Predicting habitat use and trophic interactions of Eurasian ruffe,round gobies,and zebra mussels in nearshore areas of the Great Lakes

The Laurentian Great Lakes have been subject to numerous introductions of nonindigenous species, including two recent benthic fish invaders, Eurasian ruffe (Gymnocephalus cernuus) and round gobies (Neogobius melanostomus), as well as the benthic bivalve, zebra mussel (Dreissena polymorpha). These three exotic species, or “exotic triad,” may impact nearshore benthic communities due to their locally high abundances and expanding distributions. Laboratory experiments were conducted to determine (1) whether ruffe and gobies may compete for habitat and invertebrate food in benthic environments, and (2) if zebra mussels can alter those competitive relationships by serving as an alternate food source for gobies. In laboratory mesocosms, both gobies and ruffe preferred cobble and macrophyte areas to open sand either when alone or in sympatry. In a 9-week goby–ruffe competition experiment simulating an invasion scenario with a limited food base, gobies grew faster than did ruffe, suggesting that gobies may be competitively superior at low resource levels. When zebra mussels were added in a short-term experiment, the presence or absence of mussels did not affect goby or ruffe growth, as few zebra mussels were consumed. This finding, along with other laboratory evidence, suggests that gobies may prefer soft-bodied invertebrate prey over zebra mussels. Studies of interactions among the “exotic triad”, combined with continued surveillance, may help Great Lakes fisheries managers to predict future population sizes and distributions of these invasive fish, evaluate their impacts on native food webs, and direct possible control measures to appropriate species.     

Predation by the invasive American signal crayfish, Pacifastacus leniusculus Dana, on the invasive zebra mussel, Dreissena polymorpha Pallas: the potential for control and facilitation

Non-indigenous crayfish often have major ecological impacts on invaded water bodies, and have contributed to the decline of native crayfish species throughout Europe. The American signal crayfish, Pacifastacus leniusculus, is the most widespread invasive crayfish in Great Britain, where the zebra mussel, Dreissena polymorpha, is similarly an invasive pest species. The potential for the American signal crayfish to regulate zebra mussel populations was investigated through a series of laboratory experiments. Crayfish were found to be highly size selective, consuming significantly more of the smallest size class of zebra mussels offered (7–12 mm), over medium (16–21 mm) and large (25–30 mm). Crayfish feeding rate on zebra mussels was not altered when mussels were presented clumped together in natural druses compared with mussels in a disassembled druse. Crayfish spent significantly more time foraging when mussels were unattached, and a greater proportion of attacks were on medium and large than on small mussels (83% of attacks were on medium and large mussels when unattached as opposed to 47% when on druses). Individual crayfish feeding rate decreased significantly at densities of > ~5 crayfish m⁻². Signal crayfish are, therefore, unlikely to be able to significantly impact established populations of zebra mussels in the wild, although zebra mussels have the potential to provide crayfish with a substantial food source.     

Associations between food habits and pharyngeal morphology in the round goby (<Emphasis Type="Italic">Neogobius melanostomus</Emphasis>)

Several studies have shown that round gobies (Neogobius melanostomus) undergo a dietary shift from arthropods to dreissenid mussels as they grow, and this shift is accompanied by changes in pharyngeal morphology associated with durophagy. In contrast, some populations of round gobies prey heavily on various arthropods, but it is unknown whether those populations develop pharyngeal morphology associated with durophagy or if they develop less robust and molarized structures. To test if there is a relationship between food habits and pharyngeal morphology, we compared those characters in round gobies from two sites in Erie Co., Pennsylvania: a dreissenid-present site (Presque Isle Bay, PIB, of Lake Erie) and a dreissenid-absent site (Fairview Gravel Pit, GP). Multivariate analysis of covariance (MANCOVA) revealed consistent ontogenetic changes at both sites in which lower pharyngeals of larger fish were more robust, possessed wider teeth, and had a greater area taken up by large diameter teeth. Pharyngeal morphology also differed between sites, with gobies from PIB having wider and more robust lower pharyngeals. Food habits differed markedly between round gobies from the two sites, with dreissenid mussels being the most important prey item for all length classes in PIB, and crustaceans being the most important prey item for all length classes at the GP. Canonical correlation analysis on all round gobies revealed a correlation between consumption of dreissenid prey and pharyngeal characters associated with durophagy. Although food habits and pharyngeal morphology of round gobies appear to be associated, the mechanism responsible for the association (i.e. phenotypic plasticity versus local adaptation) is not clear.     

