Feeding preferences of an invasive Ponto‐Caspian goby for native and non‐native gammarid prey

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Sublittoral abundance and food consumption of Baltic gobies

Two small demersal fishes, the sand goby Pomatoschistus minutus and the common goby Pomatoschistus microps , were quantified on soft bottoms at 20–40 m depth in the Baltic Sea, using a camera placed above the bottom. The largest numbers of gobies were seen following the settlement of young in late summer and autumn. Most recorded fishes were sand gobies. An annual average of 4·7 individuals m⁻²(0·24 g dry mass m⁻²) was recorded in 1983–1985 and 2·5 individuals m⁻²(0·13 g m⁻²) in 1997–1998. Using these densities, the annual goby food consumption was estimated to 100 kJ m⁻² in 1983–1985 and 50 kJ m⁻² in 1997–1998, corresponding to most of the annual macrobenthos production available to the gobies. The resulting goby production, assumed equal to 25% of the food consumed, must have been an important food source for the larger fishes occasionally recorded in the photographs.     

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.     

An enemy of your enemy is your friend: Impact of predators on aggregation behavior of gammarids

Predation pressure may affect many aspects of prey behavior, including forming groups and changes in social interactions. We studied the aggregation behavior of competing gammarids Dikerogammarus villosus and Pontogammarus robustoides (Amphipoda, Crustacea) to check whether they modify their preferences for conspecifics or heterospecifics in response to predator (the racer goby Babka gymnotrachelus) kairomones in the presence or absence of stone shelters (alternative protection source). Both species exhibited preferences toward shelters occupied by conspecifics over empty shelters and conspecifics apart from shelters, suggesting that their aggregation depends not only on habitat heterogeneity, but also on their social interactions. Moreover, gammarids in the presence of shelters (safer conditions) preferred conspecifics over heterospecifics, but predator kairomones made them form aggregations irrespective of species. In the predator presence, P. robustoides increased its aggregation level only in the sheltered conditions, whereas D. villosus exhibited this response only in the absence of shelters, suggesting that this behavior can protect it against predators. Therefore, we tested the antipredator effectiveness of D. villosus aggregations by exposing them to fish predation. Gobies foraged most effectively on immobile single gammarids compared to moving and aggregated individuals. Fish also avoided aggregated prey, confirming the protective character of aggregations. We have demonstrated that the predator presence increases aggregation level of prey gammarids and affects their social behavior by reducing antagonistic interactions and avoidance between competing species. This is likely to affect their distribution and functioning in the wild, where predator pressure is a standard situation.     

Diet and feeding habits of monkey goby (Neogobius fluviatilis) in a newly invaded area

The monkey goby (Neogobius fluviatilis) is one of the Ponto-Caspian species that in recent decades made substantial East-to-West invasions in European inland waters. Now the species is present in the Danube as far as its Slovakian section. In the last decade the species also invaded the Western Bug and Vistula rivers, in the latter it is numerous nearly as far as its delta. The purpose of our study was to define the diet spectrum, feeding preferences, spatial and size related changes in diet and diurnal feeding activity of this goby. Another goal was to predict which groups of native prey would be most affected by the presence of this exotic predator in newly invaded areas. The study was carried out at three sampling locations in the Vistula River system. The food spectrum of the monkey goby was broad. The fish consumed insect larvae and pupae, crustaceans, annelids, gastropods and fish. Chironomid larvae were a prevalent food category in all sampling sites, followed by amphipod crustaceans at one site and by trichopteran larvae and chironomid pupae at another. At the third site, there was no distinct subdominant food category. According to the values of the Ivlev’s selectivity index, the preferred food category were chironomid larvae. No significant differences in diet were found over the 24-h cycle. There was no variation among different fish size groups. From our studies and from the available literature data it can be concluded that the species, due to its ability to use locally available food resources, displays a generalistic and highly flexible feeding strategy.     

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.     

