Potential effects on zooplankton from species shifts in planktivorous mussels: a field experiment in the St Lawrence River

1. Suspension feeding by bivalves exceeds that by other planktivores in many North American rivers, and food webs may be altered substantially by differences in feeding patterns between native unionid mussels and invading dreissenid mussels. 2. We conducted an experiment comparing zooplanktivory by one unionid and two dreissenid species that addressed several primary questions. Is benthic planktivory important in this river? Has this linkage been altered substantially by dreissenids? Do the two dreissenid species differ in planktivory, and is this ecologically important if quagga mussels extend their geographical range? 3. Our 12‐day experiment consisted of controls (no mussels) and treatments with unionid (Elliptio complanata), quagga (Dreissena bugensis) or zebra (D. polymorpha) mussels in 3500‐L, 80‐μm mesh enclosures placed in a slackwater area of the St Lawrence River. 4. The density of the most abundant calanoid copepod Eurytemora affinis increased in the presence of dreissenids, probably as an indirect food web response. By day 12, a cumulative effect was shown by the most overwhelmingly abundant rotifer, Polyarthra, whose density declined dramatically in dreissenid enclosures compared with control and unionid enclosures. Rotifer densities in unionid enclosures were not different from controls, nor were dreissenid treatments different from each other. The effects on rotifers were probably from predation, as Chl‐a did not vary among treatments. 5. We conclude that benthic‐pelagic coupling via planktivory is important in slackwater areas. Dreissenids have strengthened this linkage, but range extension of quaggas should not appreciably alter effects produced by a similar biomass of zebra mussels.     

Soluble nitrogen and phosphorus excretion of exotic freshwater mussels (Dreissena spp.): potential impacts for nutrient remineralisation in western Lake Erie

1. Recent increases in phytoplankton biomass and the recurrence of cyanobacterial blooms in western Lake Erie, concomitant with a shift from a community dominated by zebra mussels (Dreissena polymorpha) to one dominated by quagga mussels (D. bugensis), led us to test for differences in ammonia‐nitrogen and phosphate‐phosphorus excretion rates of these two species of invasive molluscs. 2. We found significant differences in excretion rate both between size classes within a taxon and between taxa, with zebra mussels generally having greater nutrient excretion rates than quagga mussels. Combining measured excretion rates with measurements of mussel soft‐tissue dry weight and shell length, we developed nutrient excretion equations allowing estimation of nutrient excretion by dreissenids. 3. Comparing dreissenid ammonia and phosphate excretion with that of the crustacean zooplankton, we demonstrated that the mussels add to nitrogen and phosphorus remineralisation, shortening nitrogen and phosphorus turnover times, and, importantly, modify the nitrogen and phosphorus cycles in Lake Erie. The increased nutrient flux from dreissenids may facilitate phytoplankton growth and cyanobacterial blooms in well‐mixed and/or shallow areas of western Lake Erie.     

Invasive mussels modify the cycling,storage and distribution of nutrients and carbon in a large lake

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Invasive ecosystem engineers on soft sediment change the habitat preferences of native mayflies and their availability to predators

