Predation on Walleye Eggs by Fish on Reefs in Western Lake Erie

We examined diets of fishes from gillnet and egg pump collections conducted on reefs in western Lake Erie during walleye (Sander vitreus) egg incubation periods from 1994–1999 and 2004 to assess incidence of walleye eggs in fish diets. We collected no potential egg predators in samples taken in 1994 but from 1995–1999 and in 2004 we caught 22 different species of fish on reefs in addition to spawning walleye. In most years, white perch (Morone americana) stomachs contained more walleye eggs than any other species on the reefs averaging 253 eggs per stomach. We also found lower numbers of walleye eggs in the stomachs of channel catfish (Ictalurus punctatus; 53 eggs/stomach), johnny darter (Etheostoma nigrum; 2 eggs/stomach), logperch (Percina caprodes; 10 eggs/stomach), quillback (Carpiodes cyprinus; 184 eggs/stomach), rock bass (Ambloplites rupestris; 3 eggs/stomach), round goby (Neogobius melanostomus; 4 eggs/stomach), sculpin (Cottidae; 21 eggs/stomach), silver chub (Macrhybopsis storeriana; 3 eggs/stomach), spottail shiner (Notropis hudsonius; 14 eggs/stomach), trout-perch (Percopsis omiscomaycus; 30 eggs/stomach), white sucker (Catastomus commersonii; 20 eggs/stomach), and yellow perch (Perca flavescens; 181 eggs/stomach). Similar to other studies of predation on walleye eggs, our results indicate that prolonged incubation periods increase the potential for egg loss due to predation.     

Are interactions among Ponto-Caspian invaders driving amphipod species replacement in the St. Lawrence River?

In Lake Erie and Lake Ontario, the Ponto-Caspian amphipod Echinogammarus ischnus has replaced the native amphipod Gammarus fasciatus on rocky substrates colonized by dreissenid mussels, which provide interstitial refugia for small invertebrates. Based on the premise that an invader's vulnerability to predation is influenced by its evolutionary experience with the predator and its ability to compete for refugia, we hypothesized that amphipod species replacement is facilitated through selective predation by the round goby Neogobius melanostomus, a Ponto-Caspian fish that invaded the Great Lakes in the early 1990s and is now colonizing the St. Lawrence River. In laboratory experiments, we determined if E. ischnus excludes G. fasciatus from mussel patches, and if the vulnerability of G. fasciatus to predation by gobies is increased in the presence of the invasive amphipod. E. ischnus and G. fasciatus did not differ in their use of mussel patches, either when alone or in each other's presence. Both species were equally vulnerable to predation by the round goby. In field experiments, we determined if the round goby exerts a stronger impact than native predators on the relative abundance of amphipod species. Our results suggest that E. ischnus is more vulnerable to native predators, but the round goby does not have a differential impact on the native amphipod. We conclude that competition with E. ischnus does not increase the vulnerability of G. fasciatus to goby predation, and that the round goby does not promote the replacement of G. fasciatus by E. ischnus in the St. Lawrence River. The outcome of antagonistic interactions between exotic and native amphipods is mediated more by abiotic factors than by shared evolutionary history with other co-occurring exotic species.     

Comparison of Methods Needed to Estimate Population Size of Round Gobies (Neogobius melanostomus) in Western Lake Erie

The round goby (Neogobius melanostomus) is a small, demersal fish that was introduced into the Great Lakes basin in 1990. Since their arrival, the round goby has been implicated in many ecological changes—most notably changes in the flow of energy from the benthic to the pelagic food web through their consumption of dreissenid mussels. However, methods for evaluating the density and size of round gobies across different substrates are lacking, preventing the true quantification of the effects of round gobies on invaded ecosystems. In our study, we evaluated catch efficiency of numerous passive and active sampling methods for capturing round gobies. We then applied the best techniques to estimate the distribution, density, and biomass of round gobies in western Lake Erie. Visual census (underwater video transects) proved the best technique for assessing round goby size and density across a wide range of substrates. A combination of angling and bottom trawling proved most effective for obtaining biological samples. We estimated 9.9 billion round gobies in western Lake Erie in 2002. Continued efforts to describe abundance and demographics of round gobies in invaded ecosystems will enable scientists and managers to fully understand the impacts of this invading species.     

