Evidence for predatory control of the invasive round goby

We coupled bioenergetics modeling with bottom trawl survey results to evaluate the capacity of piscivorous fish in eastern Lake Erie to exert predatory control of the invading population of round goby Neogobius melanostomus. In the offshore (>20 m deep) waters of eastern Lake Erie, burbot Lota lota is a native top predator, feeding on a suite of prey fishes. The round goby invaded eastern Lake Erie during the late 1990s, and round goby population size increased dramatically during 1999–2004. According to annual bottom trawl survey results, round goby abundance in offshore waters peaked in 2004, but then declined during 2004–2008. Coincidentally, round goby became an important component of burbot diet beginning in 2003. Using bottom trawling and gill netting, we estimated adult burbot abundance and age structure in eastern Lake Erie during 2007. Diet composition and energy density of eastern Lake Erie burbot were also determined during 2007. This information, along with estimates of burbot growth, burbot mortality, burbot water temperature regime, and energy densities of prey fish from the literature, were incorporated into a bioenergetics model application to estimate annual consumption of round goby by the adult burbot population. Results indicated that the adult burbot population in eastern Lake Erie annually consumed 1,361 metric tons of round goby. Based on the results of bottom trawling, we estimated the biomass of yearling and older round goby in offshore waters eastern Lake Erie during 2007–2008 to be 2,232 metric tons. Thus, the adult burbot population was feeding on round goby at an annual rate equal to 61% of the estimated round goby standing stock. We concluded that the burbot population had high potential to exert predatory control on round goby in offshore waters of eastern Lake Erie.     

Ten polymorphic microsatellite markers in the invasive round goby (Neogobius melanostomus) and cross‐species amplification

Ten tetranucleotide, dinucleotide and compound microsatellite loci were isolated and characterized for the round goby, Neogobius melanostomus. Analysis of 64 gobies from one nonindigenous population in Lake Erie, Ontario, Canada, indicated that allele number varied from three to 12 per locus, while observed heterozygosity ranged between 0.33 and 0.86. Eight of these primers showed some amplification in other species in four genera. These newly developed microsatellite markers are a powerful tool that will provide insights into population structure and dispersal of the round goby in their novel environment.     

Expansion of tubenose gobies <Emphasis Type="Italic">Proterorhinus semilunaris</Emphasis> into western Lake Erie and potential effects on native species

The Eurasian freshwater tubenose goby Proterorhinus semilunaris (formerly Proterorhinus marmoratus) invaded the Laurentian Great Lakes in the 1990s, presumably via ballast water from transoceanic cargo ships. Tubenose gobies spread throughout Lake St. Clair, its tributaries, and the Detroit River system, and also are present in the Duluth-Superior harbor of Lake Superior. Using seines and bottom trawls, we collected 113 tubenose gobies between July 2007 and August 2009 at several locations in western Lake Erie. The number and range of sizes of specimens collected suggest that that tubenose gobies have become established and self-sustaining in the western basin of Lake Erie. Tubenose gobies reached maximum densities in sheltered areas with abundant macrophyte growth, which also is their common habitat in native northern Black Sea populations. The diet of tubenose gobies was almost exclusively invertebrates, suggesting dietary overlap with other benthic fishes, such as darters (Etheostoma spp. and Percina sp.), madtoms (Noturus spp.), and sculpins (Cottus spp.). A single mitochondrial DNA haplotype was identified, which is the most common haplotype found in the original colonization area in the Lake St. Clair region, suggesting a founder effect. Tubenose gobies, like round gobies Neogobius melanostomus, have early life stages that drift owing to vertical migration, which probably allowed them to spread from areas of colonization. The Lake St. Clair-Lake Erie corridor appears to have served as an avenue for them to spread to the western basin of Lake Erie, and abundance of shallow macrophyte-rich habitats may be a key factor facilitating their further expansion within Lake Erie and the remainder of the Laurentian Great Lakes.     