Has the invasive round goby caused new links in Baltic food webs?

The Ponto-Caspian round goby (Neogobius melanostomus, Pallas 1814) most probably was established in the Gulf of Gdańsk, Baltic Sea, in the late 1980’s and has since become one of the dominant species in the region. In this study we assess the role of round gobies as prey for two important fish species in the Gulf of Gdańsk, cod (Gadus morhua) and perch (Perca fluviatilis). We compared their present diet with stomach analyses from the area prior the round goby establishment, as well as with diet analysis from Baltic regions where round gobies are absent. There were large differences in the diet between cods from the Gulf of Gdańsk 2003–2006 compared to cods in earlier studies (1977–1981) from the Southern Baltic Sea. There were also large differences in cod and perch diets from areas with and without round goby. Presently, round goby constitutes the most important prey for medium sized cods in Gulf of Gdańsk, and perch from the same area almost exclusively feed on gobiids. Stomach analysis, trophic level estimates, and stable isotope analyses all indicated that cod and perch in Gulf of Gdańsk after the round goby establishment belonged to a similar trophic level. Beside round goby, no mussel feeding fish contributed much to the diet of cod or at all to the diet of perch. Thus, it is likely that round gobies constitute a new energetic pathway from mussels to top predators. However, due to the short time elapsed after round goby establishment, we can only speculate on the species future impacts on Baltic food webs.     

Predation on invasive zebra mussel, Dreissena polymorpha, by pumpkinseed sunfish, rusty crayfish, and round goby

The enemy release hypothesis states that invasive species are successful in their new environment because native species are not adapted to utilize the invasive. If true for predators, native predators should have lower feeding rates on the invasive species than a predator from the native range of the invasive species. We tested this hypothesis for zebra mussel (Dreissena polymorpha) by comparing handling time and predation rate on zebra mussels in the laboratory by two North American species (pumpkinseed, Lepomis gibbosus, and rusty crayfish, Orconectes rusticus) and one predator with a long evolutionary history with zebra mussels (round goby, Neogobius melanostomus). Handling time per mussel (7 mm shell length) ranged from 25 to >70 s for the three predator species. Feeding rates on attached zebra mussels were higher for round goby than the two native predators. Medium and large gobies consumed 50–67 zebra mussels attached to stones in 24 h, whereas pumpkinseed and rusty crayfish consumed <11. This supports the hypothesis that the rapid spread of zebra mussels in North America was facilitated by low predation rates from the existing native predators. At these predation rates and realistic goby abundance estimates, round goby could affect zebra mussel abundance in some lakes.     

Feeding ecology of the Bay Goby (Pisces: Gobiidae). Effects of behavioral,ontogenetic, and temporal variation on diet

The ontogenetic, diel, seasonal, and yearly variations in gut fullness, diet, and prey diversity for a California estuarine gobiid (Lepidogobius lepidus Girard) were examined. Also the feeding behavior of this species was described.Small (<50 mm, SL) and large (? 50 mm, SL) gobies consumed similar prey types in different proportions. Major prey items were polychaetes, harpacticoid copepods, gammarid amphipods, molluscs, and other crustaceans. Diets of large and small gobies were not significantly correlated, and larger fish had a more diverse diet. Small fish fed at all times while larger gobies fed primarily at night. Changes in diet may be related to differential prey preferences, feeding chronologies, and increases in fish size.Both large and small gobies displayed seasonal differences in diet and prey diversity. Year-to-year changes in diet also were noted for both size classes. The bay goby uses different feeding behaviors to capture sedentary and motile prey and appears to forage opportunistically. This behavior is probably advantageous in an environment which fluctuates drastically.     

