Fish Assemblages and Environmental Variables Associated with Gobiids in Nearshore Areas of the Lower Great Lakes

We investigated which fish species and environmental variables were associated with the invasive round goby (Neogobius melanostomus) and tubenose goby (Proterorhinus marmoratus) in nearshore Canadian waters of the Huron-Erie corridor of the lower Great Lakes. We measured a suite of environmental variables and used triplicate beach seine samples to collect fishes in summer 2006. Thirty sites were sampled in the day and a subset (n = 14) at night. Of 1,955 individuals caught in daytime samples, round goby (21.0 %), spottail shiner (17.3%) and emerald shiner (14.2%) were most abundant. Of 1,521 individuals collected at night, the most abundant species were round goby (42.3%) and emerald shiner (24.1%). Tubenose gobies represented 1% and 1.7% of all individuals caught in the day and night, respectively. Rarefaction analysis showed that overall species richness was greater in the day than night. Significantly more emerald shiner (P = 0.017), rock bass (P = 0.046) and round goby (P = 0.035) were caught at night than in the day; more logperch were caught in the day than at night (P = 0.042). Round gobies were positively associated with water temperatures up to 24°, but there was no relationship between round goby abundance and warmer temperatures. There were too few tubenose goby captured to determine their statistical association with environmental factors; however, tubenose gobies were found only where round gobies were collected. Round goby and tubenose goby were associated with yellow perch and rock bass. The benthic round goby was the most abundant species, whereas other abundant species were pelagic, schooling fishes that occupied a habitat distinct from round goby.     

Impacts of predation by the Eurasian round goby (Neogobius melanostomus) on molluscs in the upper St. Lawrence River

An invasive Eurasian fish, the round goby Neogobius melanostomus, has recently spread from the Great Lakes into the St. Lawrence River. We quantified prey preferences of this benthivore and determined whether its predatory impacts on molluscs in the river are similar to those in the Great Lakes. We measured the size structure of gastropods and dreissenid mussels at 13 St. Lawrence River sites where round goby densities ranged from 0 to 6 m^− 2. For four of these sites, data were available for multiple years before and after invasion. Contrary to studies in the Great Lakes, there were no consistent effects of round goby density on the size structure of dreissenids, although there was an ontogenetic diet shift toward dreissenids. However, the abundance and richness of small gastropods (≤ 14 mm) was negatively correlated with round goby density across all sites, and declined over time at three of four sites sampled before and after invasion. Median gastropod size also declined across sites with increasing round goby density. Gastropods (as well as chironomid larvae, caddisfly larvae, and ostracods) were consistently among the most preferred prey items consumed by gobies, whereas dreissenids (as well as leeches and freshwater mites) were consistently avoided. These results indicate the major role of the round goby in structuring gastropod populations in the St. Lawrence River, and highlight large-scale spatial variation in its predatory impact on dreissenid populations.     

Effectiveness of Piscicides for Controlling Round Gobies (Neogobius melanostomus)

Round gobies (Neogobius melanostomus) were introduced to the Great Lakes presumably as a result of ballast water releases from seagoing freighters returning from European water bodies. These unwelcome fish have become established in the Great Lakes region and are expanding their range to suitable portions of other interior drainage basins including the Mississippi River traversing the central United States and the Trent-Severn waterway spanning south-central Ontario. If the invasion continues, use of chemical toxicants as a control measure may be necessary. Toxicity tests of the currently registered piscicides antimycin, rotenone, 3-trifluoromethyl-4-nitrophenol (TFM), and Bayluscide^® were conducted with three fish species native to the Great Lakes and round gobies collected from the Illinois Waterway. Tests indicated that round gobies are sensitive to all of the piscicides, however, the level of sensitivity is similar to that of the native fish species tested. Therefore, currently registered piscicides have limited potential to selectively remove round gobies. Bottom-release formulations of Bayluscide^® and antimycin were also evaluated as control agents for the normally bottom-dwelling round goby. Avoidance behavior tests demonstrated that the round goby did not react to the presence of either chemical. Therefore, the bottom-release formulations may have some application for the selective removal of round gobies, and may be one of the few tools presently available to fishery managers to help limit the range expansion of this invasive fish.     

