Oosterschelde estuary (S.W. Netherlands): a self-sustaining ecosystem?

The Oosterschelde estuary has a special position among the Dutch North Sea estuaries. With relatively unpolluted water and high transparency it combines high biotic diversity and high primary and secondary production. Before 1970 a major part of the water of the rivers Rhine and Meuse flowed into the Oosterschelde. The building of a permeable storm-surge barrier (1986) decreased the exchange between Oosterschelde and North Sea. The construction of two additional dams (1987) reduced the very limited fresh water discharge on the estuary even further. The total effect of these changes was a decrease of the nutrient levels in the water column.The estuary is used intensively for culture of mussels and fishery of cockles. Both zoobenthos groups use together up to 60% (in the western part) of the organic food available in the Oosterschelde estuary. The storm-surge barrier resulted in reduced current velocities and increased sedimentation, accompanied by higher transparency, hence accelerating primary production. This phenomenon is counteracted by the lowered nutrient concentrations throughout the year.In the seventies a preliminary carbon budget study indicated that a substantial import of organic carbon was required to sustain the ecosystem. Later studies did not confirm this hypothesis.In the present paper three different methods are presented to answer the question, if internal production suffices to sustain the foodweb. At first a steady state model is applied, secondly, calculations with actually measured process rates are carried out and, finally, a dynamic simulation model is used.The conclusion is drawn that, before the construction of the storm-surge barrier, primary production of organic matter by phytoplankton is much more important than import, although the latter can not be neglected as additional food source for mussels, cockles and zooplankton. The simulation model predicts that this conclusion will not have to be changed in the future. The Oosterschelde will remain largely a self-sustaining ecosystem.     

Mussel culture in a changing environment: the effects of a coastal engineering project on mussel culture (Mytilus edulis L.) in the Oosterschelde estuary (SW Netherlands)

To evaluate the effects of a large scale coastal engineering project on the mussel (Mytilus edulis) bottom culture in the Oosterschelde estuary (S.W. Netherlands), mussel growth and production in the period 1980–1990 are studied in relation to food supply and the hydrodynamic conditions. Due to the construction of a storm-surge barrier and two additional dams, the risk that mussels are swept away by high current velocities decreased, resulting in an increase of the area in the Oosterschelde potentially suitable for mussel culture and in food availability now being more important as a limiting factor. For the Oosterschelde, a clear relation between mussel growth, stock sizes, and phytoplankton dynamics has been demonstrated. The meat yield of mussels landed in autumn — which is an index for growth rate — seems to be determined by the phytoplankton production in the preceding summer. In years with dense bivalve stocks, phytoplankton production and meat yields are relatively low. It is concluded that an increase of the mussel biomass cultured can result in a reduction of the primary production and, consequently, in a deterioration of the growing conditions for suspension-feeders in the estuary. This conclusion is supported by model calculations. An expansion of mussel culture in the new Oosterschelde is therefore dissuaded. Apart from primary production and stock sizes, food supply for mussels on culture lots appeared to be controlled by the horizontal advection of phytoplankton between and within the tidal channels. An observed decline in mussel landings from certain areas is attributed to the reduced mixing energy of the estuary in relation to the present distribution of the lots over the estuary. Production figures from the experimental lots, established in 1988 in the newly available areas, demonstrate that the yield of mussels can be enhanced by relaying culture lots towards the areas where the phytoplankton is produced. It is expected that by redistributing the culture lots, without expanding the biomass cultured, the carrying capacity of the Oosterschelde for mussel culture can be maintained.     

Harbour Porpoises Phocoena phocoena in the Eastern Scheldt: A Resident Stock or Trapped by a Storm Surge Barrier?