1. Dreissenid mussels (quagga mussels, Dreissena bugensis, and zebra mussels, D. polymorpha) are invasive species that function as ecosystem engineers in the Laurentian Great Lakes. Dreissena are increasingly abundant on silt, sand and other soft substrates; by altering benthic habitat, these mussels can alter benthic community structure. 2. We used laboratory mesocosm experiments to examine the effects of soft‐sediment Dreissena clusters on the habitat preference of Hexagenia, a native burrowing mayfly that is an important food source to fish. We conducted three experiments to test whether Hexagenia: (1) select for bare sediment, soft sediment covered with live Dreissena (added structure and food resources) or soft sediment with clusters made of empty Dreissena shells (added structure only), (2) prefer a specific density of live Dreissena on soft sediment and (3) select for or avoid sediment with an accumulation of empty Dreissena shells. 3. Contrary to initial expectations, we found that Hexagenia selected for sediment covered with live Dreissena clusters, followed by empty Dreissena shells clusters, and lastly what was previously thought to be the preferred habitat, bare sediment. Not only did Hexagenia prefer Dreissena‐covered sediment, but they also preferred high densities of Dreissena. 4. We also experimentally tested the effects of Dreissena‐covered soft sediment on the availability of Hexagenia to fish. We had three treatment levels representing three distinct habitat types: (1) bare sediment (no Dreissena) treatment in which water was turbid because of mayfly activity, (2) Dreissena‐covered sediment treatment in which water was clear because of Dreissena filtration and (3) Dreissena‐covered sediment with added turbidity. We found that in low light conditions, similar to many locations where both organisms are found to co‐occur, both yellow perch and round goby consumption of Hexagenia significantly decreased when Dreissena covered the bottom sediment. 5. These results suggest that by choosing Dreissena‐covered habitat, Hexagenia receive protection from fish predation in turbid/low light systems. However, protection from predation cannot be the only reason Hexagenia select Dreissena‐covered sediments, as Hexagenia selected for live clusters more often than empty clusters and may be a result of additional food resources.     

Biological control of streptococcal infection in Nile tilapia Oreochromis niloticus (Linnaeus, 1758) using filter‐feeding bivalve mussel Pilsbryoconcha exilis (Lea, 1838)

Since bivalve mussels are able to graze heavily on bacteria, in this paper it is hypothesized that when mussels are cultured with fish, the filtering efficiency of the mussels will keep the bacterial population below a certain threshold and thus assist in reducing the risk of bacterial disease outbreaks. The ability of the filter‐feeding bivalve mussel Pilsbryoconcha exilis to control Streptococcus agalactiae was tested in a laboratory‐scale tilapia culture system. Juvenile Nile tilapia (Oreochromis niloticus), the bivalve mussel as well as the bacteria were cultured at different combinations using four treatments: treatment‐1: mussel and bacteria but no fish, treatment‐2: tilapia and mussel but no bacteria, treatment‐3: tilapia and bacteria but no mussel, and treatment‐4: tilapia, mussels, and bacteria. All treatments were run in three replicates; stocking rates were 10 tilapia juveniles; five mussels; and about 3.5 × 10⁵ colony forming units (CFU) ml^?1 of bacteria in 50‐L aquaria with 40‐L volume. The mussel reduced the bacterial population by 83.6–87.1% in a 3‐week period whereas in the absence of the mussel, the bacterial counts increased by 31.5%. Oresence of the mussel also resulted in significantly higher growth and lower mortality of tilapia juveniles than when the mussel was absent. The results of this experiment suggest that the freshwater mussel P. exilis could control the population of S. agalactiae in a laboratory‐scale tilapia culture system. Future studies should focus on the dynamic interactions among fish, mussels, and bacteria as well as on how input such as feed and other organic materials affect these interactions.     

Predator size interacts with habitat structure to determine the allometric scaling of the functional response

While both predator body size and prey refuge provided by habitat structure have been established as major factors influencing the functional response (per capita consumption rate as a function of prey density), potential interactions between these factors have rarely been explored. Using a crab predator (Panopeus herbstii) – mussel prey (Brachidontes exustus) system, we examined the allometric scaling of the functional response in oyster (Crassostrea virginica) reef habitat, where crevices within oyster clusters provide mussels refuge from predation. A field survey of mussel distribution showed that mussels attach closer to the cluster periphery at high mussel density, indicating the potential for saturation of the refuge. In functional response experiments, the consumption rate of large crabs was depressed at low prey density relative to small crabs, while at high prey density the reverse was true. Specifically, the attack rate coefficient and handling time both decreased non‐linearly with crab size. An additional manipulation revealed that at low prey densities, the ability of large crabs to maneuver their claws and bodies to extract mussels from crevices was inhibited relative to small crabs by the structured habitat, reducing their attack rate. At high prey densities, crevices were saturated, forcing mussels to the edge of clusters where crabs were only limited by handling time. Our study illuminates a potentially general mechanism where the quality of the prey refuge provided by habitat structure is dependent on the relative size of the predator. Thus anthropogenic influences that alter the natural crab size distribution or degrade reef habitat structure could threaten the long‐term stability of the crab –mussel interaction in reefs.     