Spot-fixed fin digging behavior in foraging of the benthophagous maiden goby, Pterogobius virgo (Perciformes: Gobiidae)

 Several patterns of feeding behaviors have been documented in benthophagous fishes. The foraging behavior of the maiden goby, Pterogobius virgo, was studied at Kurahashi Island in the Seto Inland Sea, Japan. Pterogobius virgo foraged mostly on polychaetes by volume from among several available prey items by digging in the sandy bottom. The digging behavior comprised swing of only pectoral fins or of both pectoral fins and body. Pectoral fin swing exposed the cryptic prey within the bottom, and fins and body swing exposed the prey and washed the sediment away. The swings were repeatedly and continuously conducted at a site during the daytime, making a pit several centimeters deep in which the fish was located. After the prey was exposed, the fish immediately and rapidly picked up the prey. Polychaetes were abundant prey in the sediment, occurring in the layer 3–5 cm deep from the bottom surface in the study area. In this goby, spot-fixed fin digging, the first documentation of feeding habits in gobies, may be effective for feeding on the most valuable prey, i.e., polychaetes, which may be otherwise unavailable for this fish. Received: April 24, 2001 / Revised: April 26, 2002 / Accepted: May 7, 2002     

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.     

The Toxicity of Skin Secretions from Coral-Dwelling Gobies and their Potential Role as a Predator Deterrent

Coral-dwelling gobies in the genus Gobiodon (family Gobiidae) posses toxic skin secretions. We used bioassays to investigate interspecific variation in the toxicity of skin secretions from six species of Gobiodon from Lizard Island on the Great Barrier Reef. We then used feeding experiments with two common species of predatory fish to test if skin secretions might act as a chemical defence against predation. The skin secretions of all species were toxic to the bioassay species, Apogon fragilis, but there were marked differences in toxicity among Gobiodon species. Feeding experiments showed that both small- and large-gaped predators avoided food items to which goby skin secretions, or a whole goby, had been added. These experiments indicate that skin toxins could function as a predator deterrent in coral-dwelling gobies.     

Trophic relationships among concentrated populations of small fishes in seagrass meadows

Changes in available food and utilized foods for a densely populated guild of animals can uncover periods of niche displacement among particular consumer species or their separate size classes. Dense populations of small fishes and their foods were çensussed for 13 months and studied experimentally in shoalgrass and turtlegrass meadows of Redfish Bay, Texas. Feeding habits were determined with respect to prey availability, and illustrated the extent to which seasonal partitioning of food corresponded to food depletion among these abundant consumers in seagrass meadow food webs. The darter goby, code goby, pinfish, and Gulf pipefish were the most common species throughout the year. Although the darter goby did not show distinct ontogenetic changes in food habits, the code goby, Gulf pipefish, and pinfish demonstrated major ontogenetic progressions of foods selected.Food availability in the seagrass meadows changed seasonally. When major prey such as amphipods were abundant, during spring, many fish species showed high overlap in food use. Regardless of food availability, the code goby and Gulf pipefish fed mainly on amphipods or copepods. The more common darter goby and pinfish seasonally changed their diets with changes in food availability; the darter goby and pinfish were more carnivorous during spring, but they largely consumed epiphytic algae during summer. Cage experiments were used to monitor foods confined with elevated densities of pinfish and darter gobies, relative to control cages at natural overall densities. Prey items in the former cages decreased sharply, with corresponding dietary shifts by these common fishes. During resource depletion, these changes in resource use by these naturally concentrated consumers appeared as temporary partitioning of available resources. This shift occurred during both natural (seasonal) and experimental depletions of food, and appears to result from increased interspecific and intraspecific competition during periods of depleted preferred foods.     

Comparative Ecology of the Gobies <Emphasis Type="Italic">Nes longus</Emphasis> and <Emphasis Type="Italic">Ctenogobius saepepallens</Emphasis>, Both Symbiotic with the Snapping Shrimp <Emphasis Type="Italic">Alpheus floridanus</Emphasis>

Synopsis We re-examined the symbiotic association of the western Atlantic gobiid fishes Nes longus and Ctenogobius saepepallens with the snapping shrimp Alpheus floridanus on the basis of a critical literature review and new data. Our research confirms that N. longus interacts closely with the shrimp and is dependent on it for the cover provided by the burrow that the shrimp constructs; the goby serves as the sentinel at the burrow entrance. Ctenogobius saepepallens is often seen occupying a burrow of the alpheid, and the shrimp will leave the burrow to deposit sediment with the goby at the entrance, even pushing the goby aside at times. However, the shrimp does not make contact with the goby with its antennae, nor does the goby communicate with caudal fin fluttering at the approach of danger. We suggest that their relationship is a first step in an evolutionary process that may lead to the very close mutualistic association exhibited by N. longus and the alpheid, as well as Indo-Pacific shrimp gobies of 13 different genera and their alpheid partners. Nes longus remains close to the burrow entrance; it feeds mainly on small gastropods, decapod crustaceans, ostracods, and isopods. By contrast, C. saepepallens makes longer excusions from the shelter of the burrow; its diet is dominated by benthic copepods, followed by ostracods and lesser amounts of foraminiferans, isopods, and decapod crustaceans. By virture of its greater mobility, it can be more selective in its prey.     