Coastal protection measures are planned and executed worldwide to combat the effects of global warming and climate change, in particular the acceleration of sea level rise, higher storm surge flooding and extensive coastal inundation. The extent to which these defensive measures may impact coastal and estuarine ecosystems is still poorly understood. Since the building of a storm surge barrier, movement of harbour porpoises Phocoena phocoena in and out of the Eastern Scheldt tidal bay (SW-Netherlands) may be limited. To measure residency, porpoises stranded along the Dutch North Sea coast between 2006 and 2008 were sampled for muscle (n = 102) and bone tissue (n = 118), of which 9 muscle (8.8%) and 12 bone samples (10.2%) were collected from animals stranded within the Eastern Scheldt. Stable carbon (δ¹³C) was analysed to get insight into the habitat use and residency of porpoises in the Eastern Scheldt. Our data showed significantly higher δ¹³C values in the muscle of porpoises stranded within the Eastern Scheldt (µ = −17.7‰, SD = 0.4‰) compared to animals stranded along the Dutch coast (µ = −18.3‰, SD = 0.5‰). This suggests that most porpoises stranded in the Eastern Scheldt foraged there for a longer period. The distinct δ¹³C signature of animals from the Eastern Scheldt was not observed in bone tissue, suggesting a relatively recent shift in habitat use rather than life-long residency of porpoises within the Eastern Scheldt. The high number of strandings within the Eastern Scheldt suggests a higher mortality rate compared to the Dutch coastal zone. Our study indicates that along with other changes in the physical environment, the storm surge barrier may play an important role in determining the residency of porpoises in the Eastern Scheldt, and that the area might act as an ecological trap for porpoises entering it.     

Predator‐induced morphological defences in two invasive dreissenid mussels: implications for species replacement

相似文献   

Temporal and spatial trends in heavy metal concentrations in the marine mussel Mytilus edulis from the Western Scheldt estuary (The Netherlands)

Since the first North Sea Conference (1984, Bremen), all countries bordering the North Sea made commitments to reduce discharge of hazardous substances into the North Sea. From Belgium and The Netherlands, large reductions (upto 90) in heavy metal emissions from land-based sources have been reported between 1985 and 2000. Recently, some studies in the Western Scheldt estuary (WS) have shown that total metal concentration in the water, sediments and suspended particles have decreased compared to levels in the 70s. However, data on aquatic organisms is still very limited and it is therefore difficult to confirm whether the reductions in pollution input and generally improving water quality in the WS have a corresponding impact on the levels of heavy metals in aquatic organisms. The current study measured metal concentrations in the soft tissues of mussels, Mytilus edulis (known to be good indicators of environmental metal pollution) during the period 1996–2002. Spatial (salinity and pollution gradients), temporal and seasonal variations were also studied. Results showed a down-stream decreasing trend for the metals studied (Cd, Co, Cr, Cu, Fe, Mn, Ni, Pb and Zn) during all sampling campaigns. There was also a significant seasonal effect on tissue metal concentrations, with a peak observed around spring in both WS and the nearby less polluted Eastern Scheldt (ES). On temporal trends, a clear drop of metals in mussels was observed in the early 80s coinciding with the start of the efforts to reduce chemical pollution input into the North Sea. Since those early reductions, metal concentrations in mussels generally remained unchanged upto mid 90s. However, in recent times metal concentration in mussels have increased significantly, for example Cd in 2002 was almost 10 times the values in 1983 and similar to levels seen during the peak in the 70s. Other metals also increased in the 90s also reaching levels seen in the 70s. As there is no indication of recent increase in metal input into the estuary, we suggest that increased metal concentrations observed in mussels in recent years especially in the upper estuary are most likely a result of changes in physical and chemical speciation and metal bioavailability. Such changes may be caused by changes in some water quality parameters in the estuary (i.e. increased dissolved oxygen, concentration of organic matter), resulting in conditions that favour releases of sediment-bound metals into the water column. The relationship between metal content and season showed very similar annual profiles in the polluted WS and less polluted ES. Thus, seasonal variations in metal concentrations appear to be largely controlled by biological processes, while total body burdens are dependent on environmental levels and bioavailability.     