Comparison of thermal tolerance and standard metabolic rate of two Great Lakes invasive fish species

Round goby (Neogobius melanostomus) and western tubenose goby (Proterorhinus semilunaris) invaded the Laurentian Great Lakes at approximately the same time and area yet have shown substantial differences in their post-invasion success with more rapid establishment and development of much larger abundances of round goby populations throughout the invaded habitat. In this study, we compared differences in physiological performance (thermal tolerance and standard metabolic rate) between round and tubenose goby collected from the Huron-Erie corridor. Tubenose goby were observed to have lower thermal tolerance but exhibited similar standard metabolic rate across environmental temperatures compared to round goby. At temperatures exceeding 31 °C, tubenose goby demonstrated significantly higher mortalities and shorter times to death relative to round goby. The observed differences in thermal tolerance were consistent with differences in the native geographic ranges observed for each species at their southern ranges. The observed differences in physiological performance combined with species differences in other life history traits such body size, reproduction, feeding ecology and habitat affiliation may also explain differences in the invasiveness experienced by these two Great Lakes invasive fish including a greater ability of round gobies to occupy extreme habitats with large water temperature fluctuations.     

Occurrence and Food Habits of the Round Goby in the Profundal Zone of Southwestern Lake Ontario

Little is known about the ecology of round goby (Neogobius melanostomus), an invasive benthic fish, in the profundal zone of the Great Lakes. In April 2002–2005 we caught increasing numbers of round gobies with a bottom trawl in the 45–150 m depth range of southwestern Lake Ontario. In 2005, we examined gut contents of 30 round gobies from each of three depths, 55, 95, and 130 m, and qualitatively compared gut contents with density of benthic invertebrates determined by Ponar grabs. Round goby guts contained mostly Dreissena spp. and opposum shrimp, Mysis relicta (Mysis); the frequency of occurrence of dreissenids in guts decreased with depth, whereas the frequency of occurrence of Mysis in guts increased with depth. Abundance of these invertebrates in the environment followed the same pattern, although dreissenids of optimum edible size (3–12 mm) were still abundant (1,373/m²) at 130 m, where round gobies primarily consumed Mysis, suggesting that round gobies may switch from dreissenids to more profitable prey when it is available. Other food items were ostracods and fish, with ostracods generally eaten by smaller round gobies and fish eaten by larger round gobies. Occurrence and increasing abundance of round gobies in the profundal zone and predation on Mysis by round goby could have far-reaching consequences for the Lake Ontario fish community.     

Salinity tolerance of the invasive round goby: Experimental implications for seawater ballast exchange and spread to North American estuaries

The Eurasian round goby (Neogobius melanostomus) invaded the freshwater North American Great Lakes in ~ 1990 via accidental introduction from ballast water discharge. Its genotypes in the Great Lakes traced to estuaries in the northern Black Sea, where the round goby flourishes in a variety of salinities to 22 parts per thousand (ppt). To prevent further introductions, U.S. and Canadian Coast Guard regulations now require that vessels exchange ballast water at sea before entering the Great Lakes. Since salinity tolerance of the invasive round goby population is poorly understood, we tested 230 laboratory-acclimated fish in three experimental scenarios: (1) rapid salinity increases (0–40 ppt), simulating ballast water exchange, (2) step-wise salinity increases, as during estuarine tidal fluxes or migration from fresh to saltwater, and (3) long-term survivorship and growth (to 4 months) at acclimated salinities. Almost all gobies survived experiments at 0–20 ppt, whereas none survived ≥ 30 ppt, and at 25 ppt only 15% withstood rapid changes and 30% survived step-wise increases. Ventilation frequencies were lowest at 10–15 ppt in step-wise experiments, in conditions that were near isotonic with fish internal plasma concentrations, reflecting lower energy expenditure for osmoregulation. Growth rates appeared greatest at 5–10 ppt, congruent with the larger sizes reached by gobies in Eurasian brackish waters. Thus, we predict that the Great Lakes round goby would thrive in brackish water estuaries along North American coasts, if introduced. However, oceanic salinities appear fatal to the invasive round goby, which likely cannot withstand complete seawater ballast exchanges or oceanic habitats.     

