Fouling mussels (Dreissena spp.) colonize soft sediments in Lake Erie and facilitate benthic invertebrates

• 1 We conducted survey and transplant studies to determine whether colonization and residency on soft sediments by introduced, fouling mussels (Dreissena polymorpha and D. bugensis) were affected by physical disturbance, and whether Dreissena presence in turn influenced the diversity and population densities of other benthic invertebrates. Surveys revealed that colony density was typically higher at moderate depths than at shallower and greater ones. However, the largest, midsummer colonies and greatest coverage of sediments by mussels occurred at deeper sites.
• 2 Disturbance of transplanted colonies varied by depth and colony size, with deeper and larger colonies experiencing the lowest destruction rates. Colony destruction rate was positively correlated with current velocity adjacent to the lakebed.
• 3 Absence of mussel colonies at shallow sites was not determined by recruitment or substrate limitation, as recruit density was higher and sediment characteristics more suitable for postveliger settlement at shallow than at deeper sites. Rather, seasonal storms have much stronger effects in shallow than in deep water.
• 4 Mussel residency on soft sediment has profound effects on invertebrate biodiversity. Invertebrate species (taxon) richness and total abundance were positively correlated with mussel colony area. Mussel‐sediment habitat supported between 462 and 703% more taxa, and between 202 and 335% more individuals (exclusive of Dreissena) than adjacent soft‐sediment lacking mussels.
• 5 Results from this study illustrate that physical disturbance directly limits the distribution of mussels on soft sediments, and the diversity and abundance of other benthic invertebrates in consequence.

     

Effects of Adult Dreissena polymorpha on Settling Juveniles and Associated Macroinvertebrates

The impact of Dreissena polymorpha settlement on recruitment of juvenile mussels and density of other macroinvertebrates was studied in field experiments using blank concrete blocks and tiles (control), blocks and tiles with attached empty zebra mussel shells, and blocks and tiles with attached living mussels. On blocks, dominant invertebrate taxa showed colonization patterns coinciding with increased habitat complexity owing to zebra mussel settlement or the biodeposition of faeces and pseudofaeces. Adult and especially juvenile zebra mussels preferred blocks with empty shells to blank blocks and blocks with living mussels; this might possibly be caused by a chemical cue that induces gregarious settlement. Lower recruitment on blocks with attached living mussels compared to blocks with only shells could be the consequence of ingestion of larvae by adult mussels and of competition for food. On tiles, the sediments deposited and the organic content of the sediment were investigated. Sedimentation was significantly higher on shell‐only and live‐mussel tiles compared to blank tiles. Organic matter differed significantly between blank and live‐mussel tiles.     

Zebra mussels affect benthic predator foraging success and habitat choice on soft sediments

The introduction of zebra mussels (Dreissena spp.) to North America has resulted in dramatic changes to the complexity of benthic habitats. Changes in habitat complexity may have profound effects on predator-prey interactions in aquatic communities. Increased habitat complexity may affect prey and predator dynamics by reducing encounter rates and foraging success. Zebra mussels form thick contiguous colonies on both hard and soft substrates. While the colonization of substrata by zebra mussels has generally resulted in an increase in both the abundance and diversity of benthic invertebrate communities, it is not well known how these changes affect the foraging efficiencies of predators that prey on benthic invertebrates. We examined the effect of zebra mussels on the foraging success of four benthic predators with diverse prey-detection modalities that commonly forage in soft substrates: slimy sculpin (Cottus cognatus), brown bullhead (Ameirus nebulosus), log perch (Percina caprodes), and crayfish (Orconectes propinquus). We conducted laboratory experiments to assess the impact of zebra mussels on the foraging success of predators using a variety of prey species. We also examined habitat use by each predator over different time periods. Zebra mussel colonization of soft sediments significantly reduced the foraging efficiencies of all predators. However, the effect was dependent upon prey type. All four predators spent more time in zebra mussel habitat than in either gravel or bare sand. The overall effect of zebra mussels on benthic-feeding fishes is likely to involve a trade-off between the advantages of increased density of some prey types balanced against the reduction in foraging success resulting from potential refugia offered in the complex habitat created by zebra mussels.     

