Average annual growth and condition of mussels as a function of food source

Prior to 1986, before completion of a storm surge barrier, the average annual condition of mussels at culture plots in the Eastern Scheldt was better in the western part near the North Sea, than in the central part of the estuary. No such difference was observed in 1986 and 1987.To predict the effects of the barrier on mussel culture, the condition of the mussels, which is an index of growth rate, is analysed and correlated with food sources for the period 1981–84. In the western and central part of the Eastern Scheldt, mussel condition correlates strongly with average annual primary production, but not with chlorophyll-a concentrations. This suggests a direct link between primary production and the growth of mussels.In the western part of the estuary, the relation between condition and primary production has a less steep slope than in the central part. Import of food from the North Sea is suggested to act as an additional food source in the west of the Eastern Scheldt; hence the better condition values of the mussels, also in years with relatively low primary production.The storm surge barrier reduces the water exchange with the North Sea. Reduced import of food and consequently lower mussel condition are expected in the western part of the estuary. Preliminary data from 1987 confirm this prediction.     

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.     

The effect of seasonal variation in selective feeding by zebra mussels (Dreissena polymorpha) on phytoplankton community composition

1. To investigate the impact of zebra mussels ( Dreissena polymorpha ) on phytoplankton community composition, temporal variability in selective feeding by the mussels was determined from April to November 2005 in a natural lake using Delayed Fluorescence (DF) excitation spectroscopy.
2. Selective grazing by zebra mussels varied in relation to seasonal phytoplankton dynamics; mussels showed a consistent preference for cryptophytes and avoidance of chlorophytes and cyanobacteria. Diatoms, chrysophytes and dinoflagellates responded differentially to zebra mussel grazing depending on their size. Analysis of excreted products of the zebra mussels revealed that in addition to chlorophytes and cyanobacteria, phytoplankton >50  μ m and very small phytoplankton (≤7  μ m) were largely expelled in pseudofaeces.
3. The zebra mussel is a selective filter-feeder that alters its feeding behaviour in relation to phytoplankton composition to capture and ingest high quality phytoplankton, especially when phytoplankton occur in preferred size ranges. Flexibility of zebra mussel feeding behaviour and variation in susceptibility among phytoplankton groups to mussel ingestion indicate that invading zebra mussels could alter phytoplankton community composition of lakes and have important ecosystem consequences.     

Short-term impact of blue mussel dredging (Mytilus edulis L.) on a benthic community

The short-term effect of mussel dredging in a brackish Danish sound was studied. A commercial dredging track was identified and an analysis of the species composition inside the track and at an adjacent control area showed that dredging changed the community structure by reducing the density of polychaetes. In order to investigate the extent and the duration of the dredging impact experimental dredging was conducted. The experimental dredging removed 50% of the mussels in two dredged areas. Immediately after dredging, a significantly lower number of species was measured inside the mussel beds in dredged areas compared to control and boundary areas. This effect lasted for at least 40 days. The analysis of the species composition showed that the dredged area had a significantly lower density, particularly of polychaetes compared to the boundary area. An increased number of species was recorded outside the mussel beds just after dredging, but this effect lasted for less than 7 days. After dredging, brown shrimps, C. crangon invaded the dredged areas. This species is an important predator of smaller invertebrates, and it is suspected that it was feeding on small vulnerable polychaetes exposed at the sediment surface after dredging. The dredging process was observed to form 2–5-cm deep furrows in the seabed, but the sediment texture and the organic content of the sediment was not affected. The biomass accumulation of individual blue mussels was significantly lower in the dredged area compared to the boundary area. This indicates that the disturbance of the mussel bed structure reduced growth and that the lowering of intraspecific food competition caused by a reduced density of mussels did not increase the accumulation of biomass in the mussels which remained in the dredged area.     

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.     