Slimy sculpin depth shifts and habitat squeeze following the round goby invasion in the Laurentian Great Lakes

The collapse of Diporeia spp. and invasions of dreissenid mussels (zebra, Dreissena polymorpha; quagga, D. bugensis) and round goby (Neogobius melanostomus) have been associated with declines in abundance of native benthic fishes in the Great Lakes, including historically abundant slimy sculpin (Cottus cognatus). We hypothesized that as round goby colonized deeper habitat, slimy sculpin avoided habitat competition, predation, and aggression from round goby by shifting to deeper habitat. Accordingly, we predicted increased depth overlap of slimy sculpin with both round goby and deepwater sculpin (Myoxocephalus thompsonii) that resulted in habitat squeeze by both species. We used long-term bottom trawl data from Lakes Michigan, Huron, and Ontario to evaluate shifts in slimy sculpin depth and their depth overlap with round goby and deepwater sculpin. Lake Huron most supported our hypotheses as slimy sculpin shifted to deeper habitat coincident with the round goby invasion, and depth overlap between slimy sculpin and both species recently increased. Slimy sculpin depth trends in Lakes Michigan and Ontario suggest other ecological and environmental factors better predicted sculpin depth in these lakes.     

Metazoan Parasites of Introduced Round and Tubenose Gobies in the Great Lakes: Support for the “Enemy Release Hypothesis”

Recent invasion theory has hypothesized that newly established exotic species may initially be free of their native parasites, augmenting their population success. Others have hypothesized that invaders may introduce exotic parasites to native species and/or may become hosts to native parasites in their new habitats. Our study analyzed the parasites of two exotic Eurasian gobies that were detected in the Great Lakes in 1990: the round goby Apollonia melanostoma and the tubenose goby Proterorhinus semilunaris. We compared our results from the central region of their introduced ranges in Lakes Huron, St. Clair, and Erie with other studies in the Great Lakes over the past decade, as well as Eurasian native and nonindigenous habitats. Results showed that goby-specific metazoan parasites were absent in the Great Lakes, and all but one species were represented only as larvae, suggesting that adult parasites presently are poorly-adapted to the new gobies as hosts. Seven parasitic species are known to infest the tubenose goby in the Great Lakes, including our new finding of the acanthocephalan Southwellina hispida, and all are rare. We provide the first findings of four parasite species in the round goby and clarified two others, totaling 22 in the Great Lakes—with most being rare. In contrast, 72 round goby parasites occur in the Black Sea region. Trematodes are the most common parasitic group of the round goby in the Great Lakes, as in their native Black Sea range and Baltic Sea introduction. Holarctic trematode Diplostomum spathaceum larvae, which are one of two widely distributed species shared with Eurasia, were found in round goby eyes from all Great Lakes localities except Lake Huron proper. Our study and others reveal no overall increases in parasitism of the invasive gobies over the past decade after their establishment in the Great Lakes. In conclusion, the parasite “load” on the invasive gobies appears relatively low in comparison with their native habitats, lending support to the “enemy release hypothesis.”     

Fish and Macroinvertebrate Communities in Tributary Streams of Eastern Lake Erie with and without Round Gobies (Neogobius melanostomus, Pallas 1814)

Round gobies have had significant impacts on benthic fish and invertebrate communities in nearshore habitats of the Great Lakes. As round gobies have become more abundant in lake habitats, there has been an expansion of their populations into tributary streams and rivers. We compared stream invertebrate and fish communities in New York tributaries to Lake Erie with round gobies present and absent. Four of six benthic invertebrate metrics differed between streams with and without round gobies. Streams with round gobies present had reduced Shannon diversity, EPT richness, and EPT/chironomid ratios, and increased macroinvertebrate density relative to streams without round gobies, but there was no difference in non-Diptera density, or total taxa richness. None of the four fish metrics examined differed between streams with and without round gobies. However, darters occurred in all streams lacking round gobies, but did not occur in any streams with round gobies. Comparisons with historical fish and macroinvertebrate distributional data support our suspicion of goby-induced community changes. In these New York streams, round gobies seem to have had significant impacts on invertebrate communities via their consumptive behavior, whereas the impacts on fish communities are less evident. If round gobies continue to expand their distribution inland, the resultant alterations in macroinvertebrate communities may impact the suitability of tributary streams as spawning and nursery habitat for several sport fish species and for energy dynamics in tributary streams.     

