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.     

淡水入侵生物沼蛤的污损机制与防污措施研究进展

沼蛤是一种典型的淡水入侵贝类,能够利用其分泌的足丝牢固黏附在多种水下基质表面,引起严重的生物污损问题。沼蛤污损不但影响水生态系统健康,也给水利工程、交通运输、水产养殖等行业带来经济损失,已成为全球水生态系统安全和国民经济重要行业的潜在威胁,相关防污工作亟待开展。欲从根本上解决沼蛤污损问题,一方面需要加强对其基础生物学特性和污损机制的深入解析,另一方面也需要在此基础上研发更加经济、高效、环境友好的防污措施。本文综述了近年来国内外关于沼蛤污损生物学特性、污损机制和防污措施方面的研究进展,尤其是对沼蛤生物污损发生的主要机制如足探测识别、足丝黏附和环境影响等方面进行了总结,也从物理、化学、生物和防污材料等角度阐述了现有的沼蛤污损控制措施并对未来发展方向进行了展望,以期更加深入地理解沼蛤生物污损现象,为揭示其作用机制、制定科学有效的防污措施、维护水生态系统安全提供数据支撑,综述内容对于水下仿生材料研发也具有重要的参考价值。     

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.     

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.     

Community dynamics of intertidal soft-bottom mussel beds over two decades

Macrozoobenthos communities in the North Sea showed pronounced changes over the past decade in relation to an increasing number of invasive species and climate change. We analysed data sets spanning 22 years on abundance, biomass and species composition of intertidal soft bottom mussel beds near the island of Sylt (German Bight) in the Northern Wadden Sea, based on surveys from 1983/1984, 1990, 1993 and from 1999 to 2005. Mussel bed area and blue mussel biomass decreased, and a change in the dominance structure in the associated community comparing 1984 to mid-1990s with the period from 1999 to 2005 was observed. Coverage of the mussel beds with the algae Fucus vesiculosus decreased since the end of the 1990s. Within the study period biomass and densities of the associated community increased significantly. Dominance structure changed mainly because of increasing abundances of associated epibenthic taxa. Apart from the Pacific oyster Crassostrea gigas all other alien species were already present in the area during the study period. Community changes already started before Pacific oysters became abundant. An attempt is made to evaluate effects on the observed changes of decreasing mussel biomass, ageing of mussel beds, decreasing fucoid coverage and increasing abundances of invader. All four factors are assumed to contribute to changing community structure of intertidal mussel beds.     

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.     

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.     

Mechanisms of successional dynamics: Consumers and the rise and fall of species dominance

Insight into potential mechanisms of succession following disturbance to an ecological community can be gained by considering processes that contribute to the rise (colonization, interactions with established species) and demise (differential mortality) of specific stages within the successional sequence. Most successional theories focus on the rise to dominance, assuming demise is the result of competition, but other factors can cause differential mortality among species, including physical disturbance, senescence, and consumers. In rocky intertidal communities on the coast of Washington state, USA, gaps in mussel beds exhibit a succession from predator-susceptible to predator-resistant species following disturbance, suggesting that differential consumption by mobile species may play an important role in the demise of early succession species and the eventual dominance of the mussel Mytilus californianus. Experimental manipulation of a dominant species earlier in succession, the blue mussel Mytilus trossulus, demonstrated that this species inhibits the invasion of M. californianus in the absence of predators. Experimental manipulation of predatory snails (Nucella emarginata and Nucella canaliculata), which feed heavily on M. trossulus but not M. californianus, greatly increased the rate at which M. californianus invaded gaps. These results and those of other studies indicate that consumers frequently have important effects on the dynamics of succession in benthic marine systems, and might also play a role in other settings.     

Susceptibility of larval dragonflies to zebra mussel colonization and its effect on larval movement and survivorship

Colonization by the zebra mussel, Dreissena polymorpha, was quantified for five dragonfly species that differed in size and larval habits in a Michigan lake. Both larger size and a non-burrowing habit independently increased susceptibility to colonization. In 2005, over 50% of the final instars of the sprawlers Didymops transversa and Hagenius brevistylus were colonized, as well as younger instars. Rarely colonized were Progomphus obscurus and Dromogomphus spinosus, whose larvae burrow under sand, and the sprawler Epitheca princeps, whose final instars were lightly covered with sand. Hagenius larvae that had been preyed upon carried more mussels than those dying of other causes. More generally, mussel attachment decreased the probability that sprawlers left the water to emerge, the distance that some species traveled before emerging, and the ability of an overturned sprawler to right itself. On average, final instars of Didymops and Hagenius remaining in the water carried three times as many mussels as individuals known to emerge. Compared to uncolonized individuals, Epitheca and Progomphus with mussels emerged closer to the water line. Among colonized Didymops, the distance traveled on land before emerging decreased with increasing mussel load. Of the colonized Didymops that could right themselves, righting time increased with mussel load. Because the two common species of sprawlers were disproportionately colonized, and mussel attachment decreased their chances of emerging, our results suggest that D. polymorpha has the potential to affect the community structure of this guild of aquatic and terrestrial predators. Handling editor: R. Bailey     