Habitat engineering by the invasive zebra mussel <Emphasis Type="Italic">Dreissena polymorpha</Emphasis> (Pallas) in a boreal coastal lagoon: impact on biodiversity

Habitat engineering role of the invasive zebra mussel Dreissena polymorpha (Pallas) was studied in the Curonian lagoon, a shallow water body in the SE Baltic. Impacts of live zebra mussel clumps and its shell deposits on benthic biodiversity were differentiated and referred to unmodified (bare) sediments. Zebra mussel bed was distinguished from other habitat types by higher benthic invertebrate biomass, abundance, and species richness. The impact of live mussels on biodiversity was more pronounced than the effect of shell deposits. The structure of macrofaunal community in the habitats with >10³ g/m² of shell deposits devoid of live mussels was similar to that found within the zebra mussel bed. There was a continuous shift in species composition and abundance along the gradient ‘bare sediments—shell deposits—zebra mussel bed’. The engineering impact of zebra mussel on the benthic community became apparent both in individual patches and landscape-level analyses.     

Space as a limiting resource in freshwater systems: competition between zebra mussels (Dreissena polymorpha) and freshwater sponges (Porifera)

Species interactions between two types of sessile benthic invertebrates, the zebra mussel (Dreissena polymorpha) and freshwater sponges (Porifera), were evaluated in Michigan City IN Harbor in southern Lake Michigan during 1996. The study objective was to define whether competition plays a role in structuring benthic communities using experimental techniques commonly employed in marine systems. Sponges were uninhibited by zebra mussel presence and overgrew zebra mussel shells on hard vertical substrata. In contrast, zebra mussels did not overgrow sponge colonies, but did show an ability to re-capture hard substrata if relinquished by the sponge. The negative affect of sponges on zebra mussels through overgrowth and recruitment suggests interactions that could eventually displace zebra mussels from these benthic communities. However, seasonal reduction of sponge biomass from autumn through winter appears to allow the zebra mussel a periodic respite from overgrowth, preventing exclusion of zebra mussels from the community and allowing these two taxa to co-exist.     

Rapid range expansion of the invasive quagga mussel in relation to zebra mussel presence in The Netherlands and Western Europe

Since its appearance in 2006 in a freshwater section of the Rhine–Meuse estuary (Hollandsch Diep, The Netherlands), the non-indigenous quagga mussel has displayed a rapid range expansion in Western Europe. However, an overview characterising the spread and impacts of the quagga mussel in this area is currently lacking. A literature study, supplemented with field data, was performed to gather all available data and information relating to quagga mussel dispersal. Dispersal characteristics were analysed for rate and direction and in relation to hydrological connectivity and dispersal vectors. To determine ranges of conditions suitable for quagga mussel colonisation, physico-chemical characteristics of their habitats were analysed. After its initial arrival in the freshwater section of the Rhine-Meuse estuary and River Danube, the quagga mussel demonstrated a rapid and continued range expansion in Western Europe. Quagga mussels have extended their non-native range to the network of major waterways in The Netherlands and in an upstream direction in the River Rhine (Germany), its tributaries (rivers Main and Moselle) and the River Meuse (Belgium and France). The calculated average quagga mussel dispersal rate in Europe was 120 km year^?1 (range 23–383 km year^?1). Hydrological connectivity is important in determining the speed with which colonisation occurs. Dispersal to water bodies disconnected from the freshwater network requires the presence of a suitable vector e.g. pleasure boats transferred over land. Upstream dispersal is primarily human mediated through the attachment of mussels to watercraft. The relative abundance of quagga mussel to zebra mussel has greatly increased in a number of areas sampled in the major Dutch rivers and lakes and the rivers Main and Rhine and the Rhine–Danube Canal leading to a dominance shift from zebra mussels to quagga mussels. However, evidence for displacement of the zebra mussel is limited due to the lack of temporal trends relating to the overall density of zebra and quagga mussel.     

Effects of zebra mussel attachment on the foraging behaviour of a larval dragonfly,Macromia illinoiensis