Lake morphometry determines Dreissena invasion dynamics

Predicting the ecosystem effects of invasive species and the best control strategies requires understanding population dynamics and population regulation. Invasive bivalves zebra and quagga mussels (Dreissena spp.) are considered the most aggressive invaders in freshwaters and have become major drivers of ecosystem processes in the Laurentian Great Lakes. Combining all lake-wide studies of Dreissena spp. conducted in the Great Lakes, we found that invasion dynamics are largely governed by lake morphometry. Where both species are present, quagga mussels generally become dominant in 8–13 years. Thereafter, zebra mussels remain common in shallow lakes and embayments and lake-wide Dreissena density may remain similar, while in deep lakes quagga led to a near-complete displacement of zebra mussels and an ensuing dramatic increase in overall dreissenid density. In deep lakes, overall Dreissena biomass peaked later and achieved?~?threefold higher levels than in shallow lakes. Comparison with 21 waterbodies in North America and Europe colonized by both dreissenids confirmed that patterns of invasion dynamics found in the Great Lakes are very consistent with other waterbodies, and thus can be generalized to other lakes. Our biophysical model predicted that the long-term reduction in primary producers by mussel grazing may be fourfold less in deep compared to shallow lakes due to thermal stratification and a smaller proportion of the epilimnion in contact with the bottom. While this impact remains greatest in shallow areas, we show that when lakes are vertically well-mixed, dreissenid grazing impact may be greatest offshore, revealing a potentially strong offshore carbon and phosphorus sink.

     

Disentangling interference competition from exploitative competition in a crab–bivalve system using a novel experimental approach

In predator–prey relationships such as those between crabs and their bivalve prey, interference competition is a topic of intense investigation as it can have profound consequences on the dynamics of both predator and prey populations. However in laboratory experiments – also those on crab–bivalve systems – workers never adequately disentangled interference competition from exploitative competition, as prey depletion was never compensated. Hitherto, experimental studies on crab–bivalve systems lack direct behavioural observations and have provided only indirect and thus inconclusive evidence of interference competition. We studied interference competition in adult male shore crabs Carcinus maenas that foraged on blue mussels Mytilus edulis. We developed a novel type of experimental tank to replenish each consumed mussel, and thus to keep prey levels constant. We conducted two experiments in which we varied number of crabs (1, 2, 4) and number of mussels (first experiment: 4, 8, 16, 32; second experiment: 8, 32, 128) and directly observed the foraging behaviour of crabs (foraging area=0.25 m²). In the first experiment, feeding rates decreased with increasing crab density only at mussel density 16 because both search time and time spent in agonistic interactions increased. At other mussel densities, variation in crab density did not affect feeding rates, possibly because of low statistical power and the narrow range of mussel densities offered. In the second experiment feeding rates decreased with increasing crab density because crabs spent more time in agonistic interactions and handling their prey. Feeding rates increased with increasing mussel density. Overall, crabs spent on average 14–18% of their foraging time in agonistic behaviours, while on three out of 64 occasions feeding rates decreased because mussels were stolen (kleptoparasitism). Concluding, we have shown that interference competition occurs in absence of prey depletion, while conducting direct behavioural observations aid to identify the behavioural processes that underlie interference competition.     