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.     

Carnivorous and detritivorous feeding of Mesocyclops leuckarti Claus (Cyclopoida Copepoda)

Food preference of M. leuckarti was experimentally analysed in relation to different species of live and dead crustaceans. Also their possibility of using detritus from bottom sediments as a source of food was tested.It has been found that M. leuckarti readily feed on dead organisms, and detritus from bottom sediments is sufficient for their survival.On the basis of experimental results the feeding strategy of M. leuckarti in lakes is discussed. It seems that the availability of different species of live prey depends on the mode of their swimming and on their body cover, while the availability of dead prey depends only on the body cover. Detritus may be an important source of food, particularly when the live prey are scarce in the pelagic zone, or not easily available.     

The importance of gobies (Gobiidae,Teleostei) as hosts and transmitters of parasites in the SW Baltic

The parasite fauna of five goby species (Gobiidae, Teleostei) was investigated in the Baltic Sea during the period 1987 to 1990. 13 parasite species were found in samples from the Lübeck Bight:Bothriocephalus scorpii, Schistocephalus sp. (Cestoda);Cryptocotyle concavum, Cryptocotyle lingua, Podocotyle atomon, Derogenes varicus (Digenea);Hysterothylacium sp. (cf.auctum),Contracaecum sp.,Anisakis simplex (Nematoda);Corynosoma sp.,Echinorhynchus gadi, Neoechinorhynchus rutili, Pomphorhynchus laevis (Acanthocephala). The number of parasite species were: 10 in the sand gobyPomatoschistus minutus, 8 in the black gobyGobius niger, 7 in the two-spotted gobyGobiusculus flavescens, 6 in the common gobyPomatoschistus microps, and 5 in the painted gobyPomatoschistus pictus. Neoechinorhynchus rutili occurred only inP. minutus, andCorynosoma sp. only inG. niger. The extent to which the gobies were parasitized clearly depended on the respective ways of life and, moreover, on the kind of prey ingested by the hosts. Additionally, the age of the hosts might be important. The highest rate of parasitism, more than 60%, was reached byHysterothylacium sp. inG. niger and byCryptocotyle concavum inP. microps. Infestation incidence lay mostly below 40% which means a satellite species status (Holmes, 1991). The number of parasite species was highest in summer; the highest intensities of single parasites occurred in spring (Podocotyle atomon) or autumn (Crytocotyle concavum).Bothriocephalus scorpii, Hysterothylacium sp. andPodocotyle infested their juvenile hosts very early, but onlyHysterothylacium was accumulated byG. niger during its whole life span, whereasBothriocephalus persisted also in older gobies in low intensities. The cercariae ofCryptocotyle spp. penetrate actively into their hosts; all the other parasites named were transmitted in larval form by prey organisms which consisted mainly of planktonic and benthic crustaceans. The gobies were final hosts for only 5 parasites; but two species may be transmitted to larger fish, and 6 species to sea birds or mammals. The parasite community of the five gobies may possibly be taken to characterize the ecological quality of the environment of the Lübeck Bight.     

Invaders eating invaders: exploitation of novel alien prey by the alien shimofuri goby in the San Francisco Estuary,California

The shimofuri goby (Tridentiger bifasciatus), which is native to Asian estuaries, was recently introduced to the San Francisco Estuary, California, USA. We conducted gut content analyses to examine the gobys feeding ecology in this highly invaded estuary. Shimofuri gobies were generalist predators on benthic invertebrates, consuming seasonally abundant prey, especially amphipods (Corophium spp.). In addition, shimofuri goby utilized two novel prey items not exploited by other resident fishes – hydroids (Cordylophora caspia) and barnacle (Balanus improvisus) cirri, both of which are alien. The shimofuri gobys feeding ecology appears well-suited to the fluctuating environment of the San Francisco Estuary and may partially explain observed increases in shimofuri goby abundance compared with declines in populations of some native species.     