Swimming performance and invasion potential of the round goby

European round gobies (Neogobius melanostomus) are displacing several important native North American fish species. Controlling their invasion is contingent on understanding their swimming inclination and potential. We assessed goby swimming inclination by recording activity in a 2 m flume over a ~24 h period, and swimming potential using a critical swimming (U _crit) test, as well as burst tests in still and flowing water. When given the choice to move, gobies covered as much as 14 m/h, with a slight bias towards nocturnal activity and an overall upstream preference. When confined and coerced to perform a U _crit test, they burst-and-held to achieve 35.5 ± 1.1 cm/s. Thirty minutes following U _crit, they were able to burst-and-coast in a sprint test to almost twice this speed. In still water, they exhibited startle bursts of up to 163 cm/s. We provide a swimming endurance model that indicates flow rates would need to be >125 cm/s to prevent upstream movement, and free of refuge areas in which to recover. The current study shows that the round goby is a surprisingly powerful swimmer with the capacity to continue its invasion should hydrologic control be absent.     

Seasonal habitat use indicates that depth may mediate the potential for invasive round goby impacts in inland lakes

1. The round goby (Neogobius melanostomus) is among the fastest-spreading introduced aquatic species in North America and is radiating inland from the Great Lakes into freshwater ecosystems across the landscape. Predicting and managing the impacts of round gobies requires information on the factors influencing their distribution in habitats along the invasion front, yet this information is not available for many recently invaded ecosystems. We evaluated the seasonal habitat use and biomass of round gobies in an inland temperate lake to define the spatiotemporal scope of biological interactions at the leading edge of the round goby invasion.
2. Using novel statistical approaches, we combined hierarchical models that control for imperfect species detection with flexible smooth terms to describe non-linear relationships between round goby abundance and environmental gradients. Subsequently, we generated accurate detection-corrected estimates of the standing stock biomass of round gobies.
3. Our results show seasonally differentiated habitat niches, where suitable round goby habitat in summer months is restricted to shallow depths (<18.4 m) with a mixture of vegetative and mussel cover. We found high round goby biomass of 122 kg/ha in occupied habitats during the summer, with a total lake-wide biomass of 766,000 kg. In winter, round gobies migrate to deep offshore habitats and disperse, dramatically altering their scope for biological interactions with resident aquatic species across summer and winter seasons.
4. The results of this study indicate that the scope of biological interactions in inland lakes may be seasonally variable, with potential for high round goby biomass in shallow lakes or at the periphery of deep lakes in the summer months. Such shallow-water habitats may therefore present higher risk of ecological impacts from round gobies in invaded lentic ecosystems. As round gobies expand inland, consideration of seasonal habitat use will be an important factor in predicting the impacts of this pervasive invader.

     

Increased parental care cost for nest-guarding fish in a lake with hyperabundant nest predators

Although parental care increases offspring survival, providingcare is costly, reducing parental growth and survival and, thereby,compromising future reproductive success. To determine if anexotic benthic predator might be affecting parental care bynest-guarding smallmouth bass (Micropterus dolomieu), we comparednest-guarding behavior and energy expenditures in two systems,one with a hyperabundant recently introduced predator, the roundgoby (Neogobious melanostomus). In Lake Erie, USA, smallmouthbass vigorously defended their nests from benthic round gobies.In Lake Opeongo, Canada, smallmouth bass were exposed to fewerand predominantly open-water predators and were less activein their nest defense. From scuba and video observations, wedocumented that nest-guarding smallmouth bass chased predators(99% of which were round gobies) nine times more frequentlyin Lake Erie than in Lake Opeongo. This heightened activityresulted in a significant decline in weight and energetic contentof guarding males in Lake Erie but no change in Lake Opeongomales. Bioenergetic simulations revealed that parental careincreased smallmouth bass standard metabolic rate by 210% inLake Erie but only by 28% in Lake Opeongo. As energy reservesdeclined and offspring became increasingly independent, malesin both lakes consumed more prey and spent more time foragingaway from their nests; however, nest-guarding smallmouth bassconsumed few prey and, in Lake Erie, rarely consumed round gobies.Therefore, increased parental care costs owing to the presenceof round gobies could affect future growth, reproduction, andsurvival if smallmouth bass approach critically low energy reserves.     