Diet composition and consumption rate in round goby (<Emphasis Type="BoldItalic">Neogobius melanostomus</Emphasis>) in its expansion phase in the Trent River,Ontario

We assessed density, gut fullness and prey composition of round goby (Neogobius melanostomus) from three areas in the Trent River (Ontario) representing areas of initial introduction and subsequent expansion. Round goby had been present at the area of original introduction since 2003, and by 2007/2008, their range had expanded upstream and downstream into the outermost reaches sampled in the study. Catch per unit angling effort in nearshore sites indicated that round goby density in the area of original introduction was more than double their density in the upstream expansion area and nearly three times the density in the downstream expansion area. Gut fullness index was lower in gobies from the area of original introduction than for those at the upstream and downstream edges of their expanded range. The most dramatic difference in diet composition was with dreissenids, where large gobies (≥70 mm) occupying the area of original introduction had almost no dreissenid biomass in their guts, whereas dreissenids were the predominant prey type in gobies occupying the two expansion areas. Post-hoc zebra mussel density in the area of original introduction was an order of magnitude lower than in the two expansion areas which, combined with the differences in stomach fullness and prey composition, suggest that local, density-related reduction of this prey type was occurring in the river.     

Diel interactions between prey behaviour and feeding in an invasive fish, the round goby, in a North American river

1. We studied the diet of the invasive round goby (Neogobius melanostomus) on a diel basis in the Flint River, a warmwater stream in Michigan, U.S.A. Diet and available prey samples were collected seven times over a 24 h period in four consecutive months. The section of river studied lacked zebra mussels (Dreissena polymorpha), the primary prey of adult round gobies elsewhere in the Great Lakes region. 2. Diet changed on a diel basis with hydropsychid caddisfly and chironomid larvae predominating during the day, chironomid pupae dominating in the evening and heptageniid mayflies dominating at night. Simultaneous study of macroinvertebrate drift suggested that caddisfly and chironomid larvae were most likely picked from submerged rocks, chironomid pupae were most likely taken during their emergent ascent and mayflies were either captured from the drift or picked from rocks. 3. The Flint River lacks a diverse darter (Family: Percidae) and sculpin (Family: Cottidae) fauna and it appears that the round goby has occupied a generalised darter/sculpin niche. Our results indicate that round gobies have the potential to invade successfully riverine systems, particularly those lacking a diverse benthic fish assemblage.     

Temporal and ontogenetic variation in the diet of squid (<Emphasis Type="Italic">Loligo forbesii</Emphasis> Streenstrup) in Scottish waters

The squid Loligo forbesii is the only cephalopod species currently targeted by fisheries in the northern NE Atlantic. An active predator, it feeds primarily on fish, crustaceans and cephalopods. During 15 years since the only previous large-scale study of the diet of this species in Scottish waters, there have been substantial changes in marine fish abundances. The present study evaluates sources of variation (temporal, ontogenetic) in diet composition and prey size preferences of L. forbesii, including a comparison of contemporary (July 2006–June 2007) and historical (1990–1992) dietary datasets. Results revealed significant size-related and seasonal variation in diet composition and prey size. Teleost fish of the families Ammodytidae and Gobiidae were eaten by squid of all sampled sizes, although occurrence of gobies was generally more frequent in smaller squids, while Gadidae were eaten more frequently by larger squids. Cannibalism was also more frequent in larger squids. Compared to the 1990–1992 dataset, clupeid fish were less important in the diet of squid in 2006–2007, while the importance of gobies increased, and the size of gobies eaten also increased. The trend in gadoids differed according to the index used: their frequency of occurrence was considerably higher in 2006–2007 than in 1990–1992, but their numerical importance was slightly lower. In general, results provided little evidence that changes in the diet of L. forbesii correspond with changes in fish abundance, at least at the scales at which these are measured.     