Grazing effects of a freshwater bivalve (Corbicula leana Prime) and large zooplankton on phytoplankton communities in two Korean lakes

This study examined the effects of a freshwater filter feeding bivalve (Corbicula leana Prime) and large zooplankton (>200 μm, mostly cladocerans and copepods) on the phytoplankton communities in two lakes with contrasting trophic conditions. A controlled experiment was conducted with four treatments (control, zooplankton addition, mussel addition, and both zooplankton and mussel addition), and each established in duplicate 10-l chambers. In both lakes there were significant effects of mussel grazing on phytoplankton density and biomass. The effects were greater in mesotrophic Lake Soyang than in hypertrophic Lake Ilgam. Effects of zooplankton grazing did not differ between these lakes, and zooplankton effects on phytoplankton were much less than the effects of mussels. Although mussels exerted a varying effect on phytoplankton according to their size, mussels reduced densities of almost all phytoplankton taxa. Total mean filtering rate (FR) of mussels in Lake Soyang was significantly greater than that in Lake Ilgam (p=0.002, n=5). Carbon fluxes from phytoplankton to mussels (977–2,379 μgC l^?1d^?1) and to zooplankton (76–264 μgC l^?1 d^?1) were always greater in Lake Ilgam due to the greater phytoplankton biomass (p<0.01, n=6). Based on the C-flux to biomass ratios, the mussels consumed 170–754% (avg. 412%) of phytoplankton standing stock in Lake Soyang, and 38–164% (avg. 106%) in Lake Ilgam per day. The C-flux to biomass ratio for mussels within each lake was much greater than for large zooplankton. Mussels reduced total phosphorus concentration by 5–34%, while increasing phosphate by 30–55% relative to the control. Total nitrogen also was reduced (by 9–25%), but there was no noticeable change in nitrate among treatments. The high consumption rate of phytoplankton by Corbicula leana even in a very eutrophic lake suggests that this mussel could affect planktonic and benthic food web structure and function by preferential feeding on small seston and by nutrient recycling. Control of mussel biomass therefore might be an effective tool for management of water quality in shallow eutrophic lakes and reservoirs in Korea.     

Evaluation of the Danish mussel fishery: suggestions for an ecosystem management approach

In Limfjorden, Denmark, an extensive mussel fishery exploits the wild stocks of Mytilus edulis with annual landings of 80,000–100,000 t of mussels. During the last 10 years the impact of mussel dredging on the ecosystem has been studied, including the effect of resuspension of sediment and nutrients and the impoverishment of in- and epi-fauna assemblages. Furthermore, dredging changes the physical structure and complexity of the seabed which affects mussel growth and interactions among zoobenthic species. The blue mussel constitutes the dominant fraction of the zoobenthic suspension feeders, and is important for the transport of material and energy from the pelagic to benthic systems and the control of phytoplankton biomass. In order to evaluate the impact on clearance capacity of a reduction in mussel densities due to mussel dredging, mussel filtration activity measured in situ has been related to the mixing of the water column and the amount of near-bed phytoplankton. Fishery practice for mussel dredging in Limfjorden is discussed in relation to its known impact on the ecosystem and the ecological role of the mussels, and modifications towards an ecosystem management approach and a more sustainable fishery are suggested. The suggested modifications include: a fishery practice where the mussel beds are thinned out when the mussels have attained good quality, and a transplantation practice of mussels from areas with a high mortality to areas with a high growth rate. Both practices intensify the production in a certain area, leaving other areas open for alternative production or for permanent closure for the benefit of the benthic flora and fauna. In addition, other shellfish species represent interesting new resources for fishing or aquaculture. Habitat restoration, such as the relaying of mussel shells from the mussel industry, is another important management tool that should be included in an ecosystem management approach of the mussel fishery. Electronic Publication     

The nearshore shunt and the decline of the phytoplankton spring bloom in the Laurentian Great Lakes: insights from a three-dimensional lake model

Dreissenid mussels have been hypothesized to cause selective decreases of phytoplankton in nearshore areas (nearshore shunt hypothesis) as well as the near-complete loss of the offshore phytoplankton spring bloom in some Laurentian Great Lakes. To evaluate whether mussels can reasonably be expected to mediate such changes, we extended the three-dimensional hydrodynamic-ecological model (ELCOM-CAEDYM) to include mussels as a state variable and applied it to Lake Erie (USA-Canada). Mussel-mediated decreases in mean phytoplankton biomass were highly sensitive to the assigned mussel population size in each basin. In the relatively deep east basin, mussels were predicted to decrease phytoplankton in both nearshore and offshore zones, even during periods of thermal stratification but especially during the spring phytoplankton maximum. Spatially, impacts were associated with mussel distributions but could be strong even in areas without high mussel biomass, consistent with advection from areas of higher mussel biomass. The results supported the nearshore shunt hypothesis that mussel impacts on phytoplankton should be greater in nearshore than offshore waters and also supported suggestions about the emerging importance of deep water offshore mussels. The results of this study provide an important insight into ecological role of mussels in lowering plankton productivity in some world’s largest lakes.     