Secondary invasion of the round goby into high diversity Great Lakes tributaries and species at risk hotspots: potential new concerns for endangered freshwater species

The round goby (Neogobius melanostomus) first invaded North America in 1990 when it was discovered in the St. Clair River. Despite more than 15 years of potential invasion, many Great Lakes’ lotic systems remained uninvaded. Recently, we captured the round goby from several Great Lakes tributaries known as species-at-risk hotspots. With a combination of field sampling of round gobies and literature review of the impact of round gobies on native taxa, we assess the potential impacts of the secondary invasion to native species using three mechanisms: competition; predation; and indirect impacts from the loss of obligate mussel hosts. We estimate that 89% (17/19) of benthic fishes and 17% (6/36) of mussels that occur in these systems are either known or suspected to be impacted by the secondary invasion of round goby. In particular, we note that the distribution of potential impacts of round goby invasion was largely associated with species with a conservation designation, including seven endangered species (1 fish, 6 mussels). As these recent captures of round goby represent novel occurrences in high diversity watersheds, understanding the potential impacts of secondary invasion to native biota is fundamental to prevent species declines and to allow early mitigation.     

Population genetic structure of the round goby in Lake Michigan: implications for dispersal of invasive species

Understanding subsequent dispersal of non-native species following introduction is important for predicting the extent and speed of range expansion and is critical for effective management and risk assessment. Post-introduction dispersal may occur naturally or via human transport, but assessing the relative contribution of each is difficult for many organisms. Here, we use data from seven microsatellite markers to study patterns of dispersal and gene flow among 12 pierhead populations of the round goby (Neogobius melanostomus) in Lake Michigan. We find significant population structure among sampling sites within this single Great Lake: (1) numerous populations exhibited significant pairwise F _ST and (2) a Bayesian assignment analysis revealed three distinct genetic clusters, corresponding to different pierhead locations, and genetic admixture between these clusters in the remaining populations. Genetic differentiation (F _ST) is generally related to geographic distance (i.e., isolation by distance), but is periodically interrupted at the scale of Lake Michigan due to gene flow among geographically distant sites. Moreover, average genetic differentiation among populations exhibit a significant, negative correlation with the amount of shipping cargo at ports. Our results, therefore, provide evidence that genetic structure of the round goby in Lake Michigan results from limited natural dispersal with frequent long-distance dispersal through anthropogenic activities such as commercial shipping. Our study suggests that while round gobies can undoubtedly disperse and found new populations through natural dispersal mechanisms, their spread within and among the Great Lakes is likely aided by transport via ships. We, therefore, recommend that ballast-water treatment and management may limit the spread of non-native species within the Great Lakes after the initial introduction in addition to preventing the introduction of non-native species to the Great Lakes.     

The Round Goby, Neogobius melanostomus, a Fish Invader on both sides of the Atlantic Ocean

During the past decade, a bottom-dwelling, aggressive, multiple-spawning fish, the round goby (Gobiidae: Neogobius melanostomus), has spread from its native region in the Ponto-Caspian throughout Europe and to the Laurentian Great Lakes in North America. An international workshop, held at the Hel Marine Station, Poland, was organized to summarize population features of the round goby. Common fish predators of round gobies in the Great Lakes and in native regions are obligate and facultative benthic fishes and occasionally, pelagic fishes. In contrast, the main predator of the round goby in the Gulf of Gdansk is the Great Cormorant (Phalacrocorax carbo). In the Great Lakes, round gobies have lead to the decline of mottled sculpin (Cottus bairdi) and logperch (Percina caprodes) and reduced the hatching success of native fishes by feeding on their eggs. In the Gulf of Gdansk, round gobies have increased in abundance, while three-spined sticklebacks (Gasterosteus aculeatus) have declined. Round gobies have a broad diet throughout their range; larger specimens are molluscivores. There are fewer species of parasites and lower infection rates of round gobies in recently colonized areas than in native areas. Overall, newly colonized round gobies in brackish waters and lakes are smaller, mature earlier, have a male biased operational sex ratio and are more short-lived compared with round gobies from marine (native) habitats.     

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.     

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.     

Correlation between body size and fatty acid and essential amino acid composition of round goby (Neogobius melanostomus) and monkey goby (Neogobius fluviatilis) from the Rhine River (Germany)

In this study correlations between body size and muscle fatty and amino acid content of two species of goby, round goby (Neogobius melanostomus) and monkey goby (Neogobius fluviatilis) caught from river Rhine (Germany) were investigated. Among saturated fatty acids (SFAs), mono- (MUFA) and polyunsaturated fatty acids (PUFAs) only SFAs were significantly higher in round goby than monkey goby (P < 0.05). In general, the correlation between body size of both gobies and the content of most of the individual fatty acids was not significant. In monkey goby, the content of palmitic acid (C16:0) and oleic acid (C18:1 n-9) was positively correlated with weight (r = 0.43) and total length (r = ?0.58), respectively, and the content of docosahexaenoic acid (DHA) increased with condition factor (r = 0.50). The content of threonine, arginine, valine, phenylalanine and isoleucine in monkey goby was higher than those of round goby (P < 0.05). In round goby the three essential amino acids arginine, valine and leucine were positively (P < 0.05) correlated with body length, which indicates that longer round gobies are of higher nutritional value.     