Stable musselMytilus edulis beds in the Wadden Sea—They’re just for the birds

Predation by eiders, oystercatchers and herring gulls on natural mussel bedsMytilus edulis was studied in the Königshafen, a sheltered bay in the Wadden Sea. About 15 ha (2.5%) of the Königshafen were covered with mussel patches of a biomass of about 1300 g AFDW m^?2. The biomass on the mussel beds was dominated by old mussels and found to be constant over several years. Birds annually removed 30% of the standing stock. Eiders were by far the most important predators and consumed 346 g AFDW m^?2, followed by oystercatchers with 28 g AFDW m^?2 and herring gulls with 3.6 g AFDW m^?2. Birds consumed a substantial part of the annual production of the mussel beds which was estimated from literature data to be approx. 500 to 600 g AFDW m^?2. As other predators were absent, the production of the mussels was sufficient to sustain the high predation rate by birds. Stable mussel beds form a short and efficient link between primary production and bird predation which is unusual for the Wadden Sea, where the main part of primary food supply is thought to be unavailable for higher trophic levels.     

Secondary production of an intertidal mussel (Mytilus edulis L.) population in the Eastern Scheldt (S.W. Netherlands)

We monitored an intertidal mussel (Mytilus edulis L.) population between June 1981 and June 1982 in the Eastern Scheldt estuary (S.W. Netherlands). Density and biomass of the population remained relatively constant over the study period. The shell length growth was described by a Gompertz growth curve. The parameters of this equation were estimated from a log-log-modified Ford-Walford plot of the growth-ring data. The slope of the relationship between animal weight and shell length is season-dependent, mainly due to the spawning cycle in larger mussels.Secondary production is estimated with the growth rate method. In the calculated growth rates the change in slope of the length-weight relationship is incorporated, as well as differences in length growth rates between summer and winter. Secondary production amounts to 156 g AFDW m^–2a ^–1 (expressed per m² of mussel bank). P:B is 0.50 a^–1. The mussel productivity is probably a limiting factor for the density of overwintering Oystercatchers (Haematopus ostralegus).     

Blue mussels (Mytilus edulis) in the diet of roach (Rutilus rutilus) in outer archipelago areas of the western Gulf of Finland,Baltic Sea

The blue mussel (Mytilus edulis) is one of the key species in the Baltic Sea ecosystem and it is living at the edge of its range in the western Gulf of Finland. Roach (Rutilus rutilus) is a freshwater fish species that has benefited from recent coastal eutrophication and is at present highly abundant in the outer archipelago of the Gulf of Finland. In 2000 and 2001, a total of 516 roach were sampled for diet analysis in three study areas. Shelled molluscs formed over 95% of the diet of roach, blue mussels being the dominant single species. The proportion of this species in the food of roach in the three study areas ranged between 38–61% for smaller roach (<225 mm) and 39–85% for larger (>225 mm) roach, indicating that blue mussel is a highly important food source for roach in outer archipelago areas of the western Gulf of Finland, in contrast to reports from other parts of the northern Baltic Sea. The scarcity of large blue mussels in mussel beds in the easternmost study area was reflected in the lower proportion of blue mussels in the diet of larger roach. However, the growth of roach was not affected by the availability of blue mussels. The twofold differences observed in the annual growth of roach between warm and cold years demonstrated that temperature is an important factor controlling the growth of roach in the western Gulf of Finland.     

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.     

High survival and growth rates of introduced Pacific oysters may cause restrictions on habitat use by native mussels in the Wadden Sea

Pacific oysters Crassostrea gigas (Thunberg, 1793) were introduced to the northern Wadden Sea (North Sea, Germany) by aquaculture in 1986 and finally became established. Even though at first recruitment success was rare, three consecutive warm summers led to a massive increase in oyster abundances and to the overgrowth of native mussel beds (Mytilus edulis L.). These mussels constitute biogenic reefs on the sand and mud flats in this area. Survival and growth of the invading C. gigas were investigated and compared with the native mussels in order to predict the further development of the oyster population and the scope for coexistence of both species. Field experiments revealed high survival of juvenile C. gigas (approximately 70%) during the first three months after settlement. Survival during the first winter varied between > 90% during a mild and 25% during a cold winter and was independent of substrate (i.e., mussels or oysters) and tide level. Within their first year C. gigas reached a mean length of 35-53 mm, and within two years they grew to 68-82 mm, which is about twice the size native mussels would attain during that time. Growth of juvenile oysters was not affected by substrate (i.e., sand, mussels, and other oysters), barnacle epibionts and tide level, but was facilitated by fucoid algae. By contrast, growth of juvenile mussels was significantly higher on sand flats than on mussel or oyster beds and higher in the subtidal compared to intertidal locations. Cover with fucoid algae increased mussel growth but decreased their condition expressed as dry flesh weight versus shell weight. High survival and growth rates may compensate for years with low recruitment, and may therefore allow a fast population increase. This may lead to restrictions on habitat use by native mussels in the Wadden Sea.     