Habitat Shift in Invading Species: Zebra and Quagga Mussel Population Characteristics on Shallow Soft Substrates

Unexpected habitat innovations among invading species are illustrated by the expansion of dreissenid mussels across sedimentary environments in shallow water unlike the hard substrates where they are conventionally known. In this note, records of population characteristics of invading zebra (Dreissena polymorpha) and quagga (Dreissena bugensis) mussels from 1994 through 1998 are reported from shallow (less than 20m) sedimentary habitats in western Lake Erie. Haphazard SCUBA collections of these invading species indicated that combined densities of zebra and quagga mussels ranged from 0 to 32,500 individuals per square meter between 1994 and 1998, with D. polymorpha comprising 75–100% of the assemblages. These mixed mussel populations, which were attached by byssal threads to each other and underlying sand-grain sediments, had size–frequency distributions that were typical of colonizing populations on hard substrates. Moreover, the presence of two mussel cohorts within the 1994 samples indicated that these species began expanding onto soft substrates not later than 1992, within 4 years of their initial invasion in western Lake Erie. Such historical data provide baselines for interpreting adaptive innovations, ecological interactions and habitat shifts among the two invading dreissenid mussel species in North America.     

The influence of pre‐settlement and early post‐settlement processes on the adult distribution and relative dominance of two invasive mussel species

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Preface: Blasts from the past and back to the future

The invasion of dreissenid mussels into inland waters of the Northern Hemisphere has received considerable attention and, both zebra mussels and quagga mussels continue to spread westward. Despite studies aimed at understanding the biology of dreissenid mussels, relatively few studies have focused on water velocity and other hydrodynamic characteristics of water flow. The objective of this review was to identify, through a search of online databases, the papers that have been made available that directly have assessed the influence of hydrodynamic characteristics of water flow on dreissenid mussel biology. Using Thompson Reuters Web of Science, Google Scholar, and other resources, 46 papers were identified. These papers detailed that metrics associated with hydrodynamics of water flow, including current, wave action, velocity, flow rate, and discharge, can influence the biology of dreissenid mussels (primarily zebra mussel, which were studied far more than quagga mussel). Hydrodynamic characteristics influenced external fertilization, larval development and settlement, juvenile recruitment and attachment, and suspension feeding, growth and abundance of adults. In most cases, the impact of higher flow rates were locally negative and may present an opportunity for applications of water flow to control the spread or establishment of dreissenid mussels. Several knowledge gaps have been identified.

     

Invasive zebra mussel (Dreissena polymorpha) threatens an exceptionally large population of the depressed river mussel (Pseudanodonta complanata) in a postglacial lake

1. Freshwater mussels are in decline worldwide, with the depressed river mussel Pseudanodonta complanata being one of the rarest and most endangered species in Europe. Invasive mussels are suspected to be an important factor of decline, but there is little information on their interaction with native species.
2. This study analyzed densities, depth distribution, and individual sizes and weights in one of the largest known populations of P. complanata in Europe in relation to the co‐occurring invasive zebra mussel Dreissena polymorpha and other mussel species, using a systematic transect analysis.
3. Pseudanodonta complanata was the dominant unionid species in Lake Siecino reaching densities of up to 26 ind/m², with half of the specimens found at a water depth of 2.0–4.0 m. Densities were highest on sandy substrates in areas of underwater currents. In contrast, 67% of native Unio tumidus were found at depths < 1 m, indicating different habitat preference.
4. In the study area, 91% of P. complanata, 92% of U. tumidus, and all Anodonta individuals were fouled by D. polymorpha. The dreissenid:unionid mass ratio (mean ± SD; maximum) was 0.43 ± 0.56; 4.22 and 0.86 ± 1.87; 8.76 in P. complanata and U. tumidus, respectively. Pseudanodonta complanata fouled with D. polymorpha were impaired in their anchoring capability and had shell deformations potentially affecting shell closing and filtration activity. Fouling intensity was negatively correlated with unionid density, potentially leading to accelerated population declines.
5. The observed adverse effects of invasive zebra mussels on the depressed river mussel and the difficulties in eradicating established populations of invasive mussels suggest that D. polymorpha should be considered a serious threat to P. complanata. Therefore, the further spread of zebra mussels into habitats with native unionids needs to be avoided by all means.