Do native predators feed on non-native prey? The case of round goby in a fluvial piscivorous fish assemblage

We evaluated whether or not the invasive round goby (Neogobius melanostomus) represents an important prey source for seven native fish predators in Lake St. Pierre (St. Lawrence River, Canada). The frequency of occurrence of round goby in the stomach contents of brown bullhead (Ameiurus nebulosus) and channel catfish (Ictalurus punctatus) was very low (< 5%), while for the five other predators, it varied between 22% (yellow perch; Perca flavescens) and 65% (sauger; Sander canadensis). Several competing models linking the probability of occurrence of round goby in stomach contents to variables related to space, physical habitat, biotic interactions and predator size were tested for the five species feeding on round goby. Results indicated that space variables influenced round goby occurrence in stomachs for all species. In addition, physical habitat variables had an influence for sauger and walleye (Sander vitreus); biotic variables had an influence for yellow perch, walleye and sauger; and size had an influence for northern pike (Esox lucius), smallmouth bass (Micropterus dolomieu) and walleye. These results are discussed in light of known biological features of the round goby and native predators studied here and have important implications in terms of understanding round goby invasion success in the Great Lakes–St. Lawrence system.     

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.     

Re-engineering the eastern Lake Erie littoral food web: The trophic function of non-indigenous Ponto-Caspian species

The trophic roles of key Ponto-Caspian invaders (quagga mussels Dreissena bugensis, amphipods Echinogammarus ischnus and round goby Apollonia melanostomus) within the littoral food web of eastern Lake Erie were quantified using stable isotopes (δ¹³C, δ¹⁵N). A dual stable isotope parameter search with a mass balance component was used to assess the isotopic importance of quagga mussels and amphipods as dietary items to two size classes of round goby. The utility of the mass balance simulation was also evaluated as a tool to approximate isotopic contributions of feasible prey and identify gaps incurred by “missing” prey items not included in the sampling. The mass balance dietary simulation, confirmed by stomach content data, indicated that isotopically important prey to small round goby (< 11.2 cm) were chironomids and Ponto-Caspian amphipods, while large round goby (≥ 11.2 cm) showed strong preference for quagga mussels. The dietary mass balance simulation output also supported the isotopic importance of round goby to the somatic growth of smallmouth bass, rock bass and freshwater drum. The isotopic mass balance output for yellow perch was more ambiguous, which may be in line with their known broadly omnivorous diet. The white bass output was in line with published data indicating increasing consumption of round goby for this species, while the brown trout output strongly favoured alewife isotopic contributions. However for white perch and walleye, the mass balance simulations were not in line with their known published diets in Lake Erie, probably due to a lack of key prey items in the sample set (e.g. zooplankton for white perch and shiner species for walleye). As expected, the Ponto-Caspian species have integrated themselves into the littoral food webs, and the “quagga mussel–round goby–smallmouth bass” food chain forms one of the key components within the trophodynamics of Lake Erie.     

Regional Differences in Size-at-age of the Recovering Burbot (Lota lota) Population in Lake Erie

The burbot Lota lota population in Lake Erie increased dramatically between 1995 and 2003, due mainly to control of the sea lamprey Petromyzon marinus, which began in the late 1980s. We estimated total length- and weight-at-age at capture for burbot caught in annual gillnet surveys of eastern Lake Erie during August 1994–2003. Mean total length was generally greater for burbot age 4–9 years that were caught in New York waters than in either Ontario or Pennsylvania waters of Lake Erie. Similarly, mean weight was greater for burbot at ages 4 through 6 years in New York waters than in either Ontario or Pennsylvania waters. Age-9 burbot caught in Ontario waters had greater mean weight and mean total length than did age-9 burbot caught in Pennsylvania waters. One possible explanation for greater length- and weight-at-age for New York burbot may be greater abundance of prey fishes, particularly rainbow smelt Osmerus mordax and round goby Neogobius melanostomus in New York waters. Total lengths at ages 4–10 years were generally greater for burbot caught in Lake Erie during 1994–2003 than those from published studies of other large lakes in North America that we considered, including for Lake Erie in 1946. The regional differences in size-at-age have important management ramifications, particularly because a commercial fishery targeting burbot has been considered for Ontario waters of Lake Erie.     

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.”     

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.     

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.     

The Zebra Mussel Dreissena polymorpha Found in North America in 1986 and 1987

The zebra mussel Dreissena polymorpha was first detected in the western basin of Lake Erie, Ontario, Canada, on natural gas wellheads and well markers between April and November 1986. It was found again in 1987 on the north shore of Lake Erie in a water treatment plant, and in vessel fouling. The population increased in Lake Erie in 1988. Dreissena may have spread from Lake Erie to Lake St. Clair, where it was then discovered on 1 June 1988.     