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.     

Impact of suspended mussels (Mytilus edulis L.) on plankton communities in a Magdalen Islands lagoon (Québec, Canada): A mesocosm approach

The Grande-Entrée Lagoon (Magdalen Islands, Canada) has supported mussel (Mytilus edulis) cultivation for the last 25 years. Algal biomass in this lagoon is relatively low while heterotrophic plankton biomass is high. Although often considered herbivorous, it is known that filter-feeding bivalves can consume various types of food, from bacteria to zooplankton. We hypothesize that along with phytoplankton, heterotrophs constitute an important food resource for the Grande-Entrée mussels. In situ mesocosm experiments were undertaken at different seasons using short socks filled with mussels from the same cohort taken from an aquaculture farm, in order to determine the impact of cultured mussels on local plankton communities and assess the role of heterotrophs. Filtration activity by the mussels and associated epibionts present in the socks was expressed as clearance rates (CR). The average CR over all taxa was lowest in June and highest in October. Diatoms, dinoflagellates and heterotrophic protists constituted the bulk of planktonic carbon removed by mussels. While smaller-sized taxa contributed little (< 5%) to mussel carbon intake, large-sized heterotrophs (namely ciliates) contributed 69 to 88%. Taxon-marker pigment analyses generally confirmed these observations for groups containing phototrophic pigments. The high heterotrophic biomass retained by mussels indicates they are a major food source for mussels in this environment and should be considered both in the evaluation of mussel feeding and in assessing the influence of cultured mussels on local plankton ecosystems.     

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).     

Physical forces constrain the depth distribution of the abundant native mussel Elliptio complanata in lakes

1. Unionid mussels often account for a large portion of benthic biomass and contribute to nutrient cycling and sediment processes, but are thought to be limited to shallow areas (<2–3 m). 2. The depth distribution and body size of Elliptio complanata were compared in seven Canadian Shield lake basins of different sizes to test what factors determine the upper and lower limit of their depth range. Specifically, I tested whether (i) the upper range of their distribution is limited by exposure to winds and wave action and (ii) the lower range of their distribution is limited by the depth of the thermocline or by the boundary of mud deposition. 3. The average depth distribution of E. complanata shifted to greater depths in larger lake basins. When comparing individual transects, maximum mussel density was found deeper at more exposed sites. Mussel size decreased with increasing depth and was larger, on average, in larger lake basins. These results suggest that physical forces limit the upper range of mussel distribution in lakes. 4. The maximum depth at which mussels were found in different lakes was closely related to thermocline depth. However, mussels were commonly observed below the predicted depth of the mud deposition boundary. The thermocline limits the lower range of mussel distribution in lakes, probably by limiting food availability and by determining water temperature. Substratum type does not limit the lower distribution of mussels. 5. These results suggest that unionid mussels are present in the deeper parts of the littoral zone, especially in large lakes. Therefore, comparisons of mussel populations between sites and between lakes would be biased unless the full depth distribution of these mussels is considered. These results also suggest that long‐term changes in the thermal structure of lakes could affect the range of unionid mussel populations and their functional role in littoral ecosystems.     

Spatial patterns of abundance and growth of Mytilus edulis on boulders in the Northern Baltic Sea proper

Small scale spatial patterns of abundance, growth and condition of the mussel Mytilus edulis on sub-littoral boulders (approx. 1–2 m high) were investigated at the island of Askö in the northern Baltic proper. The effect of side (exposed/sheltered with respect to wave action and sunlight) of boulder and position (up/down) on boulder was investigated. A large spatial variability in abundance of M. edulis between boulders and between various sites within boulders were found. The highest numbers of mussels were found on the wave exposed side, near the top of boulders. Shell growth was favoured by a sheltered side and a down position. The body condition (meat weight/shell weight) of mussels was, on the other hand, affected only by position, the condition of mussels being better at the up position. Consequently, there seems to be temporal differences in the condition for growth within a spatial position. The body condition of the mussels was best near the top of boulders in the spring, but long-term shell growth was favoured by a sheltered side and a down position. This may reflect changes in the composition and availability of food during the year with phytoplankton as the major food source during the spring bloom and resuspension of benthic production and detritus as relatively more important during the rest of the year.     