Phenology and species diversity in a Lake Huron ichthyoplankton community: Ecological implications of invasive species dominance

Ichthyoplankton communities are dynamic and vary spatiotemporally based on factors such as wind, water currents, and phenology. Nonetheless, ichthyoplankton are an indicator of spawning success in fish populations and examining their community diversity and composition can serve to provide information on ecosystem integrity. Although some ichthyoplankton species may be transient, understanding their distribution in space and time provides information on species composition, abundance, and habitat use during critical early life stages. We sampled the spring-summer ichthyoplankton community during 2008 and 2009 in northern Lake Huron to determine species succession, abundance, and species diversity along physical and environmental gradients. Seasonal succession of species was similar during both years, indicating well-defined patterns in spawning by northern Lake Huron fish populations. Invasive alewife, rainbow smelt, and round goby were the dominant species during both years, with native stickleback species also abundant. Shannon Entropy (H′) increased with increasing water temperature until late summer when H′ declined. H′ decreased with increasing bottom depth and distance to tributary mouth indicating the important ecological role of these habitat features during early life stages. Although ichthyoplankton diversity was comparable to or higher than that reported for other areas of the Great Lakes, the prominence of invasive species in our study is reflective of the disturbed state of the Lake Huron fish community, despite large reductions in invasive planktivorous fish since 2004. Continued monitoring of ichthyoplankton communities will be important for measuring the impacts of species invasions or other ecosystem stressors on fish community structure in the Great Lakes.     

Diet shift of double-crested cormorants in eastern Lake Ontario associated with the expansion of the invasive round goby

The proliferation of the invasive round goby (Apollonia melanostoma) in the Great Lakes has caused shifts in the trophic ecology in some areas. We examined the diet of double-crested cormorants (Phalacrocorax auritas) prior to, and immediately after, round goby population expansion at two colonies, Pigeon and Snake Islands, in eastern Lake Ontario from 1999 to 2007. Cormorant diet was determined from the examination of 10,167 pellets collected over the nine-year period. By the second year round gobies were found in the diet (2002 at Snake Island and 2003 at Pigeon Island) they were the main species consumed by cormorants at each colony. The dominance of round goby in cormorant diets had a significant effect on both daily fish consumption and seasonal trends in fish consumption compared to the pre-goby years. Seasonal differences that were observed during the pre-goby years were lost once gobies became the main diet component of cormorants. The rapid switch to a benthic prey such as round goby, from a largely limnetic fish diet demonstrates the adaptive foraging ability of cormorants. Round goby may act as a buffer for yellow perch and smallmouth bass, two sport fish impacted by cormorant predation in eastern Lake Ontario.     

Response to putative round goby (Neogobius melanostomus) pheromones by centrarchid and percid fish species in the Laurentian Great Lakes

Pheromone trapping is an increasingly viable strategy to reduce invasive fish populations, largely due to the pheromones' function of evoking behavioral responses among conspecifics. Prior to attempting such population control techniques, the pheromones must be identified and their possible influences on non-target species addressed. The round goby (Neogobius melanostomus) is a species invasive to the Great Lakes region, and negatively impacts the ecosystem by interfering with local fish populations. At least two 5β-reduced 3α-hydroxyl steroids released by reproductive N. melanostomus (11-O-ETIO and 11-O-ETIO-3s) evoke olfactory sensory responses from the olfactory epithelium of conspecifics, and water conditioned by reproductive males (containing these steroids) attracts female round gobies. In this study, we examined whether these putative pheromones, along with simultaneously-released 11-O-ETIO-17s, stimulate olfactory sensory responses from alternative fish species sharing the same ecosystem as N. melanostomus in the Great Lakes region. Rock bass (Ambloplites rupestris), bluegill sunfish (Lepomis macrochirus), pumpkinseed sunfish (Lepomis gibbosus), smallmouth bass (Micropterus dolomieu), and yellow perch (Perca flavescens) were the targets of an electro-olfactogram experiment designed to record responses to odors. When compared to round goby responses from previous studies, amino acids and the bile acid consistently elicited electro-olfactogram responses across all species, but only round gobies showed a response to the putative pheromones. This study supports the concept of conducting a pheromone trapping trial in the field without adversely affecting the olfactory responses of non-target fish in the area.     

