle layer were covered by layers of cyanobacteria.About 50 species of macroinvertebrates were found,mainly species that are normally associated withliving kelp, such as the detritivorous species Ophiura robusta and Gammarus oceanicus, and fewspecies that are specific inhabitants of organic-richbiotopes in the White Sea such as Capitellacapitata, Ophryotrocha irinae and Nebaliabipes. It was remarkable that in the shallow waterbasins of the White Sea, the process of decompositionof brown algae in the sublittoral takes place withoutsea urchins, and no other macrofaunal form plays anecological role in the mechanical breakdown of theplant substratum, even not in the large accumulationsof detached kelp.Along the open rocky shoreline, communities associatedwith dead detached kelp were situated at a depth of60–90 m, 40–50 m below the belt of living kelp. Inthis deep zone, no macroinvertebrates typical of thekelp community in the photic zone were found. Duringthe passage from the shoreline to the deeper benthiccommunity, where sea urchins were dominant, all plantdebris became fragmented. These deeper benthiccommunities appeared to be the zone for decompositionof the detached kelp.     

Production of planktonic crustaceans in the White Sea

This paper analyzes different methods for assessment of the production of marine organisms. The estimated values of annual production and ration of planktonic copepods of the White Sea are 5.0 × 10¹² and 2.2 × 10¹³ kcal, respectively. Comparison of the planktonic copepod ration with phytoplankton production shows that the energy demands of the copepod community are met by consumption of phytoplankton and, probably, also bacterioplankton and detritus.     

On the dynamics of the stocks of blue mussels (Mytilus edulis L.) in the Danish Wadden Sea

As biological basis for the monitoring programme for the commercially exploited stock(s) of mussels (Mytilus edulis L.) in the Danish Wadden Sea, samples of mussels have been collected regularly since 1986, both from sub-tidal and inter-tidal mussel beds. These samples are the basis for the estimation of total biomass. They also provide data on size frequency distributions, which have been analysed for cohort identification resulting in length at age data, which again have been used for estimating parameters (L and K) for the von Bertalanffy growth equation (VBGE) as well as mortality parameters. By applying these in the Beverton & Holt model, estimates of average biomass and annual production (P) of the mussels have been obtained together with possible fisheries yields from the beds. The growth and mortality parameters and the figures for annual production and P/B are compared with figures from other investigations. These analyses have been the basis for annual assessments of the mussel stocks, which again are used in the current management of mussel fishery in the Danish Wadden Sea.     

Life History of the Barnacle Balanus Amphitrite Darwin and Its Role in Fouling Communities of Peter the Great Bay, Sea of Japan

The life history of the barnacle Balanus amphitrite Darwin and its role in fouling communities of Golden Horn Bay (Peter the Great Bay), which is subjected to thermal pollution, were studied. The warm-water B. amphitrite occurs as a common minor species on operational vessels and waterworks in Peter the Great Bay, where it was brought by ocean-going ships operating on Russia–Japan lines. Even in the conditions of the higher temperature regime of Golden Horn Bay, the reproductive season of B. amphitrite is confined to the summer and autumn months. The adult individuals brought by ships in summer produce 2–3 generations of larvae. The development of larvae and their settling on the substrate occurs from August to October within a broad temperature range from 22.5 up to 12°C. Even in the low temperatures of Golden Horn Bay the larvae attain a greater size than those in tropical and subtropical waters. The juveniles have time to reach maturity and to produce their own progeny, but most often they perish with winter drop in the water temperature. It was shown that in Peter the Great Bay there is dependent population of B. amphitrite inhabiting the anthropogenic substrates only in the warm season: water works, idle vessels, and operational offshore vessels. The water temperature is the limiting factor of successful acclimation of that species.     

Reproduction of the colonial hydroid Obelia geniculata (L., 1758) (Cnidaria,Hydrozoa) in the White Sea

Populations of the colonial hydroid Obelia geniculata in the White Sea reproduce asexually by frustule formation. Young medusae appear in the plankton during July and August. The number of medusae rarely exceeds 36 per m³, and the average number varies every year from 0.4 to 10 per m³. The size of medusae is smaller than reported from other regions. The umbrella of the largest recorded medusa was only 0.57 mm in diameter and the specimen had just 35 tentacles. Only a few mature medusae were found during the study. The colonies in the White Sea are epiphytic and grow only on laminarian thalli. At the beginning of July there are no colonies on thalli from the upper subtidal zone. By the end of August, colonies of O.␣geniculata had increased in density to 30 per m². Hydroid recruitment was attributed to active frustule production by colonies living below that zone. The frustules detach from the stems of the hydroids and are found in plankton. Production of frustules on branches occurs continuously during colony growth until water temperatures climb above 0 °C. We found that water temperature in this Arctic environment is generally too low for medusa maturation and planula development in the species. Propagation by frustule formation is the principal means of reproduction in Obelia geniculata within the White Sea, and this phenomenon accounts for the species being a dominant epiphyte on laminarian thalli there.     

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.