     

Change in diel catchability of young-of-year yellow perch associated with establishment of dreissenid mussels

1. Non‐native mussels have increased water clarity in many lakes and streams in North America and Europe. Diel variation in catchability of some fish species has been linked to visibility during survey trawls (used to measure escapement). 2. Water clarity increased in nearshore areas of western Lake Erie by the early 1990s, following passage of legislation in 1972 to improve water quality (e.g. reduce phosphorus loading) and the invasion of dreissenid mussels (Dreissena spp.) beginning in 1987. 3. We hypothesised that increased water clarity in Lake Erie resulted in decreased catchability of young‐of‐year (age‐0) yellow perch (Perca flavescens Mitchill) during daylight compared to during night. We used a two‐tiered modelling approach to test this hypothesis on the ratio (R) of catch per hour (CPH) during night to CPH during daylight in bottom trawl surveys conducted during 1961–2005. 4. First, we examined seven a priori models. The first model, the ‘null’ model, represented no change in R over time. Three more models tested whether the timing of the change in R was associated with passage of water quality legislation only, dreissenids only (two‐period models) and both legislation and dreissenids (three‐period models). Three additional models included a 3‐year lag before the effects of legislation, dreissenids or both occurred. Secondly, all possible two‐ and three‐period models with a minimum of 2 years per time period were explored a posteriori. The a posteriori procedure determined the temporal transitions to higher R that were best supported by the data, without regard to a priori hypotheses. 5. Night CPH was greater than daylight CPH in 3 of 11 years during 1961–72, in 10 of 15 years during 1973–87, and in 14 of 18 years during 1988–2005. During 1991–2005 night CPH exceeded daylight CPH in all years except one, and night CPH was more than twice daylight CPH in 10 years during this period. 6. The best a priori model had two periods, with a break between 1990 and 1991, corresponding to 3 years after the dreissenid invasion. Similarly, the best two‐ and three‐period a posteriori models both had breaks between 1990 and 1991. The results supported our hypothesis that age‐0 yellow perch exhibited a transition to lower catchability during daylight compared to night, and the timing of the transition coincided with the establishment of dreissenid mussels. 7. The most plausible mechanism for our results was increased visibility of the trawl during daylight, resulting in increased avoidance of the trawl. These results have potential applications wherever non‐native mussels have increased water clarity.     

Transformation of the offshore benthic community in Lake Michigan: recent shift from the native amphipod Diporeia spp. to the invasive mussel Dreissena rostriformis bugensis

1. The native amphipod Diporeia spp. was once the dominant benthic organism in Lake Michigan and served as an important pathway of energy flow from lower to upper trophic levels. Lake‐wide surveys were conducted in 1994/1995, 2000 and 2005, and abundances of Diporeia and the invasive bivalves Dreissena polymorpha (zebra mussel) and Dreissena rostriformis bugensis (quagga mussel) were assessed. In addition, more frequent surveys were conducted in the southern region of the lake between 1980 and 2007 to augment trend interpretation. 2. Between 1994/1995 and 2005, lake‐wide density of Diporeia declined from 5365 to 329 m⁻², and biomass (dry weight, DW) declined from 3.9 to 0.4 g DW m⁻². The percentage of all sites with no Diporeia increased over time: 1.1% in 1994/1995, 21.7% in 2000 and 66.9% in 2005. On the other hand, total dreissenid density increased from 173 to 8816 m⁻², and total biomass increased from 0.4 to 28.6 g DW m⁻². Over this 10‐year time period, D. r. bugensis displaced D. polymorpha as the dominant dreissenid, comprising 97.7% of the total population in 2005. In 2007, Diporeia was rarely found at depths shallower than 90 m and continued to decline at greater depths, whereas densities of D. r. bugensis continued to increase at depths greater than 50 m. 3. The decline in Diporeia occurred progressively from shallow to deep regions, and was temporally coincident with the expansion of D. polymorpha in nearshore waters followed by the expansion of D. r. bugensis in offshore waters. In addition, Diporeia density was negatively related to dreissenid density within and across depth intervals; the latter result indicated that dreissenids in shallow waters remotely influenced Diporeia in deep waters. 4. With the loss of Diporeia and increase in D. r. bugensis, the benthic community has become a major energy sink rather that a pathway to upper trophic levels. With this replacement of dominant taxa, we estimate that the relative benthic energy pool increased from 17 to 109 kcal m⁻² between 1994/1995 and 2005, and to 342 kcal m⁻² by 2007. We project that previously observed impacts on fish populations will continue and become more pronounced as the D. r. bugensis population continues to expand in deeper waters.     