Piscivory in recovering Lake Michigan cisco (Coregonus artedi): The role of invasive species

Contemporary conditions in Lake Michigan where cisco (Coregonus artedi) populations are expanding are vastly different from those encountered by the historic fish community. Invasive species introductions have substantially altered the Lake Michigan ecosystem in the last half century. Successful management efforts for cisco in Lake Michigan hinge on our ability to understand their contemporary ecology, especially diet. We collected 725 cisco stomachs opportunistically from commercial fisheries (2%) and in agency surveys (98%) over six years (2014–2019). The majority (70%) of stomachs were from East Grand Traverse Bay and 96% of these were collected at Elk Rapids. Additional samples were collected from Charlevoix (8%), Little Traverse Bay (11%), other sites in northern Lake Michigan (4%), Central Lake Michigan (6%), and Green Bay (1%). Our results indicated a high degree of piscivory, in contrast to historical and contemporary accounts of planktivory for cisco in the other Laurentian Great Lakes. The top three prey items by mass were not native to the Great Lakes and these accounted for 87% of all observed prey mass consumed: round goby (Neogobius melanostomus) (58%), Bythotrephes longimanus (15%), and alewife (Alosa pseudoharengus) (14%). Round goby dominated the prey in the spring and summer, while B. longimanus and alewife occurred more in summer and fall diets. The contemporary population of cisco in Lake Michigan has been able to uniquely capitalize on abundant invasive prey resources, which may be less limiting and more energy-rich than a more typical planktivorous cisco diet.     

Round goby (Neogobius melanostomus) and native fishes as potential nest predators of centrarchid species in the upper St. Lawrence River

Widespread invasion of Round Goby (Neogobius melanostomus) throughout the Great Lakes has raised concerns regarding increased egg predation on fish species. To better understand nest predation, we examined nesting habitat selected by three upper St. Lawrence River Centrarchid species and the predator assemblage at nests during the 2011 and 2012 egg incubation and larval periods. Following removal of guarding males by angling, 5-min observations were used to identify and enumerate predators at rock bass (Ambloplites rupestris; n = 81), pumpkinseed (Lepomis gibbosus; n = 80), and smallmouth bass (Micropterus dolomieu; n = 40) nests. Differences in nesting habitat among centrarchids corresponded with differences in nest predator assemblages along a gradient defined primarily by depth and substrate. Pumpkinseed nests in shallow depths with soft substrate were visited principally by minnow spp., but few round goby. Smallmouth bass nests at greater depth with hard substrates were frequented nearly exclusively by round goby, while rock bass nests at intermediate depth with a mix of hard and soft substrates were visited by round goby and yellow perch (Perca flavescens). Rock bass nests had a higher predator burden than pumpkinseed nests in 2011, but no differences were observed among centrarchid species in 2012. Round goby were a major component of the predators at rock bass and smallmouth bass nests. However, predation burden imposed by yellow perch was higher than round goby at rock bass nests. We conclude nesting habitat selection influences native and non-native egg predator assemblages, but whether round goby predation is additive or compensatory remains unclear.     

Updated invasion risk assessment for Ponto-Caspian fishes to the Great Lakes

Majority of invasive species discovered in the Great Lakes since 1985 are native to the Ponto-Caspian region, including species that have had strong negative impacts in the Great Lakes (for example, dreissenid mussels and the round goby). The rich biota of the Ponto-Caspian region coupled with a high volume of commercial shipping traffic strongly suggests that this region will continue to be a major source of invasive species to the Great Lakes. To assess invasion risk by Ponto-Caspian fishes that have not been included in previous studies, we reviewed English-language publications and untranslated European literature (published primarily in Russian), focusing on physiological and ecological traits that have proven useful in previous risk assessments. We then used discriminant analysis to identify fishes that had a high probability of becoming established, spreading, and having significant negative impacts in the Great Lakes. Our updated listing of high-risk Ponto-Caspian fishes includes five species identified previously (the Black and Caspian Sea sprat, Eurasian minnow, big-scale sand smelt, European perch, and monkey goby) and five additional species (the Black sea shad, Caspian tyulka, Volga dwarf goby, Caspian bighead goby, and black-striped pipefish). Of these ten species, four (the monkey goby, big-scale sand smelt, Caspian tyulka, and black-striped pipefish) are likely to survive ballast water exchange as eggs, larvae, or adults based on salinity tolerances. Our results can be used to focus ongoing surveillance and rapid response efforts by highlighting Ponto-Caspian fishes that pose the greatest risk of becoming established and having significant negative impacts in the Great Lakes.     