Dynamic measurements of black oystercatcher (Haematopus bachmani) predation on mussels (Mytilus californianus)

Intertidal zone mussels can face threats from a variety of predatory species during high and low tides, and they must balance the threat of predation against other needs such as feeding and aerobic respiration. Black oystercatchers (Haematopus bachmani) on the Pacific coast of North America can depend on the mussel Mytilus californianus for a substantial portion of their diet. Observations suggest that oystercatchers tend to focus on mussels beginning to gape their valves during rising tides, following periods of aerial emersion. We present detailed, autonomous field measurements of the dynamics of three such predation events in the rocky intertidal zone. We measured accelerations of up to 4 g imposed on mussels, with handling times of 115–290 s required to open the shell and remove the majority of tissue. In each case a single oystercatcher attacked a mussel that had gaped the shell valves slightly wider than its neighbors as the rising tide began to splash the mussel bed, but no other obvious characteristic of the mussels, such as body temperature or orientation, could be linked to the oystercatcher's individual prey choice.     

Genetic variability and phylogeographical patterns of a nonindigenous species invasion: a comparison of exotic vs. native zebra and quagga mussel populations

There have been few investigations of the number of founding sources and amount of genetic variability that lead to a successful nonindigenous species invasion, although genetic diversity is believed to play a central role. In the present study, population genetic structure, diversity and divergence patterns were analysed for the zebra mussel Dreissena polymorpha [n=280 samples and 63 putative randomly amplified polymorphic DNA (RAPDs) gene loci] and the quagga mussel D. bugensis (n=136 and 52 loci) from 10 nonindigenous North American and six Eurasian sampling sites, representing their present‐day ranges. Results showed that exotic populations of zebra and quagga mussels had surprisingly high genetic variability, similar to those in the Eurasian populations, suggesting large numbers of founding individuals and consistent with the hypothesis of multiple colonizations. Patterns of genetic relationships indicate that the North American populations of D. polymorpha likely were founded by multiple source populations from north‐western and northcentral Europe, but not from southcentral or eastern Europe. Sampling areas within North America also were significantly divergent, having levels of gene flow and migration about twice those separating long‐established Eurasian populations. Samples of D. bugensis in Lakes Erie and Ontario were significantly different, with the former being more closely related to a native population from the Dnieper River, Ukraine. No evidence for a founder effect was discerned for either species.     

Optimal foraging versus shared doom effects: interactive influence of mussel size and epibiosis on predator preference

In the western Baltic Sea, the highly competitive blue mussel Mytilus edulis tends to monopolize shallow water hard substrata. In many habitats, mussel dominance is mainly controlled by the generalist predator Carcinus maenas. These predator-prey interactions seem to be affected by mussel size (relative to crab size) and mussel epibionts.There is a clear relationship between prey size and predator size as suggested by the optimal foraging theory: Each crab size class preferentially preys on a certain mussel size class. Preferred prey size increases with crab size.Epibionts on Mytilus, however, influence this simple pattern of feeding preferences by crabs. When offered similarly sized mussels, crabs prefer Balanus-fouled mussels over clean mussels. There is, however, a hierarchy of factors: the influence of attractive epibiotic barnacles is weaker than the factor ‘mussel size’. Testing small mussels against large mussels, presence or absence of epibiotic barnacles does not significantly alter preferences caused by mussel size. Balanus enhanced crab predation on mussels in two ways: Additional food gain and, probably more important, improvement in handling of the prey. The latter effect is illustrated by the fact that artificial barnacle mimics increased crab predation on mussels to the same extent as do live barnacles.We conclude that crab predation preferences follows the optimal foraging model when prey belong to different size classes, whereas within size classes crab preferences is controlled by epibionts.     