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

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Stoichiometry of regenerated nutrients differs between native and invasive freshwater mussels with implications for algal growth

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Investigation of the effects of current velocity on mussel feeding and mussel bed stability using an annular flume

An annular flume was used to measure the effect of increasing current velocity on mussel (Mytilus edulis) feeding rate and the stability of mussel beds sampled from the mouth of the Exe estuary (SW England). It was found that, in contrast to earlier flume studies, the feeding rates of mussels from open coast sites were unaffected by current velocities up to 0.8 m s^–1. Algal cell depletion in the water column above mussels was a function of current velocity, increasing with declining currents below 0.05 m s^–1. The erodability/stability of the mussel bed, measured in terms of critical erosion velocity, sediment mass eroded and mean erosion rate, was found to be a function of the nature of the substrate and the density of the mussels. Erosion of mussel beds on sandy substrate showed a non-linear relationship with mussel bed density. In comparison with the sand (0% mussel cover), sediment resuspension was about five and four times higher for 25% and 50% cover, respectively. This was due to the increased turbulence and scouring around the clumps of mussels in low-density parts of the bed, and this resulted in some mussels detaching from the bed. At ~100% mussel cover, the sandy bed was more protected by the dense surface layer of mussels, and none became detached during erosion due to the high number of byssal attachments between individuals. The sediment resuspension from the 100% mussel cover was about three times lower than the 0% cover. Erosion of the bed with 50% cover resulted in burial of a large proportion of the mussels, with a 6 cm increase in sediment level. However, the mussels returned to the surface and recovered in 1–2 days, due to a combination of migration upwards and substrate settlement. Channels on the edge of the main Exmouth mussel bed were characterised by a more stable substrate comprising pebbles and sand with varying mussel densities. At these sites, where mussels experience high current velocities on spring tides (up to 0.9 m s^–1), there was no difference between the erodability of pebble/sand substrate with 0% and 100% mussel cover. The sediment erosion was also lower than the 100% mussel cover on the sandy substrate, particularly at currents >0.4 m s^–1. Sampling of different parts of the mussel bed at Exmouth showed mussels at low densities were made up of smaller clumps with a lower mass ratio of mussels to attached substrate (pebbles/sand), thus providing a greater degree of anchorage. Electronic Publication     

The effects of predation and unionid burrowing on bivalve communities in a Laurentian Great Lake coastal wetland