Use of wetland versus open habitats by round gobies in lakes Michigan and Huron: Patterns of CPUE,length, and maturity

It has been suggested that some Great Lakes coastal wetlands may be resistant to invasion by several non-indigenous species including round goby, Neogobius melanostomus. However, there is inconclusive evidence regarding how susceptible exposed fringing coastal wetlands, in particular, are to round goby invasion. Therefore, we quantified round goby catch per unit effort (CPUE) using fyke nets in the Beaver Archipelago of Lake Michigan, and the Les Cheneaux islands and Saginaw Bay regions of Lake Huron. In addition, we examined the influence of body size and maturity on round goby habitat use. Catch per unit effort from fyke nets was highest in the Beaver Archipelago, where wetlands were dominated by small, immature round gobies and open water habitats were dominated by large adults. Fyke net catches within Les Cheneaux sites were similar between habitats and differences in size and maturity were not observed. Conversely, very few round goby were captured in wetlands of Saginaw Bay where CPUE was moderate in open water. This indicates that some exposed fringing wetlands in the Great Lakes, specifically those with high productivity, could have a higher degree of resistance to round goby invasion.     

Activity,Aggression, and Habitat Use of Ruffe (Gymnocephalus cernuus) and Round Goby (Apollonia melanostoma) under Laboratory Conditions

Potential negative ecological interactions between ruffe Gymnocephalus cernuus and round goby Apollonia melanostoma (formerly Neogobius melanostomus) might affect the colonization dynamics of these invasive species where they are sympatric in the Great Lakes. In order to determine the potential for ecological interactions between these species, we examined the activity, aggression, and habitat use of round gobies and ruffe in single species and mixed species laboratory experiments. Trials included conditions in which food was concentrated (in light or darkness) or scattered. Results showed that ruffe were more active than gobies, particularly when food was scattered. Activity of both species was significantly lower during darkness. Round gobies were significantly more aggressive than ruffe, and total aggression was lower in mixed species trials. Habitat use by ruffe and round gobies overlapped considerably, but we observed significant differences between species in their use of specific habitats that depended on experimental conditions. Overall, ruffe used open habitats more often than did round gobies, primarily when food was scattered. Round gobies used rocks significantly more frequently than did ruffe, but their use of rock habitat decreased during dark conditions. Ruffe were found more often in plant habitats and less often near the wall of the pool in trials during daylight with concentrated food. Activity and habitat use of ruffe and round goby did not significantly differ between single and mixed species trials. Overall, we found little evidence for negative ecological interactions between ruffe and round goby in these laboratory experiments.     

Exploring potential drivers of spatiotemporal variation in length-at-age and condition of two common nearshore fishes in southwestern Lake Michigan

Understanding factors like fish size and condition can be crucial to fisheries management. In the Great Lakes, long-term observations of these factors are used to describe the effects of ecosystem dynamics on fish ecology. Considering the diversity of available benthic habitat and dynamic environmental conditions in the nearshore zone of Lake Michigan, we sought to understand the variability in length-at-age and condition of yellow perch and round goby, two important members of the nearshore fish community. We measured these metrics from both species captured in gillnets across three locations in Illinois waters of southwest Lake Michigan from June through October 2008–2012 and related the spatiotemporal differences seen to observed variation in temperature, prey abundance, and competitor abundance. Yellow perch length-at-age varied annually for all three age-classes, with differences relating to thermal regime (age-0 and age-2) and alewife abundance (age-1). Yellow perch condition, measured as residual weight, varied spatially and annually, showing a negative relationship with round goby abundance. Round goby length-at-age was greater at the southern location, likely due to warmer temperatures, while condition was greater at the rockier location. In general, model selection analysis revealed several potential relationships between the relative abundances of competitor species and length and condition of yellow perch and round goby, while prey abundance may be related to round goby condition. These results demonstrate the importance of understanding the diversity in nearshore habitats and the influence of this diversity on factors affecting fish production, including size and condition, in southwest Lake Michigan.