Size-selective predation by roach (Rutilus rutilus) on zebra mussel (Dreissena polymorpha): field stuides

Summary Studies of predation by roach (Rutilus rutilus) on zebra mussel (Dreissena polymorpha) in a large, eutrophic lake showed that there was a clearly marked size threshold ( 160 mm SL) above which roach began to feed on mussels. Roach preying on various sizes of mussels selected them in proportions different from their abundance and accessibility in the habitat. The mean size of mussels ingested by roach of 220 mm and larger, which fed predominantly on Dreissena, closely followed the pattern expected for a constant ratio of mean prey size to mean predator mouth size = 0.59. To explain the size selection we applied an optimal foraging approach, based on the ability of different-sized fish to crush (cost) mussels of different sizes, and hence crushing resistance, and energy contents (benefit). We found that fish smaller than 160 mm, which showed no inclination to eat Dreissena, would only be able to take small mussels with a very high cost/benefit ratio. The real switch to Dreissena would be expected in fish of 230–240 mm that could take most of their prey from highly profitable, numerous, and easily accessible size classes while keeping the mean prey size at the optimal level relative to mean predator mouth size.     

Diet-shifts by an estuarine goby (Pomatoschistus microps) in the face of variable prey availability

Ontogenetic shifts in diet provide a mechanism for maximising fitness throughout development and are common where predators exhibit large increases in size. In order to maximise their fitness throughout development, benthic feeding fish can show diet-shifts that centre on the transition from meiofaunal to macrofaunal prey. Here we assessed whether such a shift was influenced by natural variation in prey-size availability by comparing the sizes of prey consumed by naturally foraging common gobies (Pomatoschistus microps). We tested explicitly for the presence of an ontogenetic shift by analysing the length of prey consumed and an index for prey importance for gobies of different lengths. We also tested the match between actual diets and those predicted by a foraging model. The goby size at which the diet-shift occurred was consistent among locations that differed in their availability of prey and through temporal changes in densities and types of prey. The mean sizes of ingested prey increased for gobies > 35 mm in length and the relative importance of macrofauna increased at 30 mm. The foraging model predicted that gobies > 30 mm would eat larger prey than would smaller gobies which differed from the observed changes in prey-size at 35 mm. Availability of prey did not appear to influence the lengths at which gobies changed diet but did affect the size of prey taken after the diet-shift. A relatively large abundance of large-bodied chironomids at two sites was reflected in the mean size of prey consumed by gobies > 30 mm at these sites. Our study indicates that intrinsic mechanisms can be more important than fluctuating environments in determining prey-choice and shifts in diet, although for the common goby, variability in prey-size may have implications for prey-choice later in ontogeny.     

An indirect effect of biological invasions: the effect of zebra mussel fouling on parasitisation of unionid mussels by bitterling fish

Invasive species represent a major threat with both direct and indirect effects on natural ecosystems, including effects on established and coevolved relationships. In a series of experiments, we examined how the interaction between two native species, a unionid mussel (Unio pictorum) and the European bitterling (Rhodeus amarus), a fish that parasitises unionids, was affected by the non-native zebra mussel (Dreissena polymorpha). The zebra mussel fouls hard substrates, including shells of living unionids, and its presence is often associated with a decrease in population density of native unionid mussels. Bitterling lay their eggs into live unionids and the embryos develop inside their gills. Using a range of zebra mussel densities, we demonstrated that zebra mussel fouling had a negative effect on the number of bitterling eggs inside the mussel host, with abundances of 5–10 zebra mussels (shell size 15–25 mm) per unionid critical for bitterling ability to utilise the host. In a further experiment, we found that bitterling did not discriminate between unfouled unionids and those fouled with five zebra mussels. Most ovipositions into fouled hosts, however, were unsuccessful as eggs failed to reach the unionid gills. We discuss implications of such unsuccessful ovipositions for bitterling recruitment and population dynamics.     