Unionid (Mollusca: Unionidae) densities have declined dramatically throughout the Laurentian Great Lakes after the introduction of dreissenid mussels (Mollusca: Dreissenidae). Recent surveys in some Great Lake coastal wetlands have found abundant unionid populations, but the factors that reduce zebra mussels on unionids in these habitats are not well understood. In 2001–2002, we tested effects of predation and unionid burrowing on corbiculids, sphaeriids and dreissenids in a Great Lake coastal wetland in western Lake Erie. In one experiment, we reduced access by molluscivores using exclosures with two mesh sizes (1.3 cm × 1.3 cm; 5 cm × 10 cm) and sampled bivalves after 15 months. Small mesh exclosures had higher numbers of dreissenids, Corbicula fluminea and sphaeriids (54.9, 3.8, 22.6 individuals/m², respectively) than large mesh exclosures (0.0, 1.13, 0.13 individuals/m², respectively) or open controls (0.3, 1.0, 0.1 individuals/m², respectively). Numbers of dreissenids on C. fluminea were higher in small mesh exclosures (3.8 dreissenids/Corbicula) than in large mesh exclosures (0.1 dreissenids/Corbicula) or cageless controls (0␣dreissenids/Corbicula). In a second experiment, we held two species of live unionids (Leptodea fragilis, Quadrula quadrula) and immobile Pyganodon grandis shells in exclosures (2.5 cm × 2.5 cm mesh) with either 5 cm, 10 cm, or 20 cm deep sediments and sampled bivalves after 2 months. There were fewer dreissenids on L. fragilis than P. grandis shells, but there was no difference in the number of dreissenids on Q. quadrula and P. grandis shells. Numbers of attached dreissenids were higher inside (189–494 dreissenids/unionid) than outside (8–11 dreissenids/unionid) exclosures, and densities of sphaeriid and C. fluminea clams were also higher inside (21.8, 4.7 individuals/m², respectively) than outside (0.4, 0.9 individuals/m², respectively) exclosures. Numbers of attached dreissenids were higher on unionids that could burrow below the sediments (20 cm depth) than unionids in shallow sediments (5 cm depth) for unexplained reasons. Our data suggest that molluscivores can play a pivotal role in limiting numbers of bivalves including dreissenids in coastal wetlands.     

Effectiveness of zebra mussels to act as shelters from fish predators differs between native and invasive amphipod prey

Biological invasions cause organisms to face new predators, but also supply new anti-predator shelters provided by alien ecosystem engineers. We checked the level of anti-predator protection provided to three gammarid species by an invasive Ponto-Caspian zebra mussel Dreissena polymorpha, known for its habitat modification abilities. We used gammarids differing in their origin and level of association with mussels: Ponto-Caspian aliens Dikerogammarus villosus (commonly occurring in mussel beds) and Pontogammarus robustoides (not associated with mussels), as well as native European Gammarus fossarum (not co-occurring with dreissenids). The gammarids were exposed to predation of two fish species: the racer goby Babka gymnotrachelus (Ponto-Caspian) and Amur sleeper Perccottus glenii (Eastern Asian). This set of organisms allowed us to check whether the origin and level of association with mussels of both prey and predators affect the ability of gammarids to utilize zebra mussel beds as shelters. We tested gammarid survival in the presence of fish and one of five substrata: sand, macrophytes, stones, living mussels and empty mussel valves. D. villosus survived better than its congeners on all substrata, and its survival was highest in living dreissenids. The survival of the other gammarids was similar on all substrata. Both fish species exhibited similar predation efficiency. Thus, D. villosus, whose affinity to dreissenids has already been established, utilizes them as protection from fish predators, including allopatric predators, more efficiently than other amphipods. Therefore, the presence of dreissenids in areas invaded by D. villosus is likely to help the invader establish itself in a new place.     

Do Zebra Mussels Grow Faster on Live Unionids than on Inanimate Substrate? A Study with Field Enclosures

The zebra mussel Dreissena polymorpha has invaded numerous freshwaters in Europe and North America and can foul many types of solid substrates, including unionid bivalves. In field experiments we compared growth rates of dreissenids on live specimens of the freshwater bivalve Anodonta cygnea to growth rates of dreissenids on stones. Dreissena density in the study lake was about 1000 m^–2 in most places, Anodonta density approximately 1 m^–2 and about 50% of the Anodonta were infested with 10–30 Dreissena . In summer/autumn small dreissenids generally grew faster on live Anodonta than on stones. Similar trends were observed for spring, but differences of growth increments between dreissenids on live Anodonta and stones were usually not significant. Dreissenids settled down or moved towards the ingestion/egestion siphons of Anodonta and ingestion siphons of dreissenids were directed towards siphons of Anodonta . These results suggest that dreissenids can use the food provided by the filter current of the large Anodonta . (© 2006 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)