Eutrophication and mussel culture in the western Dutch Wadden Sea: Impact on the benthic ecosystem; a hypothesis

Since 1950, two large-scale changes have taken place in the western Dutch Wadden Sea, namely the eutrophication of the area and the introduction of an extensive mussel culture. Although eutrophication in the fresh waters started already around 1950, nutrient concentrations in the western Wadden Sea remained fairly constant until about 1970, due to the retention of nutrients in Lake IJssel, the main source. From 1970–1980 concentrations increased strongly, and during the last years the situation has stabilized. Mussel culture was introduced in 1950 and expanded during the next decade to an area of 70 km², all situated in the sublittoral area. From 1960 the area of mussel culture remained about constant with fluctuating yields of between 35 and 120 millions of kg fresh weight. Due to a lack of data for the period until 1970 the impact of eutrophication and mussel culture on the ecosystem cannot be assessed. From 1970 onwards an increased biomass and production of the macrofauna in the intertidal zone has been observed, which is attributed to eutrophication. The hypothesis is postulated that the introduction of mussel culture between 1950 and 1960 has resulted in an increased food competition in the area, leading to a decreased stock of the macrofauna in the intertidal. Eutrophication from about 1970 onwards has improved the food conditions and as a result both the macrofauna in the intertidal and the mussel in the sublittoral area would have increased in biomass, allowing higher maximum yields of the mussel culture. The importance of monitoring programs is stressed to follow these trends in the near future and to check the above hypothesis in areas where it is decided to introduce or intensify mussel culture. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988) Publication No. 30 of the project “Ecological Research of the North Sea and Wadden Sea” (EON)     

Long-term demography of a zebra mussel (Dreissena polymorpha) population

1. We used long‐term data and a simulation model to investigate temporal fluctuations in zebra mussel populations, which govern the ecological and economic impacts of this pest species. 2. The size of the zebra mussel (Dreissena polymorpha) population in the Hudson River estuary fluctuated approximately 11‐fold across a 13‐year period, following a cycle with a 2–4 year period. 3. This cycling was caused by low recruitment during years of high adult population size, rapid somatic growth of settled animals, and adult survivorship of 50% per year. 4. Adult growth and body condition were weakly correlated with phytoplankton biomass. 5. The habitat distribution of the Hudson's population changed over the 13‐year period, with an increasing proportion of the population spreading onto soft sediments over time. The character of soft‐sediment habitats in the Hudson changed because of large amounts (mean = 34 g DM m^?2) of empty zebra mussel shells now in the sediments. 6. Simulation models show that zebra mussel populations can show a range of long‐term trajectories, depending on the balance between adult space limitation, larval food limitation, and disturbance. 7. Effective understanding and management of the effects of zebra mussels and other alien species depend on understanding of their long‐term demography, which may vary across ecosystems.     

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.     

Secondary production of the brackish copepod communities and their contribution to the carbon fluxes in the Westerschelde estuary (The Netherlands)

The zooplankton community of the brackish part of the Westerschelde estuary (November 1989–October 1990) was dominated by two calanoid copepods, Eurytemora affinis and Acartia tonsa. Eurytemora was present during a longer period of the year and was much more important in terms of total abundances and biomasses than Acartia.The secondary production of these species was estimated by means of the growth rate method, using weight-specific growth rates obtained from laboratory cultures (Eurytemora) or from the literature (Acartia).Due to the substantially higher growth rates of Acartia compared to Eurytemora, total yearly productions of both communities were comparable, notwithstanding the large discrepancies in biomass. They amounted to about 5 and 6 g C m^–2 y^–1 by Acartia and Eurytemora respectively.The food needed to realise this production was estimated to be about 14 and 17 g C m^–2 y^–1 by Acartia and Eurytemora respectively. Provided that the copepods are able to selectively ingest the phytoplankton, in situ net production provides sufficient carbon for zooplankton demands for a short period of the year only. As phytoplankton standing stock is very low and net phytoplankton productivity is negative from late fall to early spring, nutritional demands of the copepods have to be fulfilled by other than algal food at least during this period of the year.Although the copepods in the brackish part can have an important impact on some food items, their contribution to total carbon fluxes in the brackish zone is negligible: each year some 6% of all consumed carbon in the brackish part of the estuary passes through the copepod food web.     