Retention of N and P by zebra mussels (<Emphasis Type="Italic">Dreissena polymorpha</Emphasis> Pallas) and its quantitative role in the nutrient budget of eutrophic Lake Ekoln,Sweden

We quantified cover, population densities, size distribution and biomass of zebra mussels along 7 transects in eutrophic Lake Ekoln (Sweden). We also analyzed the elemental (C, N, P) composition of zebra mussel soft tissue and computed their retention rates of N and P their quantitative role in the lake’s nutrient budget. We hypothesized that zebra mussels play an important role in the nutrient budget of the lake and speculate that the successive harvesting of cultured mussels could contribute to the lake’s rate of recovery from cultural eutrophication. At depths exceeding 5 m, mussels covered consistently less than 5% or were absent. Similarly, mean densities were 3,158 ± 2,143 ind m⁻² between 2 and 4 m, but rapidly declined at larger depths. Calculated clearance rates averaged 19.4 ± 2.3 km³ y⁻¹, implying the entire lake is filtered every 8–10 days. Concentrations of N and P in mussel soft tissue averaged 100.9 ± 1.5 mg N g⁻¹ DW and 9.3 ± 0.2 mg P g⁻¹ DW. The lake population was estimated to 22.2 ± 2.6 × 10¹⁰ mussels, corresponding to a standing stock biomass of 362 ± 42 ton DW, or conservative estimates of 36.6 ± 4.3 ton N and 3.4 ± 0.4 ton P. Assuming a life span of 2–3 years gives a retention estimate of 1.2–1.8 ton P y⁻¹ by mussels, corresponding to 50–77% of the annual P influx from Uppsala sewage treatment plant to the lake. Similarly, annual N-retention by zebra mussels makes up 13–20 ton N y⁻¹, largely equaling the annual N-deposition from atmospheric sources on the lake’s surface. These retention rates correspond to only a few percent of the annual P-load from agricultural sources, but we argue that the quantitative role of zebra mussels in nutrient budgets is much larger if these budgets are adjusted for the bias introduced by coarse estimates of N and P pools that include a large share of refractory P.     

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.     

Predatory blue crabs induce byssal thread production in hooked mussels

Abstract. Blue crabs (Callinectes sapidus) prey on hooked mussels (Ischadium recurvum) growing epizoically on oyster clumps in estuaries along the Louisiana coast. In prey size‐selection experiments, blue crabs preferred small mussels (<30‐mm shell length) to larger mussels, possibly because handling time increased with mussel size. When crabs were given a choice of solitary mussels versus mussels in clumps on oysters in the laboratory, mortality was lower by 86% in clumped mussels. However, no size selection by crabs occurred with mussels in clumps, likely because smaller mussels escaped predation in crevices between larger mussels or oysters. When individuals of two size classes of mussels were exposed to water containing the scent of crabs and of mussels consumed by blue crabs, an increase in byssal thread production was induced in all mussels, but byssal thread production rate was higher for small mussels than for large mussels. We conclude that increased predation risk for small mussels has resulted in higher size‐specific production of byssal threads, and that predator‐induced production of byssal threads, which may increase clumping behavior, may reduce their risk of mortality to predatory blue crabs.     