Short-term predator avoidance behavior by invasive and native amphipods in the Great Lakes

Understanding predator avoidance behavior by prey remains an important topic in community and invasion ecology. Recently, the Ponto-Caspian amphipod Echinogammarus ischnus (Stebbing 1898) was accidentally introduced into the Great Lakes. Since its introduction, it has displaced the native amphipod, Gammarus fasciatus (Say 1818), from several locations in the lower lakes. To assess whether behavioral differences in predator avoidance might be a causal mechanism increasing the success of the invasive amphipods, two experiments were conducted examining (1) native and invasive amphipod behavioral responses to five fish species with different foraging behaviors, and (2) amphipod responses to different densities of round gobies, a hyper-abundant benthic invertivore. Echinogammarus reduced its distance moved in the presence of all fish species tested, whereas Gammarus reduced its distance moved only after exposure to round gobies, black crappies, and rainbow darters. Both amphipod species increased the time spent motionless following exposure to round gobies, but not after encountering the scent of most of the remaining fish predators. The exception was that Echinogammarus also responded to black crappie scent whereas Gammarus did not. Although both amphipod species exhibited behavioral responses to many of the fish predators, the magnitude of their responses differed only after exposure to the brown bullhead. In the bullhead trials, Echinogammarus reduced its distance traveled significantly more than Gammarus. Both amphipod species increased their avoidance response to increasing goby density, however, the pattern of avoidance behavior was different. Invasive E. ischnus exhibited a consistently strong avoidance response to round gobies over the test duration. Native G. fasciatus initially avoided goby scent, but then either ceased their avoidance response or showed a hyper-avoidance response, depending on goby density. These results suggested (1) both species of amphipods were able to differentiate and react to a variety of fish predators, (2) invasive Echinogammarus amphipods avoided a larger range of fish predators than the native Gammarus, (3) increased avoidance behavior was associated with an increased density of fish, and (4) the avoidance response patterns of invasive Echinogammarus when faced with round goby predators might lead to increased predation on native Gammarus in habitats where they co-occur.     

The round goby (Neogobius melanostomus) in the Gulf of Gdańsk – a species introduction into the Baltic Sea

In recent years, information concerning the awareness of organisms accidentally introduced into the Baltic Sea has substantially improved. Non-indigenous Estuarine and Marine Organisms (NEMO's) are hazardous for the Baltic ecosystem. Currently, about one hundred species are identified as accidentally or intentionally introduced into the Baltic Sea. Ballast waters and escape from aquaculture are the most important invasion vectors. During the last decade, an invasion of the round goby (Neogobius melanostomus) – a Ponto-Caspian fish species has been observed in the Gulf of Gdańsk. The first record of this fish in the Baltic Sea is from 1990. Early detection of the invader enabled the study of population growth and changes in the area. The first years of invasion were characterized by low numbers of individuals and a limited distribution. Later, the round goby gradually colonized all shallow waters in the western part of the Gulf of Gdańsk. Initially the fish inhabited stony and rocky habitats, but later it also occupied sandy bottoms. The round goby is now the dominating fish species in most of the shallow waters of the Gulf of Gdańsk. Two main factors account for the successful invasion of this fish in the region: the state of ecosystem at the time of the invasion and the biological features of N. melanostomus. In the late 1980s, the shallow waters of the Gulf of Gdańsk were almost devoid of piscivorous fishes. Concurrently, bivalves (a preferred prey of the round goby) have increased. Important is also parental care of laid eggs and reproductive strategy. Population growth potential enables the colonization of nearby regions. The first round gobies in the Vistula Lagoon were collected in 1999 and colonization of other Baltic Sea areas is anticipated.     

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.     

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.     

Predator and prey: the role of the round goby Neogobius melanostomus in the western Baltic

Different studies on the position of the non-indigenous species Neogobius melanostomus within the coastal food web of the Pomeranian Bay (western Baltic) were performed, resulting in a quantitative and qualitative species list of prey organisms found in the stomachs of the invader and an estimation concerning the importance of round goby as prey for different resident predators. It seems that the colonization process is not fully completed yet, but the results reveal that the species is already established in the food web 16 years after the first observation within the study area. The results show that N. melanostomus feed upon a wide range of different resident organisms. While a direct predation effect on native fish species appears rather unlikely, indirect effects such as competition cannot yet be excluded. In addition, our results reveal an ontogenetic diet shift and that the round goby itself already serves as an important prey for piscivorous fish and seabirds. Finally, we formulate different hypotheses based on our results which will require further research.