Efficiency of blue mussel (Mytilus edulis) spat collectors in highly dynamic tidal environments of the Lower Saxonian coast (southern North Sea)

Traditional mussel culture in the Wadden Sea, southern North Sea, is carried out by taking seed mussels of about 1-4 cm shell length from natural beds and transplanting them to permanently water covered sites. Besides the damage done to the natural beds, the ratio of seeded to harvested mussels is only about 1:1-1.3, i.e. about the same tonnage of mussels seeded is recovered. In addition, this technique relies exclusively on natural spat falls, which do not occur regularly. In order to overcome these difficulties spat collectors have been deployed in the Jade Bay, southern North Sea. These provided suitable settlement grounds for mussel larvae. Blue mussel weights reached weights of about 8-9 kg/m collector rope with maximum shell lengths of 4-5 cm within one growing season.     

Mytilus galloprovincialis (Mollusca: Bivalvia) in a warm-temperate South African estuarine embayment

The distribution, population structure, condition and gonad indices, and the attachment strengths of the invasive Mediterranean mussel Mytilus galloprovincialis are described from the Knysna Estuary, South Africa. Mussels were found in the rocky intertidal at the mouth of the estuary and on all man-made hard substrata as far as 12?km from the mouth in 2013. However, mussel beds were found only in the lower reach of the estuary, where mussel densities were as high as 120 per 0.01?m². When compared to mussels from the embayment regions of the estuary, those at the estuary mouth, where wave action was high, were generally smaller, had significantly lower condition and gonad indices, and greater attachment strength. This suggests that mussels within the sheltered regions of the estuary invest more energy in growth and reproduction, and that this is one reason for their success as an invasive species in the Knysna estuarine embayment.     

Commensals of Mytilus galloprovincialis in the Black Sea: Urastoma cyprinae (Turbellaria) and Polydora ciliata (Polychaeta)

The intensity of infection of the mussel Mytilus galloprovincialis Lamarck in the Black Sea by the turbellarian Urastoma cyprinae (Graff), which lives on its gills, was found to be higher in larger hosts, reaching a maximum in mussels of 50–70 mm length. Greater numbers occured in mussels inhabiting a silty bottom than in cultivated mussels suspended above the bottom. Over the period 1982–1987, U. cyprinae was most numerous in winter and especially so in years that were colder. The spionid polychaete Polydora ciliata (Johnston) also infects M. galloprovincialis, burrowing into the shell. Young spionids of up to 1 mm length occured in mussels with a shell length of 35 mm. Numbers of this commensal reached a maximum in mussels of intermediate size.     

Body size and food thresholds for zero growth in Dreissena polymorpha: a mechanism underlying intraspecific competition

1. Dreissena polymorpha is an extraordinarily successful invasive species that shows high recruitment of small juvenile mussels on established mussel banks. Such juvenile settlement on, and overgrowth of, large adult mussels; however, leads to competition with adults, and often at high densities and low‐food concentrations. 2. The concept of food thresholds for zero growth has been a powerful approach to explaining size‐related exploitative competition in different zooplankton species. We applied it to investigate whether food threshold concentrations for zero growth (C₀) differ between juvenile and adult zebra mussels. 3. By determining body mass growth at various concentrations of a diet mixture (Nannochloropsis limnetica and Isochrysis aff. galbana) we demonstrate that the threshold food concentration for growth of juvenile mussels (C₀ = 0.08 mg C L⁻¹) is substantially lower than that for adults (C₀ = 0.36 mg C L⁻¹). 4. This indicates that, at low food availability, juvenile zebra mussels are competitively superior to their larger conspecifics. Within zebra mussel banks plankton food is substantially depleted and so the observed mechanism might ensure juvenile success and therefore the regeneration of mussel banks in nature.     

