Long-Term Changes in Shallow-Water Fouling Communities of the White Sea

The replacement of an ascidian (Styela rustica L.) fouling community by a blue mussel (Mytilus edulis L.) community was described for the White Sea. The alternation of populations of these two species takes place in fouling communities developing in the upper 3- to 5-m layer of water. The life span of each type of fouling probably depends on interannual climate fluctuations.     

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.     

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.     

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.     

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.     

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.     

Testing the hypothesis of tolerance strategies in Hiatella arctica L. (Mollusca: Bivalvia)

The physiological and biocenotic optima of Hiatella arctica L. inhabiting shallow water fouling communities of the White Sea were compared. The biomass and proportion of H. arctica in communities were used for the estimation of biocenotic optima or community success. The physiological state of populations was assessed by means of the fluctuating asymmetry. The fluctuating asymmetry of H. arctica was calculated using the valve weights. It was determined that the shell of H. arctica possesses a slight directional asymmetry, the right valve usually being larger (and heavier) than the left one. The relationship between fluctuating and directional asymmetries is discussed. High biomass and proportion of H. arctica in the community generally correspond with high levels of fluctuating asymmetry. Thus, a discrepancy between physiological and ecological optima is observed, which is recognised as being characteristic of a tolerance strategy. However, in the absence of pressure from major filter-feeding competitors such as the mussel Mytilus edulis L. and the solitary ascidian Styela rustica L., populations of H. arctica possess a high biomass and low levels of fluctuating asymmetry. It appears that H. arctica prefers to inhabit shelters or depressions, or to be covered by other organisms. Populations of H. arctica existing in the absence of shelter had extremely high levels of fluctuating asymmetry despite the absence of filter-feeding competitors. Thus, the strategy adopted by H arctica L. in the investigated upper 5 m layer of water in the White Sea can be described as a stress- and competitor-tolerant strategy.     

Variability of morphological characteristics in <Emphasis Type="Italic">Oithona similis</Emphasis> (Copepoda: Cyclopoida) in the White Sea

The spatial variability of the body size and morphological structures of the planktonic copepod Oithona similis in the White Sea was studied. From north to south, there was a tendency for the male cephalothorax length to decrease. In moving from the central to the northern part of the sea and from the eastern to the western part, the relative antennule length in both sexes increased. The average body size of copepods was positively and the absolute (and relative) lengths of the antennules were negatively correlated with water salinity. The morphological variations have adaptive significance and are determined by environmental factors.     

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.     

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.     

Tolerance and Resistance in Gastropod Mollusks Hydrobia ulvaeand H. ventrosafrom the White Sea to Abiotic Environmental Factors

Tolerance to salinity changes and resistance to desiccation, fresh water, and freezing were studied in two hydrobiid species from the White Sea, Hydrobia ulvae(Pennant) and H. ventrosaMontagu. It was shown that H. ventrosahas a greater tolerance to low salinities in the range of 6–10 but is less tolerant to high salinities (35–45). The interspecies differences in low salinity tolerance persisted after the acclimation of snails to 20, 16, and 12. A comparison of survival in fresh water and under desiccation conditions suggests that H. ulvaeis more resistant to these factors. In fresh water, LT₅₀was 30 and 60 days for H. ventrosaand H. ulvae, respectively. Under desiccation conditions, LT₅₀was 6 and 25 days for H. ventrosaand H. ulvae, respectively. At subzero temperatures, H. ventrosawas shown to be a superior survivor. Within the same species of mud snail, the detrimental effect of freezing depended on the salinity: survival decreased with decreasing salinity. These data suggest significant differences in the mechanisms of resistance and tolerance to abiotic environmental factors between the two species. Despite the partial overlapping of their ecological niches, the interspecies differences may play an important role in the distribution and spatial structure dynamics of coexisting populations of these species in the White Sea region.     

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.     

Life history characteristics of the clam<Emphasis Type="Italic"> Mya arenaria</Emphasis> in the White Sea

Dynamics of Mya arenaria beds in two bights of the Chupa Inlet (Kandalaksha Bay, White Sea) were studied on a long-term basis. Observations were carried out at 1– to 3-year intervals from 1979 up to 1999. The studied soft-shell clam beds were characterised by a substantial instability of age structure. Since 1988, only one year-class has dominated in the beds while other generations have been scarce and recruitment was not observed. This pattern of Mya bed dynamics was related neither to interannual environmental changes nor to differential reproduction success or predation effects in the benthic assemblages. Favourable conditions for spat formation in 1988 (low abundance of other M. arenaria generations), as well as for juvenile survival during the following winter, resulted in high abundance of juveniles in both investigated locations in 1989. The mortality rate (μ) in this 1988 generation varied throughout the period of investigation and was related to age. The mortality level decreased for the first 2–4 years of the life cycle, then stabilised for the next 3–4 years, and eventually increased in subsequent years. Overall μ values ranged from 0 to 1.68 year^–1. The oldest specimens observed were 17 years old and had a maximum shell length of 79 mm. Significant differences in average growth rates were observed between molluscs of different locations. Communicated by H.-D. Franke     

The Polychaete Polydora limicola in Benthic Communities of the Black Sea

Many papers dealing with nonindigenous species in the Black Sea do not mention the polychaete Polydora limicola Annenkova, a new invader in the Pontic-Azov Basin. In this connection, we will analyze here the taxonomic status of this species and discuss the history of its introduction and distribution in the northwestern Black Sea and related coastal lagoons. It is shown that P. limicola is now one of the most common species of the benthos in the Black Sea, especially in estuarine areas and coastal lagoons. The naturalization of this species resulted in changes in the structure of some bottom biocenoses of this region.     

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.     