Potential Impact of Filter-feeding Invaders on Temperate Inland Freshwater Environments

Since the 1990s, biological invasions have captured the attention of the scientific community as an important element of global change and a major threat to biodiversity. The inland waters of South America provide two examples of biological invasions. This review examines bivalve invasions in South America, summarizes the research results for two species, the Asian clam (Corbicula fluminea) and the golden mussel (Limnoperna fortunei), and suggests further studies. The rapid expansion of invasive bivalves into these environments involves significant changes. Until now, C. fluminea, the Asian clam, did not produce generalized macrofouling in the Neotropical region, as is common in the Holarctic region. However, the first specific cases of macrofouling by C. fluminea were recently detected in heat interchangers of power stations in Brazil. On the other hand, L. fortunei is provoking new economic impacts in South American freshwaters through macrofouling. Before the invasion by the golden mussel, macrofouling was recorded only in the marine and estuarine environments of the Neotropical region. The impact caused by invasive bivalves in this region is not only economic, however. Rapid changes in the benthic community, favoring the presence of Oligochaeta and Hirudinea, as well as the displacement of native species of mollusks, are among the problems related to the presence of the golden mussel. Another issue is the settlement of golden mussels on native bivalves. This bivalve is now a new element in the diet of some native fish species, being the main food item in some cases.     

Fouling of European freshwater bivalves (Unionidae) by the invasive zebra mussel (Dreissena polymorpha)

1. The zebra mussel (Dreissena polymorpha) is well known for its invasive success and its ecological and economic impacts. Of particular concern has been the regional extinction of North American freshwater mussels (Order Unionoida) on whose exposed shells the zebra mussels settle. Surprisingly, relatively little attention has been given to the fouling of European unionoids. 2. We investigated interspecific patterns in fouling at six United Kingdom localities between 1998 and 2008. To quantify the effect on two pan‐European unionoids (Anodonta anatina and Unio pictorum), we used two measures of physiological status: tissue mass : shell mass and tissue glycogen content. 3. The proportion of fouled mussels increased between 1998 and 2008, reflecting the recent, rapid increase in zebra mussels in the U.K. Anodonta anatina was consistently more heavily fouled than U. pictorum and had a greater surface area of shell exposed in the water column. 4. Fouled mussels had a lower physiological condition than unfouled mussels. Unlike tissue mass : shell mass ratio, tissue glycogen content was independent of mussel size, making it a particularly useful measure of condition. Unio pictorum showed a stronger decline in glycogen with increasing zebra mussel load, but had a broadly higher condition than A. anatina at the time of study (July). 5. Given the high conservation status and important ecological roles of unionoids, the increased spatial distribution and fouling rates by D. polymorpha in Europe should receive more attention.     

The impact of the sea anemone Actinothoe sphyrodeta on Mytilus galloprovincialis mussel cultivation (Galicia,Spain)

Abstract

Marine mussel aggregations act as a substratum and refuge for many fouling species. Mussel cultivation in Galicia, Spain, is carried out on hanging ropes in subtidal systems. The fauna associated with this cultivation includes a large number of invertebrates that compete for space or food with the mussels, or use their clusters as a refuge from predators or water turbulence. Outbreaks of the epibiont anemone Actinothoe sphyrodeta have been reported in cultivated Galician mussels since 2013, but their impact has not been investigated rigorously. Here, the temporal and spatial variability of Actinothoe sphyrodeta on mussel shells throughout one year is presented. Sampling of mussel size, weight and byssus attachment strength allowed mussel tenacity (attachment strength relative to size) to be calculated. A higher presence of Actinothoe sphyrodeta correlated with lower mussel tenacity and greater biomass losses, suggesting that this species could be an economically important biofouling component.     