Testing a typology system of running waters for conservation planning in Hungary

The zebra mussel (Dreissena polymorpha) and its congener the quagga mussel (Dreissena rostriformis bugensis) are both invaders in freshwater, but have very different invasion histories, with zebra mussels attaining substantially faster rates of spread at virtually all spatial scales. However, in waterbodies where they co-occur, D. r. bugensis can displace D. polymorpha. To determine if the mechanisms for this displacement are associated with different survival and growth, we kept mussels in flow-through tanks for 289 days with two temperature regimes that mimicked the natural surface water (littoral zone) and hypolimnion conditions of Lake Erie. For the littoral zone regime, we used water directly from the surface of Lake Erie (range 4–25°C, average 11.9 ± 0.6°C). For the profundal zone treatment, Lake Erie surface water was chilled to about 6°C (range 5–8°C, average 6.2 ± 0.6°C) for the full duration of the experiment. For each of these temperature regimes, we used three replicate tanks with only zebra mussels present and three replicate tanks with only quagga mussels (150 ind./tank each), and three replicate tanks with both species (75 ind./tank of each species). Quagga mussels had higher survivorship and grew more than zebra mussels in all treatments. For both species, the size of the mussel entering the winter was critical for survivorship. Larger mussels had a higher survival over the winter in all treatments. For both species, there was a survivorship and growth tradeoff. In the warmer littoral zone treatment both species had higher growth, but lower survival than in the colder profundal zone treatment. Surprisingly, although quagga mussels outperformed zebra mussels, zebra mussel survivorship was better when they were faced with competition by quagga mussels than with just intraspecific competition. In addition, quagga mussels suffered size-specific mortality during the growing season only when facing interspecific competition with zebra mussels. Further experiments are needed to determine the possible mechanisms for these interspecific effects.     

Ontogenetic changes in habitat use by whitefish,Coregonus lavaretus

Synopsis The whitefish, Coregonus lavaretus, in the lake Mj?sa exhibited two niche changes during their life cycle. Juveniles (< 25 cm body length) were confined to the shallow (0–30 m) epibenthic zone. Medium sized whitefish (25–35 cm body length) expanded their habitat use to include the deep (30–90 m) epibenthic zone as well as the pelagic zone. From a body length of 35 cm, habitat use was restricted to the deep epibenthic zone. Small fish in the shallow epibenthic zone ate small and medium-sized prey (zooplankton, insect larvae and surface insects). Medium-sized fish in this zone were in addition feeding on the larger amphipod, Pallasea quadrispinosa. In the pelagic zone, the diet of medium-sized whitefish was dominated by zooplankton, although some larger prey like surface insects and age-0 smelt, Osmerus eperlanus, were also eaten. In the deep epibenthic zone, the diet of both medium-sized and large (< 35 cm) whitefish consisted mainly of the large prey P. quadrispinosa.     

Northern map turtles (Graptemys geographica) derive energy from the pelagic pathway through predation on zebra mussels (Dreissena polymorpha)

1. Zebra mussels ( Dreissena polymorpha ) derive their energy from the pelagic energy pathway by filtering plankton. Because zebra mussels occur in high densities in littoral habitats, they potentially constitute an important trophic link between littoral consumers and pelagic energy sources. Northern map turtles ( Graptemys geographica ) are widespread in North America and consume zebra mussels.
2. We used stable isotopes analyses to quantify the flow of energy from the pelagic pathway to northern map turtles and to infer the contribution of zebra mussels to map turtle biomass. We then built a bioenergetic model to estimate the annual intake of zebra mussels by northern map turtles in Lake Opinicon, Ontario, Canada.
3. Stable isotopes analyses indicated that zebra mussels constitute between 0% and 14% of the diet of males and between 4% and 36% of the diet of females. Assuming that zebra mussels account for all of the pelagic contribution, we estimated that map turtles consume 3200 kg of zebra mussels annually. Because female map turtles are much larger than males and consume more zebra mussels, they are responsible for 95% of the zebra mussel biomass ingested annually.
4. The pelagic pathway supports an important part of the standing crop biomass of map turtles in Lake Opinicon. We highlight the importance of freshwater turtles in lake ecosystems. Unravelling the trophic interactions mediated by freshwater turtles will lead to a more integrated picture of lake ecosystems.     