Iceberg Disturbance and Successional Spatial Patterns: The Case of the Shelf Antarctic Benthic Communities

High-latitude, shelf Antarctic benthic communities are highly diversified and structured, dominated by benthic suspension feeders, and are subject to major natural disturbances. This study focuses on spatial patterns of the Antarctic benthos emphasizing the succession process after iceberg disturbance. For this purpose, underwater photographs (1 m² each) from the southeastern Weddell Sea shelf (<300 m depth) were analyzed using techniques from the field of landscape ecology. Here, we examine measurements of spatial patterns (landscape indices) to describe changes in structural patterns along successional stages on these Antarctic benthic communities. We show a gradual separation from the early to older stages of succession based on sessile benthic cover area, size, shape, diversity, and interspersion and juxtaposition indices. Conceptually, the results describe a gradient from samples belonging to first stages of recovery with low cover area, low complexity of patch shape, small patch size, low diversity and patches poorly interspersed to samples from later stages with higher values of these indices. Cover area was the best predictor of recovery. We conclude that a variety of factors affect the observed successional sequences of Antarctic shelf benthic communities after iceberg disturbance, including the existence and dispersal abilities of propagules, growth rates, and competition between species. Overall, changes in the magnitude, frequency, and duration of disturbance regimes and alterations of ecosystem resilience pose major challenges for conservation of Antarctic benthos. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Dynamics of a bloom of halophilic archaea in the Dead Sea

After a period of more than ten years in which bacterial and algal community sizes were extremely small, a dense bloom of halophilic archaea developed in the upper 5–10 m of the Dead Sea water column in the summer of 1992. The development of this bloom followed a dilution of the upper water layer by winter rainfloods, which enabled the development of a short-lived dense bloom of the unicellular green alga Dunaliella parva. The dense archaeal community (up to 3.5 × 10⁷ cells m1^–1 in June 1992) imparted a red coloration to the Dead Sea, due to its high content of bacterioruberin. Bacteriorhodopsin was not detected. High levels of potential heterotrophic activity were associated with the bloom, as measured by the incorporation of labeled organic substrates. After the decline of the algal bloom, archaeal numbers in the lake decreased only little, and most of the community was still present at the end of 1993. The amount of carotenoid pigment per cell, however, decreased 2–3-fold between June 1992 and August 1993. No new algal and archaeal blooms developed after the winter floods of 1992–1993, in spite of the fact that salinity values in the surface layer were sufficiently low to support a new algal bloom. A remnant of the 1992 Dunaliella bloom maintained itself at the lower end of the pycnocline at depths between 7 and 13 m (September 1992–August 1993). Its photosynthetic activity was small, and very little stimulation of archaeal growth and activity was associated with this algal community.     

Free-living nematodes of the genus <Emphasis Type="Italic">Alaimella</Emphasis> Cobb 1920 (Nematoda: Leptolaimidae): A description of <Emphasis Type="Italic">A. macramphis</Emphasis> sp. n. from the White Sea and a revision of the genus

Alaimella cincta Cobb 1920 and Alaimella macramphis sp. n. are described and illustrated. Both the species were first recorded for the White Sea in northern Russia. A. cincta occurs in the shells of the agglutinated foraminiferan Reophax curtus, as well as freely in bottom sediments. A. macramphis sp. n. is described from a single male sampled from bottom sediments. A. macramphis sp. n. is distinguished from two previously known species of Alaimella (A. cincta Cobb 1920 and A. truncate Cobb 1920) by having a longer body, longer cephalic setae, and a wide amphid equal to the respective body diameter. The new species also differs from A. truncate by the distinct striation of the cuticular annulations. The Alaimella species are additionally characterized by having a posterior glandular widening of the esophagus. An emended diagnosis of the genus Alaimella Cobb 1920 and a key for species identification are provided.     

The Distribution of Nearshore Fishes in Kelp and Eelgrass Communities in Prince William Sound, Alaska: Associations with Vegetation and Physical Habitat Characteristics

The nearshore (less than 20m depth) demersal fish community in Prince William Sound, Alaska, is dominated by Pacific cod, Gadus macrocephalus, pricklebacks (mostly Arctic shanny Stichaeus punctatus), gunnels (mostly crescent gunnels Pholis laeta), a variety of greenlings (Hexagrammidae) and sculpins (Cottidae). During summer, the spatial distribution of fishes, over scales of 100's of m to 10's of km, varied by habitats characterized by different vegetation types. Juvenile Pacific cod and greenlings were numerically dominant in eelgrass, Zostera marina, beds. Pricklebacks and sculpins were dominant in areas with an understory of the kelps Agarum cribrosum and Laminaria saccharina. Greenlings and sculpins were the most abundant demersal fishes in more exposed sites with a canopy of Nereocystis luetkeana and an understory of L. bongardiana. Measured habitat variables, including vegetation type, slope, vegetation biomass, and substratum type, explained a significant proportion of the variation in the presence or absence of most fishes. The relative importance of different habitat characteristics varied between taxonomic groups of fishes. Vegetation type explained a significant proportion of variation for cod, rockfishes, and ronquils. Juvenile cod were closely associated with eelgrass, while rockfish and ronquils were associated with kelps. Pricklebacks and rockfishes were more frequently observed on steeply sloped shorelines, while ronquils were more often found at sites with higher biomass of vegetation. Within A. cribrosum habitats, more greenlings and sculpins were present at sites where algal biomass was higher. Also, sculpins were more abundant in deeper water and gunnels were more abundant in shallow water within this habitat. These associations may not have been causative. However, evidence suggests that some differences between fish communities in eelgrass and Agarum beds may have been causally related to vegetation characteristics. The possible roles of different vegetation types as refugia from predators or as sources of prey are discussed.