The occurrence of non-native tubenose goby Proterorhinus semilunaris in the pelagic 0+ year fish assemblage of a central European reservoir

In July 2008, early juvenile tubenose goby Proterorhinus semilunaris were found in nocturnal pelagic waters of the Vranov Reservoir, Czech Republic. Presence of benthic-living prey in the guts of these fish suggested migration between benthic and pelagic habitats.     

Age,growth and reproductive traits of invasive goby Taenioides cirratus in the Chaohu Lake,China

The eel goby Taenioides cirratus (Blyth, 1,860) is a small fish inhabits muddy bottoms of brackish-water in the Indo-West Pacific. It has invaded many inland freshwater lakes in China, such as the Chaohu Lake, Gaoyou Lake and Nansi Lake, and its population increased rapidly in these freshwater lakes in recent years. The age, growth and reproductive traits of T. cirratus invading the Chaohu Lake were studied. A total of 482 specimens (210 females, 204 males and 68 juveniles) with total length (TL) ranging from 9.4 to 20.6 cm were collected using the benthic fyke nets at monthly intervals from March 2018 to February 2019. The sagittal otolith was used for age determination. Monthly variation of marginal increment ratio indicated that the annual forming of opaque band on sagittal otolith was completed during March and April. For both sexes, only four (from 0+ to 3+ years) age groups were observed and 1+ and 2+ years age individuals dominated the population. Back calculated length at age showed males grew faster than females. Both sexes reached maturity at 1+ year age and the TL at first maturity (TL₅₀) was 12.6 cm for females and 11.9 cm for males. Monthly variation of gonado-somatic index indicated that the spawning occurred from May to August. The fecundity ranged from 967 ova to 5,114 ova, with a mean of 3,205 ova. Our study provides a comprehensive data on the key life history traits of T. cirratus for the first time.     

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.     

Krill-feeding behaviour of gentoo penguins as shown by animal-borne camera loggers

Animal-borne camera loggers were used to examine the patterns of prey encounter and feeding behaviour of gentoo penguins at King George Island, Antarctica. The still images from the camera loggers showed that the penguins encountered the swarms of krill for 25.5% (range: 8–38%) of their dives (>5 m) on average, during their foraging trips (mean duration of 5.4 h, n = 7 trips). They encountered krill swarms during the dives to 10–70 m depth, in pelagic as well as benthic habitats. In the benthic habitat, the penguins swam just above the sea floor and headed downward over a krill swarm, probably using the sea floor to assist them to feed on mobile swarms. The shallow coastal waters would be the important foraging habitat of gentoo penguins breeding in King George Island.     

Predation of zebra mussels by round gobies, Neogobius melanostomus

Preliminary gut analysis of a recent Great Lakes invader, the round goby, Neogobius melanostomus (7.0–8.4 cm), collected from the Detroit River, showed that they ate zebra mussels (58%), snails (6%), and other invertebrates (36%), including aquatic insects (Hexagenia), softshelled crayfish, and zooplankton. Because zebra mussels, Dreissena polymorpha, predominated as prey, we investigated the ability of round gobies to consume different size classes of zebra mussels. In laboratory experiments, we examined feeding preferences of three size classes of round gobies (5.5–6.9 cm; 7.0–8.4 cm; 8.5–10.3 cm standard length) on four different size classes of zebra mussels (6.0–9.9 mm, 10.0–12.9 mm, 13.0–15.9 mm, 16.0–18.9 mm). All sizes of round gobies ate zebra mussels < 10.0 mm. Only the largest size class of round gobies ate larger zebra mussels (10.0–12.9 mm) when all prey sizes were presented. The association between the total mass of zebra mussels available and the amount consumed by round gobies increased positively up to about 6.5 g of available mussels and then levelled off. Round gobies consumed an average of 1.0 g of mussels in 24 h. There was a significant positive relationship between gape size and standard length of round gobies. Although larger round gobies (over the size range of fish in our study) are able to consume larger zebra mussels, small mussels were preferred. Our findings suggest that the preference of small zebra mussels by round gobies has the potential to alter the size structure of zebra mussel populations. This revised version was published online in July 2006 with corrections to the Cover Date.     