Sediment accumulation predicts the distribution of a unionid mussel (Elliptio complanata) in nearshore areas of a Canadian Shield lake

1. The importance of native freshwater mussels for ecosystem processes depends on their density, size distribution and activity. In lakes, many of the factors that affect mussels (fish hosts, habitat, food) could be directly or indirectly related to wind‐driven physical processes. 2. We tested whether the abundance and size of Elliptio complanata in the shallow, nearshore areas of a medium‐sized lake were related to site exposure, substratum type and fish distribution. To disentangle some of the correlated variables known to affect mussel distribution, we used paired exposed and sheltered sampling sites along the 7‐km fetch of the lake basin. 3. The distribution of sediment characteristics in nearshore areas was highly predictable. The mean depth of accumulated soft sediments decreased with increasing fetch at wind‐exposed sites, but increased with increasing fetch at sheltered sites. Sediments were deeper along the main shoreline than around islands. Deeper sediments tended to be finer and higher in silt content and organic fraction. 4. The density and proportion of juvenile mussels along the main shoreline varied in a unimodal way with sediment depth. These results suggest that wind‐driven physical forces affect the transport of young juveniles to sediment depositional areas and create sediment conditions that influence their growth and survival. In contrast, the proportion of juvenile mussels around islands was not related to sediment characteristics, but decreased with remoteness of the island, suggesting that the distribution of juvenile mussels may be limited by fish movements. These results are tentative since they do not include buried juvenile mussels. 5. We also found a unimodal relationship between total mussel density (juveniles and adults) and sediment depth but, in contrast to the relationship for juveniles only, it applied to all sites with soft sediments, including islands. We conclude that factors related to sediment depth affect the growth and survival of adult mussels around islands and that these factors are strong enough to modify the pattern of distribution established via dispersal during earlier life stages. 6. The mean shell length of adults at different sites within the lake basin ranged from 60 to 85 mm. Mussel shell length decreased with increasing fetch at sites exposed to the prevalent winds, but was relatively constant on the sheltered side of peninsulas and islands. The size of unionid mussels in different parts of the lake seems to be determined both by their exposure to physical forces and by sediments. 7. The local distribution of E. complanata is determined, directly and indirectly, by wind‐driven forces. These processes are likely to be important for other benthic organisms affected by similar habitat conditions (e.g. sediment characteristics, physical disturbance).     

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

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Consumption of benthic invertebrates by waterbirds in the Oosterschelde estuary,SW Netherlands

The number of waders in the Oosterschelde, S.W. Netherlands, declined after a reduction in intertidal area due to the construction of a storm surge barrier and secondary dams, suggesting that the carrying capacity had been reached (Schekkerman et al., 1993). In this paper we present data on consumption and predation pressure by birds to explore whether the reduction in their numbers is due to prey depletion or to other factors.The total annual consumption of benthic invertebrates by birds in the Oosterschelde amounted to 1573 × 10³ g ADW y^–1 in the period before the coastal engineering works (pre-barrier) and 1500 × 10³ kg ADW y^–1 in the post-barrier period. More than half of the total amount of biomass is eaten by the Oystercatcher, and only seven (pre-barrier) or even six (post-barrier) bird species together take 90% of the total.Although the consumption by individual species may vary considerably among years, the total consumption was remarkably stable, with a CV of only 3–4% of the mean, especially compared to the variability of the prey populations. In the pre-barrier period, consumption was lowest in mid summer, increased sharply from August onwards until a peak was reached in January. A sharp decrease took place in March. In the post-barrier period, consumption peaked in October.The total consumption per unit area per year does not differ much between different sectors of the Oosterschelde, apart from a distinctly lower value in the eastern part. Of the total amount of food taken by birds, only 0.1–0.4% is taken in the subtidal compartment. In several study plots on an individual tidal flat, there was a clear relation between consumption and benthic biomass.The predation pressure was 13 and 23% of the standing stock, in the post- and pre-barrier period respectively. When cockles, mussels and their main predator, the Oystercatcher, are excluded from the calculations, the predation pressure of the other species was 30 and 37% of the biomass, respectively.Predation pressure of Oystercatchers in individual study plots varied from less than 10% to more than 70% of the standing stock. On cockle beds the predation pressure was positively related to the average length of the cockles present.Based on these results and a comparison with the literature we conclude that, at least for several species that feed intertidally, carrying capacity could be limited by the stocks of food. This does not mean that birds face food shortage each season. As the variability of the benthos populations is much higher than that of the bird densities it is likely that at some times food is not limiting, at other times it is. On the other hand, consumption is very low in the subtidal compartment and species feeding here could potentially increase substantially in numbers in the Oosterschelde.