Mechanisms and impact of differential fouling of the zebra mussel Dreissena polymorpha on different unionid bivalves

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Fouling community of the red king crab, <Emphasis Type="Italic">Paralithodes camtschaticus</Emphasis> (Tilesius 1815), in a subarctic fjord of the Barents sea

We examined the species composition of red king crab (Paralithodes camtschaticus) fouling communities in Dolgaya Bay, a small fjord of the Barents Sea, in August 2005 and 2006. In total, there were 13 species observed on 301 crabs collected from water depths of 5–90 m. Barnacles (Balanus crenatus; prevalence 42.9%) and blue mussels (Mytilus edulis; 11.6%) were the most common epibionts, while amphipods (Ischyrocerus commensalis) were the most common symbionts (28.6%). Infestation rates in Dolgaya Bay were different from those in an “open” area of the Barents Sea (Dalnezelenetskaya Bay), probably due to differences in hydrodynamic conditions. Differences in infestation prevalence and intensity were detected neither between male and female crabs nor between crabs collected at 5–35 m versus 90 m depths. Prevalence of common fouling species increased with host size. Amphipods I. commensalis colonized the carapace and limbs in Dolgaya Bay less frequently than in Dalnezelenetskaya Bay, probably due to interspecific competition with barnacles occupying the dorsal parts of the host. Juvenile barnacles and mussels dominated the fouling communities on the crabs. The age of barnacles did not exceed 2–4 months. However, the presence of 4-year-old mussels suggests that these older mollusks have been directly transferred from mussel beds to the hosts. Our results indicate that colonization by epibionts and symbionts is generally not disadvantageous for the crab hosts, except for some possible negative impacts of amphipods occupying the gills.     

Indigenous microflora and opportunistic pathogens of the freshwater zebra mussel, Dreissena polymorpha

Freshwater fouling invertebrate zebra mussels (Dreissena polymorpha) harbor a diverse population of microorganisms in the Great Lakes of North America. Among the indigenous microorganisms, selective species are opportunistic pathogens to zebra mussels. Pathogenicity to zebra mussels by opportunistic bacteria isolated from the mussels was investigated in this study. Among the more than 30 bacteria isolated from temperature-stressed mussels, Aeromonas media, A. veronii, A. salmonicida subsp. salmonicida, and Shewanella putrefaciens are virulent pathogens to juvenile zebra mussels. Inoculation of a bacterial concentration of A. media, A. salmonicida subsp. salmonicida and S. putrefaciens at 10⁷ cells per zebra mussel resulted in 100% mortality within 5 days, and only 64.9% for A. veronii. In contrast, mortality was less than 12.3% following inoculation of a sterile phosphate buffer solution as a control. In addition, mortality was dependent on the size of the pathogen population used in inoculation and the incubation temperature, indicating the close relationship between the bacterial population and subsequent death. On the mussel tissue, a dense microbial population was evident from the moribund mussels viewed with Scanning Electron Microscope (SEM). Opportunistic bacteria invaded and destroyed the D. polymorpha tissue after 7 days of incubation when the bacterial inoculation was larger than 10⁵ per zebra mussel. Our results suggest that mussels are reservoirs of opportunistic pathogenic microorganisms to aquatic organisms and humans and a better understanding of the microbial ecology of the mussels will provide insights to the possible health hazards from these microorganisms.     

Seasonal settlement and succession of fouling communities in Kalpakkam, east coast of India

Seasonal distribution and community succession of macrofoulants were studied using concrete panels in the coastal waters of Kalpakkam, east coast of India, for a period of two years. The panels were suspended at 1 m, 4 m and 7 m depths and categorised into short-term and long-term exposures. A high total of 105 fouling taxa were recorded. The major fouling organisms observed were hydroids, barnacles, mussels, anthozoans and ascidians. Considerable faunistic and biomass variations were noticed both with respect to season and depth. The month of panel exposure had a significant influence on the succession of fouling communities. On the short-term panels, the maximum fouling biomass was 64 kg m^–2 in 30 days at 4 m depth, whereas on the long-term panels, it was 250 kg m^–2 after 216 days at 4 m depth. A comparison with the biomass values reported from elsewhere shows that biomass build-up in Kalpakkam coastal waters is one of the highest ever reported. Such a very high biomass accumulation is due to the extremely dense settlement of mussels, especially the green mussel,Perna viridis (L).     

Seasonal variations in the fouling diatom community structure from a monsoon influenced tropical estuary

Seasonal variations in the fouling diatom community from a monsoon influenced tropical estuary were investigated. The community composition did not differ significantly between stainless steel and polystyrene substrata due to dominance by Navicula spp. The experimental evidence suggests that Amphora, which is a dominant fouling diatom in temperate waters, ultimately dominates the community in tropical regions when conditions are favourable. These observations reveal that a faster onset of macrofouling interferes with the microfouling community wherein the faster recruiters that have a higher abundance in ambient waters, dominate the community. Seasonal variations were observed in the fouling diatom community. Navicula delicatula dominated during the post-monsoon and tychopelagic diatoms (Melosira and Odontella) were observed during the monsoon. Low diatom abundance was recorded during the pre-monsoon season. The results indicate that although the fouling diatom community composition does not vary between substrata, there is a seasonal change in the community depending on the physical, chemical and biological interactions.     