1. Larvae of Macromia illinoiensis Walsh are often colonised by the zebra mussel, Dreissena polymorpha Pallas, a recent invader to North America. To determine how mussel attachment affects an individual's foraging behaviour, we quantified capture of Hexagenia limbata Hexes mayfly prey and the distance moved by newly‐molted final instars before and after an individual's colonisation with zebra mussels. 2. In night trials, larvae sprawled above the sand, and caught more mayflies than individuals in daytime trials, but the estimated distance travelled did not differ. When resting under a layer of sand with only its eyes exposed during the day, an individual could capture a mayfly prey using a sit‐and‐wait ambush strategy. When sprawled above the sand, some larvae caught prey that rested on their legs. 3. When mussel‐free, individuals captured more prey than they did when carrying zebra mussels, although mussel attachment per se did not affect the estimated distance that a larva moved. 4. During day trials, but not night ones, the increasing mussel load of colonised individuals decreased prey capture and the distance moved in an apparent step‐wise function. Although the number of mussels carried did not differ, night foragers carried a heavier load. Independent of time of the day, the distance an individual travelled when mussel‐free was predictive of the number of prey it caught when colonised, suggesting that the greater general activity of some individuals helped mitigate negative effects that mussel attachment had on prey capture. 5. Our results add to a growing number of negative effects of zebra mussel colonisation on sprawling and hiding dragonfly larvae. Although the impact of these costs on dragonfly populations remains to be determined, a decrease in this guild of predators whose life cycle spans aquatic and terrestrial habitats might have cascading effects across ecosystems.     

Predation on exotic zebra mussels by native fishes: effects on predator and prey

SUMMARY 1. Exotic zebra mussels, Dreissena polymorpha, occur in southern U.S. waterways in high densities, but little is known about the interaction between native fish predators and zebra mussels. Previous studies have suggested that exotic zebra mussels are low profitability prey items and native vertebrate predators are unlikely to reduce zebra mussel densities. We tested these hypotheses by observing prey use of fishes, determining energy content of primary prey species of fishes, and conducting predator exclusion experiments in Lake Dardanelle, Arkansas. 2. Zebra mussels were the primary prey eaten by 52.9% of blue catfish, Ictalurus furcatus; 48.2% of freshwater drum, Aplodinotus grunniens; and 100% of adult redear sunfish, Lepomis microlophus. Blue catfish showed distinct seasonal prey shifts, feeding on zebra mussels in summer and shad, Dorosoma spp., during winter. Energy content (joules g⁻¹) of blue catfish prey (threadfin shad, Dorosoma petenense; gizzard shad, D. cepedianum; zebra mussels; and asiatic clams, Corbicula fluminea) showed a significant species by season interaction, but shad were always significantly greater in energy content than bivalves examined as either ash-free dry mass or whole organism dry mass. Fish predators significantly reduced densities of large zebra mussels (>5 mm length) colonising clay tiles in the summers of 1997 and 1998, but predation effects on small zebra mussels (≤5 mm length) were less clear. 3. Freshwater drum and redear sunfish process bivalve prey by crushing shells and obtain low amounts of higher-energy food (only the flesh), whereas blue catfish lack a shell-crushing apparatus and ingest large amounts of low-energy food per unit time (bivalves with their shells). Blue catfish appeared to select the abundant zebra mussel over the more energetically rich shad during summer, then shifted to shad during winter when shad experienced temperature-dependent stress and mortality. Native fish predators can suppress adult zebra mussel colonisation, but are ultimately unlikely to limit population density because of zebra mussel reproductive potential.     

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.     

Context-dependent effects of freshwater mussels on stream benthic communities

1. We asked whether unionid mussels influence the distribution and abundance of co‐occurring benthic algae and invertebrates. In a yearlong field enclosure experiment in a south‐central U.S. river, we examined the effects of living mussels versus sham mussels (shells filled with sand) on periphyton and invertebrates in both the surrounding sediment and on mussel shells. We also examined differences between two common unionid species, Actinonaias ligamentina (Lamarck 1819) and Amblema plicata (Say 1817). 2. Organic matter concentrations and invertebrate densities in the sediment surrounding mussels were significantly higher in treatments with live mussels than treatments with sham mussels or sediment alone. Organic matter was significantly higher in the sediment surrounding Actinonaias than that surrounding Amblema. Actinonaias was more active than Amblema and may have increased benthic organic matter through bioturbation. 3. Living mussels increased the abundance of periphyton on shells and the abundance and richness of invertebrates on shells, whereas effects of sham mussels were similar to sediment alone. Differences in the amount of periphyton growing on the shells of the two mussel species reflected differences in mussel activity and shell morphology. 4. Differences between living and sham mussel treatments indicate that biological activities of mussels provide ecosystem services to the benthic community beyond the physical habitat provided by shells alone. In treatments containing live mussels we found significant correlations between organic matter and chlorophyll a concentrations in the sediment, organic matter concentrations and invertebrate abundance in the sediment and the amount of chlorophyll a on the sediment and invertebrate abundance. There were no significant correlations among these response variables in control treatments. Thus, in addition to providing biogenic structure as habitat, mussels likely facilitate benthic invertebrates by altering the availability of resources (algae and organic matter) through nutrient excretion and biodeposition. 5. Effects of mussels on sediment and shell periphyton concentrations, organic matter concentrations and invertebrate abundance, varied seasonally, and were strongest in late summer during periods of low water volume, low flow, and high water temperature. 6. Our study demonstrates that freshwater mussels can strongly influence the co‐occurring benthic community, but that effects of mussels are context‐dependent and may vary among species.     