In situ experiments on predatory regulation of a bivalve mollusc (Dreissena polymorpha) in the Mississippi and Ohio Rivers

1. In situ exclosure experiments in the Mississippi and Ohio Rivers determined the importance of fish predation in regulating zebra mussels (Dreissena polymorpha), an increasingly important constituent of the benthic invertebrate assemblages in both rivers. 2. We evaluated the effects of predatory fish on the density, biomass and size distribution of zebra mussels in a floodplain reach of the upper Mississippi River and in a naturally constrained reach of the Ohio River. Fifty, six-sided, predator-exclusion cages and fifty ‘partial’ cages (mesh at the upstream end only) were deployed, with half the cages containing willow snags and half clay tiles suspended 12–16 cm above the bottom. A single snag or tile sample unit was removed from each cage at approximately monthly intervals from July to October 1994. Types and relative abundances of molluscivorous fish were evaluated by electrofishing near the cages in both rivers. Actual and potential recruitment of young zebra mussels on to the substrata were measured using benthic samples in both rivers and estimated (Ohio River only) from counts of planktonic veligers. 3. Zebra mussels were consumed by at least three fish species in the upper Mississippi River (mostly carp, Cyprinus carpio, and redhorse suckers, Moxostoma sp.) and five species in the Ohio River (primarily smallmouth buffalo, Ictiobus bubalus, and channel catfish, Ictalurus punctatus), but potential recruitment seemed adequate to replace consumed mussels, at least in the Ohio River. The number of juvenile benthic mussels showed no apparent link with the density of veligers soon after initiation of reproduction. Recruitment of juveniles on snags and tiles was not affected by cage type (thus eliminating a potentially confounding ‘cage effect’). 4. Fish significantly influenced mussel populations, but the impact was often greatest among low density populations in the upper Mississippi. Density and biomass differed in both rivers for cage type (higher inside cages), substratum (greater on tiles), and date (increased over time). Presumed size-selective predation was present in the Mississippi (greater on larger size classes) but was not evident in the Ohio. We hypothesize that fish in the Mississippi can more easily select larger prey from the low density populations; whereas size-selective predation on tightly packed zebra mussels in the Ohio would be difficult. 5. Although fish can reduce numbers of Dreissena polymorpha in the two rivers, current levels of fish predation seem insufficient to regulate zebra mussel densities because of its great reproductive capacity. The recent invasion of zebra mussels, however, could lead to larger fish populations while promoting greater carbon retention and overall ecosystem secondary production.     

Feeding habits of horse mackerel, Trachurus trachurus (Linneaus, 1758), from the central Adriatic Sea

The feeding habits of the horse mackerel, Trachurus trachurus, from the central Adriatic Sea were investigated with respect to fish size and season. Stomach contents of 1200 specimens 12.9–37.6 cm total length (TL) taken at monthly intervals (January–December 1995) were analyzed. Of the total number of stomachs examined, 597 were empty (49.7%). This percentage varied significantly with season; the maximum number of empty stomachs was observed during January (80%) and February (82%) and a minimum number observed during July (15%) and August (19%). A total of 30 different prey species belonging to five major groups: Crustacea (Euphausiacea, Mysidacea, Decapoda), Cephalopoda, and Teleostei was identified in the stomach contents. Euphausiids were the most important ingested prey in all seasons as well as in small‐ and medium‐size classes (<28 cm TL). Two species of euphausiids, Nyctiphanes couchii (%IRI = 36.0) and Euphausia krohni (%IRI = 12.9), were the most frequent prey. The proportion of teleosts in stomach contents increased with the increasing size class of horse mackerel and was 15.5% IRI in >30 cm TL individuals. The mean weight of stomach contents increased significantly for fish larger than 24.1 cm TL, while the mean number of prey items significantly declined in the large‐size classes.