Distribution of the Ponto-Caspian Amphipod Echinogammarus ischnus in the Great Lakes and Replacement of Native Gammarus fasciatus

The gammarid amphipod Echinogammarus ischnus was found to be widespread from the south end of Lake Huron, downstream in the St. Clair River and across Lake Erie to the Niagara River outlet into Lake Ontario. The presence of this exotic species was first reported in the Detroit River, where it now dominates; this species has been common in western Lake Erie since the summer of 1995. The species has replaced the native amphipod Gammarus fasciatus on rocky habitats in the St. Clair, Detroit, and Niagara rivers, and is the dominant amphipod on rocky shores in western Lake Erie. In one year, E. ischnus became the dominant amphipod at the Lake Ontario end of the Welland Canal, although the fecundity of E. ischnus is less than G. fasciatus. E. ischnus has not yet been reported from the north shore of Lake Ontario or the outlet into the St. Lawrence River but occurs 100 km further downstream at Prescott.     

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.     

Effects of Food Type,Habitat, and Fish Predation on the Relative Abundance of Two Amphipod Species,Gammarus fasciatus and Echinogammarus ischnus

We evaluated the abundance patterns of Gammarus fasciatus and Echinogammarus ischnus in dreissenid and macrophyte areas in Hatchery Bay, Lake Erie before (1997) and after round goby (2001, 2002) invaded the area. Total amphipod abundance was higher before round goby invasion in both habitats. In mussel beds, E. ischnus abundance was similar or significantly higher than G. fasciatus. In macrophytes, G. fasciatus was significantly more abundant than E. ischnus. In laboratory experiments, we compared amphipod survivorship and growth when fed mussel feces and pseudofeces (F+P) or macrophytes with epiphytes (M+E). Gammarus fasciatus survivorship and growth were higher when fed F+P than M+E. Echinogammarus ischnus showed similar survivorship under both diets, but significantly higher growth when fed M+E than F+P. Therefore inter-habitat differences in food resources cannot explain the abundance patterns observed in the lake. We also estimated the relative vulnerability of G. fasciatus and E. ischnus to yellow perch (Perca flavescens) and round goby (Neogobius melanostomus) predation in laboratory feeding trials using mussel colonies or macrophyte beds as substrate. Both fish strongly preferred E. ischnus in macrophytes, but consumed relatively more G. fasciatus than E. ischnus in dreissenid habitats. Our results suggest that dreissenid establishment may have facilitated the invasion of E. ischnus. However, habitat-specific differences in vulnerability to fish predation may mediate the coexistence of G. fasciatus and E. ischnus by minimizing expansion of E. ischnus to macrophyte areas. Our results also suggest that round goby invasion can alter amphipod abundance patterns in Lake Erie.     

Dietary niche and growth rate of the nonnative tubenose goby (Proterorhinus semilunaris) in the Lake Superior basin

The tubenose goby (Proterorhinus semilunaris) entered the Great Lakes in the 1990s via ballast water, but remains poorly studied within North America, making it difficult to predict its effects on native ecosystems. Dietary breadth and somatic growth rate have important ramifications for survival, competitiveness, and dispersal ability of a fish species, and thereby its ecological impact. We studied diet and growth of age-0 tubenose goby within the St. Louis River, a tributary to Lake Superior that contains the largest population within the Lake Superior basin. We sampled tubenose gobies from shallow, vegetated habitats during summer and fall. Stomach contents were identified and weighed to measure fullness and dietary breadth between seasons and several locations. We aged fish based on otolith daily increments to model somatic growth. Diet was dominated by isopods and amphipods, and dietary breadth was low and not significantly different between locations and seasons. Tubenose goby diet strongly overlapped with that of tadpole madtom (Noturus gyrinus), a native, demersal species. We tested several candidate growth models; the Gompertz growth function was the most parsimonious model among those examined. The model demonstrates that tubenose goby obtains a small maximum size and is short-lived. We conclude that tubenose goby presents a unique risk to the Great Lakes and other freshwater bodies because their life history is typical of invasive species, their diet overlaps with native fish, and because they occupy shallow, vegetated habitat which functions as both nursery and foraging habitat for many native fishes.     

The effects of riprap in enhancing the abundance and coexistence of Gobiidae along the southern Caspian Sea coast

Many natural coastal habitats around the world are being replaced by hard coastal defense structures as a result of rising sea levels and increasing intensity of storms. To examine the potential effects of riprap on gobies in the nearshore zone of the southern Caspian Sea, we used angling to compare their abundance and diversity in 13 riprap and natural habitats (sandy and gravel shores) from April 2012 through April 2013. In total, 819 gobies of three species, Caspian sand goby Neogobius pallasi (N?=?433), round goby Neogobius melanostomus (N?=?206), and Caspian goby Neogobius caspius (N?=?180) were collected. No round gobies were caught from sandy shores and only two Caspian gobies were caught from gravel habitats, while all three species were abundant in riprap shores. A significant difference was observed in CPUE of all three goby species among shore types, with riprap sites having the highest abundance. PERMANOVA and nMDS demonstrated differences in goby assemblages among all three habitats. There were significant differences in round goby and Caspian goby sizes among riprap and natural habitats, while there were no significant differences in Caspian sand goby size among habitat types. In general deployment of riprap, especially in an area characterized as having natural shores with low structural complexity, could enhance the abundance and coexistence of nearshore gobies and act as a dispersal vector.     