Shell morphometrics and growth rate of the invasive bivalve mollusc Mytilus galloprovincialis in the Knysna estuarine embayment,South Africa

Shell morphometrics of the invasive mussel Mytilus galloprovincialis were compared at five sites and growth rate at four sites (in four seasons) in the Knysna estuarine embayment. Mussels from two sites (The Heads, Leisure Isle) where wave action was present had shells significantly lower for any length when compared with other more sheltered sites. There was no significant difference in shell width of mussels for any given length among sites. Mussels from The Heads had thicker shells than other sites, and those from Leisure Isle thicker shells than three other embayment sites where shells did not differ in thickness. Growth rate of mussels at two embayment sites (Thesen’s Wharf and Thesen Islands Marina) was greatest in autumn and summer whereas at The Heads and Leisure Isle there was little seasonal difference in growth rate. Growth rate of mussels at Thesen’s Wharf and Thesen Islands Marina was mainly greater in all seasons when compared with mussels at The Heads and Leisure Isle. The more rapid growth rate of mussels at the sheltered embayment sites might in part explain why M. galloprovincialis now dominates the mid- to lower intertidal on hard substrata in this region of the Knysna estuary.     

Comparing grazing by Dreissena polymorpha on phytoplankton in the presence of toxic and non-toxic cyanobacteria

SUMMARY 1. The feeding behaviour of the zebra mussel ( Dreissena polymorpha ) was studied in the laboratory on different combinations of food, including a green alga ( Chlamydomonas reinhardtii ) and toxic and non-toxic strains of the cyanobacterium Microcystis aeruginosa .
2. The highest clearance rate of phytoplankton by zebra mussels was found when the mussels were feeding on a mixture of Chlamydomonas and non-toxic Microcystis , the lowest on a mixture of Chlamydomonas and toxic Microcystis .
3. The differences found in the clearance rates between food combinations can be partly explained by the production of pseudofaeces containing live phytoplankton cells. Zebra mussels expelled significantly more live phytoplankton cells in the presence of toxic Microcystis than in the presence of non-toxic Microcystis . The pseudofaeces contained predominantly live Chlamydomonas cells. Proportionately much less live Microcystis cells were encountered in the pseudofaeces.
4. Consequently, grazing of zebra mussels on a combination of Chlamydomonas and Microcystis may finally result in a dominance of Chlamydomonas over Microcystis . The presence of toxic Microcystis may even strengthen this shift.     

The role of the blue mussel Mytilus edulis in the cycling of nutrients in the Oosterschelde estuary (The Netherlands)

The fluxes of particulate and dissolved material between bivalve beds and the water column in the Oosterschelde estuary have been measured in situ with a Benthic Ecosystem Tunnel. On mussel beds uptake of POC, PON and POP was observed. POC and PON fluxes showed a significant positive correlation, and the average C:N ratio of the fluxes was 9.4. There was a high release of phosphate, nitrate, ammonium and silicate from the mussel bed into the water column. The effluxes of dissolved inorganic nitrogen and phosphate showed a significant correlation, with an average N:P ratio of 16.5. A comparison of the in situ measurements with individual nutrient excretion rates showed that excretion by the mussels contributed 31–85% to the total phosphate flux from the mussel bed. Ammonium excretion by the mussels accounted for 17–94% of the ammonium flux from the mussel bed. The mussels did not excrete silicate or nitrate. Mineralization of biodeposition on the mussel bed was probably the main source of the regenerated nutrients.From the in situ observations net budgets of N, P and Si for the mussel bed were calculated. A comparison between the uptake of particulate organic N and the release of dissolved inorganic N (ammonium + nitrate) showed that little N is retained by the mussel bed, and suggested that denitrification is a minor process in the mussel bed sediment. On average, only 2/3 of the particulate organic P, taken up by the mussel bed, was recycled as phosphate. A net Si uptake was observed during phytoplankton blooms, and a net release dominated during autumn. It is concluded that mussel beds increase the mineralization rate of phytoplankton and affect nutrient ratios in the water column. A comparison of N regeneration by mussels in the central part of the Oosterschelde estuary with model estimates of total N remineralization showed that mussels play a major role in the recycling of nitrogen.