The role of environmental factors in the induction of oxidative stress in zebra mussel (<Emphasis Type="Italic">Dreissena polymorpha</Emphasis>)

The aim of this study was to investigate the influence of specific environmental factors, such as temperature, pH, oxygen concentration, and phosphate, nitrate, chloride, sodium, potassium, sulphate, magnesium and calcium ions concentration, as well as microcystins, on the seasonal variations in the activity of the antioxidant system of the zebra mussel. We examined changes in lipid peroxidation (LPO) levels, glutathione content and the catalase activity of mussels inhabiting the two ecosystems, which differ due to their trophic structure and the presence of toxic cyanobacteria. The results show a relationship between the activity of the antioxidant system of zebra mussels and the seasonal fluctuations of environmental parameters: the symptoms of oxidative stress were generally the highest during spring and the lowest during summer in both ecosystems. Our study also revealed that regardless of the study area the most important factors determining the activity of the antioxidant defences of mussels were the mineral composition (particularly magnesium and calcium ions concentrations) and physical parameters of the water (oxygen concentration and pH). However, factors resulting from the trophic status of studied ecosystems, such as limitations in food resources or high concentration of microcystins during cyanobacterial blooms, were periodically responsible for increased level of LPO in the tissues of zebra mussel. These findings may indicate a limited tolerance of the zebra mussel to the local environmental conditions.     

Nutrient loading associated with agriculture land use dampens the importance of consumer‐mediated niche construction

The linkages between biological communities and ecosystem function remain poorly understood along gradients of human‐induced stressors. We examined how resource provisioning (nutrient recycling), mediated by native freshwater mussels, influences the structure and function of benthic communities by combining observational data and a field experiment. We compared the following: (1) elemental and community composition (algal pigments and macroinvertebates) on live mussel shells and on nearby rocks across a gradient of catchment agriculture and (2) experimental colonisation of benthic communities on live vs. sham shells controlling for initial community composition and colonisation duration. We show that in near pristine systems, nutrient heterogeneity mediated by mussels relates to greater biodiversity of communities, which supports the notion that resource heterogeneity can foster biological diversity. However, with increased nutrients from the catchment, the relevance of mussel‐provisioned nutrients was nearly eliminated. While species can persist in disturbed systems, their functional relevance may be diminished or lost.     

Invasive mussels induce community changes by increasing habitat complexity

Habitat complexity is among the most important factors affecting the diversity, structure, and density of natural communities. The invasive byssate bivalves zebra mussel (Dreissena polymorpha (Pallas, 1771)) and golden mussel (Limnoperna fortunei (Dunker, 1857)) are sessile suspension feeders that form aggregations (druses), physically changing sediments and increasing habitat complexity, and providing shelter and food for other benthic organisms. In this study, we explored the impact of the change in habitat complexity on benthic community created by druses of L. fortunei and D. polymorpha, formed on various sediments. D. polymorpha was studied in Europe (Naroch Lake, Belarus) and in North America (Glen Lake and Lower Nashotah Lake, USA), and L. fortunei was studied in South America (Río Tercero Reservoir, Argentina). Druses of D. polymorpha or L. fortunei and samples of bare nearby sediment (without druses of exotic mussels) were collected at each sampling site. We found significant changes in species richness, density, biomass, taxonomic, and trophic structure of communities formed in druses compared to the nearby bare sediments. Community taxonomic richness increased threefold, and density increased sevenfold with increasing complexity of habitat from sand to druse. The feeding functional group approach indicated that the impact of increased complexity was reinforced by an increase in food supply in D. polymorpha and L. fortunei druses. Along with increasing species richness and densities, byssate bivalves homogenized benthic communities.     

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.     

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.     