Factors governing the distribution and fish-community associations of the round goby in Michigan tributaries of the Laurentian Great Lakes

The round goby (Neogobius melanostomus) is increasingly being reported in tributaries of the Laurentian Great Lakes where these fish have been shown to adversely impact native stream biota. Determining the characteristics and distribution of invaded streams are the first steps toward effective round goby management. We sampled 30 tributaries in the Great Lakes basin and characterized each in terms of nine physical reach-scale attributes. Round goby were detected in 14 streams where abundances ranged from 4% to 53% of the fish sampled in each stream. Round goby was the single most abundant fish species sampled, constituting 14% of all fish encountered across all sites, and 30% of individuals in round goby-present sites. Round goby-present sites were larger, had lower channel slopes, less large wood, and less canopy cover than round goby-absent sites, suggesting that these attributes may promote round goby establishment. Mottled sculpin, cyprinids, brook stickleback, white sucker and rainbow trout were associated with goby absence while centrarchids, percids, yellow bullhead, and mud minnow were associated round goby presence. Collectively these results demonstrate that round goby are widespread in eastern Michigan tributaries to the Great Lakes, present in streams with a range of physical habitat characteristics, and that round goby presence is associated with certain fish species.     

How Non-native Species in Lake Erie Influence Trophic Transfer of Mercury and Lead to Top Predators

Lake Erie's food web has been dramatically modified by exotic species. Both exotic dreissenid mussels and the round goby Neogobius melanastomus have shifted the food web from a pelagicbased to a benthic-based one, potentially creating a new pathway for contaminant transfer to top predators. Before the invasion of round gobies, few predators of dreissenids occurred in Lake Erie, allowing contaminants to be confined to these benthic organisms. The invasion of the round goby has produced a new pathway through which these contaminants can enter the food web. To characterize heavy-metal transfer through this new food web and to assess risk to humans, water, surficial sediment, dreissenid, round goby, and smallmouth bass Micropterus dolomieui samples were collected at three sites during summers, 2002 and 2003, and analyzed for total lead (Pb), total mercury (Hg), and methyl mercury (MeHg). In addition, we compared smallmouth bass Pb and Hg concentrations to those measured in 1993/1994, before round gobies were prevalent. Pb biodiminished and MeHg biomagnified through the food web to smallmouth bass; patterns were similar among our three sites. Total Pb concentrations in smallmouth bass were higher before the incorporation of round gobies into their diet. We attributed this decline to changes in food web structure, changes in contaminant burdens in prey, or declines in sediment Pb concentrations in Lake Erie. By comparison, Hg concentrations in smallmouth bass changed little, before and after the round goby invasion, possibly due to a shift in diet that increased growth. Despite a decline in sediment Hg concentrations in Lake Erie, smallmouth bass continued to accumulate Hg at historical rates possibly because of their high consumption rates of benthivorous round gobies. As smallmouth bass continue to consume round gobies during their lives, their Hg concentrations may well continue to increase, potentially increasing the risk of Hg contamination to humans.     

Genetic and Biogeographic Relationships of the Invasive Round (Neogobius melanostomus) and Tubenose (Proterorhinus marmoratus) Gobies in the Great Lakes Versus Eurasian Populations

Population genetic structure and systematic relationships were investigated for two exotic fishes introduced to the Great Lakes in 1990, the round goby Neogobius melanostomus and the tubenose goby Proterorhinus marmoratus, using DNA sequences from the left domain of the mitochondrial DNA control region. Samples of round gobies were compared from different sites in the Great Lakes, an introduced population from the Gulf of Gdansk in Poland, and a native population from the northern Black Sea. The round goby was characterized by relatively high genetic variability, and 17 haplotypes were identified from 64 individuals. Levels of genetic variation for the round goby were similar in the invasive and native sampling sites, suggesting relatively large founding populations and lack of bottlenecks. The northern Black Sea was eliminated as a probable founding source for both the Great Lakes and the exotic population in Poland. Substitutions in the left domain of the control region revealed significant differences among samples from the Great Lakes and Eurasia, and between Lakes Erie and St. Clair, suggesting non-random mating. No variation was detected in the tubenose goby population in the Great Lakes, which has been less successful in terms of spread and population growth. A molecular clock calibration suggested that the genera Neogobius and Proterorhinus diverged about 5.2 ± 1.0 million years ago, apparently separating from a common ancestor shared with Gobius during the isolation of the Paratethys basin from the Mediterranean Tethys Sea.