Predictors of the distribution and abundance of a tube sponge and its resident goby

Microhabitat specialists offer tractable systems for studying the role of habitat in determining species’ distribution and abundance patterns. While factors underlying the distribution patterns of these specialists have been studied for decades, few papers have considered factors influencing both the microhabitat and the inhabitant. On the Belizean barrier reef, the obligate sponge-dwelling goby Elacatinus lori inhabits the yellow tube sponge Aplysina fistularis. We used field data and multivariate analyses to simultaneously consider factors influencing sponge and goby distributions. Sponges were non-randomly distributed across the reef with density peaking at a depth of 10–20 m. Sponge morphology also varied with depth: sponges tended to be larger and have fewer tubes with increasing depth. Knowing these patterns of sponge distribution and morphology, we considered how they influenced the distribution of two categories of gobies: residents (≥18 mm SL) and settlers (<18 mm SL). Maximum tube length, number of sponge tubes, and depth were significant predictors of resident distribution. Residents were most abundant in large sponges with multiple tubes, and were virtually absent from sponges shallower than 10 m. Similarly, maximum tube length and number of sponge tubes were significant predictors of settler distribution, with settlers most abundant in large sponges with multiple tubes. The presence or absence of residents in a sponge was not a significant predictor of settler distribution. These results provide us with a clear understanding of where sponges and gobies are found on the reef and support the hypothesis that microhabitat characteristics are good predictors of fish abundance for species that are tightly linked to microhabitat.     

Dispersal strategies, secondary range expansion and invasion genetics of the nonindigenous round goby, Neogobius melanostomus, in Great Lakes tributaries

Dispersal strategies are important mechanisms underlying the spatial distribution and colonizing ability of all mobile species. In the current study, we use highly polymorphic microsatellite markers to evaluate local dispersal and colonization dynamics of the round goby (Neogobius melanostomus), an aquatic invader expanding its range from lake to river environments in its introduced North American range. Genetic structure, genotype assignment and genetic diversity were compared among 1262 round gobies from 20 river and four lake sites in three Great Lakes tributaries. Our results indicate that a combination of short-distance diffusion and long-distance dispersal, collectively referred to as 'stratified dispersal', is facilitating river colonization. Colonization proceeded upstream yearly (approximately 500 m/year; 2005-2009) in one of two temporal replicates while genetic structure was temporally stable. Contiguous dispersal from the lake was observed in all three rivers with a substantial portion of river fish (7.3%) identified as migrants. Genotype assignment indicated a separate introduction occurred upstream of the invasion front in one river. Genetic diversity was similar and relatively high among lake and recently colonized river populations, indicating that founder effects are mitigated through a dual-dispersal strategy. The remarkable success of round goby as an aquatic invader stresses the need for better diffusion models of secondary range expansion for presumably sessile invasive species.     

Range expansion dynamics of the invasive round goby (Neogobius melanostomus) in a river system

The rates of upstream and downstream range expansion of the round goby (Neogobius melanostomus) were examined in the Trent-Severn Waterway in 2009 and 2010. Relative abundance, demographics, and habitat use were also compared between areas of range expansion and a longer established area to determine how these characteristics varied along the invasion pathway. Round gobies were sampled using an angling removal method in May and August of 2009 and 2010 at 75 sites at each area of range expansion and 25 sites near the center of their range where they first became established in the waterway. Areas of range expansion had initially low abundance and low site occupancy in May 2009 relative to the longer established area. Large increases in abundance and site occupancy were observed over the first summer of occupation, but with limited range expansion. Rapid range expansion was observed during the non-reproductive season at the upstream edge of range. Individuals sampled in the expanded upstream range were small and female biased relative to other range locations. Round gobies also exhibited higher habitat selectivity for rocky substrates at range edges than in the longer established area. These characteristics of seasonal dispersal, biased demographics of dispersers, and high habitat selectivity may be important components of successful range expansion of the round goby in invaded ecosystems.