Experimental measurements of zebra mussel (Dreissena polymorpha) impacts on phytoplankton community composition

1 . To investigate direct effects of zebra mussel ( Dreissena polymorpha ) feeding activities on phytoplankton community composition, short‐term microcosm experiments were performed in natural water with complex phytoplankton communities. Both gross effects (without resuspension of mussel excretions) and net effects (with resuspension) were studied.
2. Gross clearance rates were not selective; essentially all taxa were removed at similar rates ranging from 24 to 63 mL mussel^–1 h^–1. Net clearance rates were highly selective; different plankton taxa were removed at very different rates, ranging from 12 to 83% of the gross rates, leading to consistent changes in the phytoplankton community composition. Thus, although zebra mussels can cause most phytoplankton to decline, there is considerable variation among taxa in either pre‐digestive selection or post‐digestive survival.
3. The direct, short‐term effects of zebra mussels on phytoplankton community composition are consistent with some of the major changes observed in the Hudson River since establishment of zebra mussels.
4. We show, with simple calculations, how zebra mussel filtration rate, its selective efficiency on various taxa, and phytoplankton growth rates interact to produce changes in the phytoplankton composition.     

Zebra mussels (Dreissena polymorpha) in Ireland, AFLP-fingerprinting and boat traffic both indicate an origin from Britain

1. The zebra mussel (Dreissena polymorpha) is an aquatic nuisance species that invaded Ireland around 1994. We studied the invasion of the zebra mussel combining field surveys and genetic studies, to determine the origin of invasion and the vector of introduction. 2. Field surveys showed that live zebra mussels, attached to the hulls of pleasure boats, were transported from Britain to Ireland. These boats were lifted from British waters onto trailers, transported to Ireland by ferry and lifted into Irish waters within a day. Length‐frequency distributions of dead and living mussels on one vessel imported 3 months earlier revealed a traumatic occurrence caused by the overland, air‐exposed transportation. Results show that a large number of individuals survived after re‐immersion in Irish waters and continued to grow. 3. Zebra mussels from populations in Ireland, Great Britain, the Netherlands, France and North America, were analysed using amplified fragment length polymorphisms (AFLP)‐fingerprinting to determine the origin of the Irish invasion. Phylogenetic analysis revealed that Irish and British mussels clustered closely together, suggesting an introduction from Britain. 4. Ireland remained un‐invaded by the zebra mussel for more than 150 year. The introduction of the zebra mussel to Ireland occurred following the abolition of value added tax in January 1993 on imported second‐hand boats from the European Union (UK and continental Europe). This, together with a favourable monetary exchange rate at that time, may have increased the risk of invasion of the zebra mussel.     

Effects of invasive zebra mussels on phytoplankton,turbidity, and dissolved nutrients in reservoirs

Few experiments have quantified the effects of invasive zebra mussels (Dreissena polymorpha) on man-made reservoirs relative to other aquatic habitats. Reservoirs, however, are the dominate water body type in many of the states that are at the current front of the zebra mussel invasion into the western United States. The objective of this research, therefore, was to determine how zebra mussels affected phytoplankton, turbidity, and dissolved nutrients in water that was collected from three Kansas reservoirs that varied in trophic state (mesotrophic to hypereutrophic), but all experienced frequent cyanobacterial blooms. Laboratory mesocosm experiments were conducted to document the effects of zebra mussels on cyanobacteria and general water quality characteristics in the reservoir water. Zebra mussels significantly reduced algal biomass, and the total biovolume of cyanobacteria (communities were dominated by Anabaena) in each reservoir experiment. The effects of zebra mussels on other major algal groups (diatoms, flagellates, and green algae) and algal diversity were less consistent and varied between the three reservoir experiments. Similarly, the effects of zebra mussels on nutrient concentrations varied between experiments. Zebra mussels increased dissolved phosphorus concentrations in two of the reservoir experiments, but there was no effect of zebra mussels on dissolved phosphorus in the mesotrophic reservoir experiment. Combined, our results strongly suggest that zebra mussels have the potential to significantly impact reservoirs as they continue to expand throughout the western United States. Moreover, the magnitude of these effects may be context dependent and vary depending on the trophic state and/or resident phytoplankton communities of individual reservoirs as has similarly been reported for natural lakes.     

Size‐structured vulnerability of the colonial cyanobacterium,Microcystis aeruginosa,to grazing by zebra mussels (Dreissena polymorpha)

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