Facilitation and competition between invasive and indigenous mussels over a gradient of physical stress

The interactions between invasive exotic and indigenous species can have profound harmful effects on the recipient community; however, not all such interactions are negative. Facilitation is increasingly recognised as important in shaping natural communities and is believed to vary under different conditions. Earlier studies have shown that the indigenous intertidal mussel Perna perna initially facilitates survival of the invasive Mytilus galloprovincialis in the low mussel zone by providing protection against waves, but later excludes M. galloprovincialis through interference competition for space. Here, we examined interactions between these species in the mid and upper mussel zones, moving mussels to experimental plots in different combinations of densities and species. Mussels were left on the shore for more than a year and treatment effects on mortality, shell length and condition were compared. In the high zone, treatment had no effects and P. perna showed greater mortality than M. galloprovincialis, indicating that its exclusion from the high shore is due to emersion stress. In the mid zone, treatment had no significant effects on M. galloprovincialis, but multiple comparisons among treatments involving P. perna showed that facilitation occurred. P. perna survived better at higher densities, but survived even better when mixed with the physiologically more tolerant M. galloprovincialis. Length data indicated both inter- and intraspecific competition for P. perna in the mid zone. Whereas facilitation occurs strongly in the low zone (P. perna facilitates M. galloprovincialis) and weakly in the mid zone (M. galloprovincialis facilitates P. perna), the lack of facilitation in the high zone suggests that the probability of facilitation is not linearly linked to increasing physical stress. Instead it is likely to be hump shaped: relatively unimportant under conditions that are benign for a particular species, significant under more severe conditions, and overridden by physical stress under very harsh conditions.     

Effects of the exotic invader Musculista senhousia on benthic communities of two Mediterranean lagoons

The impact that the alien mussel Musculista senhousia has on benthic biodiversity and community structure was investigated in two Italian transitional environments, where the species was particularly abundant: the Sacca di Goro (Adriatic Sea), and the Rio Padrongiano Deltaic area (Sardinia, Tyrrhenian Sea). Sampling campaigns were conducted by sampling mussel beds from each of two different patch size classes (small: <1.0 m², and large: >10.0 m²), and the surrounding bare sediments. At both study sites, mussel density varied greatly with time within and between patches. Despite the huge difference in benthic species complement at Goro and Padrongiano (33 and 115 taxa, respectively), at both sites the presence of mussel patches had a gross positive effect on the benthic community. At Goro, many trophic guilds showed a positive relationship with mussel abundance. At Padrongiano, a continuous and directional process of patches colonization (both large and small) through time seemed evident, while at Goro the process was interrupted by high-summer mortalities. At both study sites, M. senhousia played a double role: (1) as secondary substratum, enhancing the environmental structural complexity, and (2) as densely aggregated, living organisms, thus giving rise to a number of possible interactions with other invertebrates. The outcome, however, was strictly dependent by the level of anthropogenic stress, in this case water oxygen deficiency, which was different at the two study sites. Guest editors: A. Razinkovas, Z. R. Gasiūnaitė, J. M. Zaldivar & P. Viaroli European Lagoons and their Watersheds: Function and Biodiversity     

Process of invasiveness among exotic tunicates in Prince Edward Island, Canada

Over the past decade, four exotic tunicates (Styela clava, Ciona intestinalis, Botrylloides violaceus and Botryllus schlosseri) have been reported in the Brudenell estuary in Prince Edward Island (PEI), Canada. Styela clava was the first exotic tunicate to arrive in 1997, rapidly establishing, spreading, invading, and eventually becoming a nuisance in several estuaries of PEI. In the Brudenell estuary, S. clava remained the only exotic nuisance tunicate until 2003. In the fall of 2004, the vase tunicate C. intestinalis, was reported in low abundance, followed by the two colonial species, B. schlosseri and B. violaceus, reported in the spring of 2005. The abundance of C. intestinalis rapidly increased post-introduction, eventually replacing S. clava as the foremost nuisance species on mussel farms in the estuary. To date, C. intestinalis continues to colonize this estuary at epidemic proportions, resulting in the continuing drop of S. clava abundance. The current abundance of C. intestinalis is estimated at 5 cm⁻², which is similar to S. clava abundance at its height in 2003. The 2006 abundance of S. clava is estimated to have fallen to near 0 cm⁻². The dominance of C. intestinalis as a fouling organism on mussel farms is considered a serious threat to this aquaculture industry, mainly due to its unmanageable weight. The process of the detection, establishment, invasiveness, and eventual rise to nuisance level of exotic tunicates in the Brudenell River is presented.