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.     

Invasive round goby Neogobius melanostomus has sex‐dependent locomotor activity and is under‐represented in catches from passive fishing gear compared with seine catches

The higher proportion of males of the invasive round goby Neogobius melanostomus in samples from two activity selective passive fishing gears compared with one activity non‐selective fishing gear in three Dutch lakes is related to higher male locomotory activity and is a sex‐dependent trait. This difference in activity reflects the different ecology of male and female N. melanostomus.     

Metazoan parasites of Neogobius fishes in the Slovak section of the River Danube

Metazoan parasites were investigated in three non‐native fishes (monkey goby Neogobius fluviatilis, bighead goby Neogobius kessleri and round goby Neogobius melanostomus) collected from the former and current main channel of the River Danube and from the River Hron, Slovakia, in November 2003. Thirteen parasite species were identified: Triaenophorus crassus, Diplostomum sp., Tylodelphys clavata, Metorchis xanthosomus, Nicolla skrjabini, Gyrodactylus proterorhini, Pomphorhynchus laevis, Contracoecum sp., Raphidascaris acus, Anguillicola crassus, Unio tumidus, Anodonta anatina and Pseudoanodonta complanata. The maximum parasite diversity was found in N. fluviatilis. Total parasite abundance was significantly higher in N. kessleri, but no significant differences among sampling sites were observed. Pomphorhynchus laevis and glochidia of Anodonta anatina reached 100% prevalence in N. kessleri in the new channel of the Danube and, in general, these species were also the most prevalent parasites in all three goby species. For endoparasites, gobies served mostly as intermediate (digenean, cestodes and nematodes) or paratenic (acantocephalan and nematodes) hosts. All parasite species found are common parasites in the Middle Danube basin. No parasites specific to Neogobius, known from their native populations, were observed.     

Features of biology of the round goby <Emphasis Type="Italic">Neogobius melanostomus</Emphasis> (Perciformes,Gobiidae) in waters of Kuibyshev Reservoir

Based on materials of 2003–2005, the size-age composition, dates of reproduction, distribution, as well the qualitative and quantitative composition of the food of the round goby Neogobius melanostomus in the coastal part of Kuibyshev Reservoir are considered. It is shown that the round goby prefers coastal parts of the reservoir with depths to 10 m and rocky grounds and that its main food are mollusks of the genus Dreissena.     

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.     

Seasonal plasticity in telencephalon mass of a benthic fish

To gain a deeper understanding of how environmental conditions affect brain plasticity, brain size was explored across different seasons using the invasive round goby Neogobius melanostomus. The results show that N. melanostomus had heavier telencephalon in the spring compared to the autumn across the two years of study. Furthermore, fish in reproductive condition had heavier telencephala, indicating that tissue investment and brain plasticity may be related to reproductive needs in N. melanostomus.     

The recent distribution and abundance of non-native Neogobius fishes in the Slovak section of the River Danube

The distributions of invasive Neogobius species were investigated in the Slovak section of the River Danube from Bratislava downstream to the village of Chl'aba. During October 2004, the main channel of the Danube was sampled, including by‐pass, head‐race and tail‐race canals of the Gab?íkovo dam, backwaters and the lower‐most sections of the tributaries Malý Dunaj, Hron, Váh and Ipel’. Three Neogobius species already documented in Slovakia were captured (monkey goby Neogobius fluviatilis, bighead goby N. kessleri, round goby N. melanostomus), with the latter two species being found in almost all stretches of the Slovak Danube. Monkey goby had a most limited distribution, and no racer goby N. gymnotrachelus were observed. The abundance of particular Neogobius species appeared to depend on the character of the shoreline habitat, and a possible association between larger towns and the abundance of bighead and round gobies requires further investigation.     

Ontogenetic variability in external morphology of bighead goby Neogobius kessleri from the Middle Danube, Slovakia

Over the last decade, four species of goby have invaded the Middle Danube area, and all of them have spread rapidly. In the early 1990s, bighead goby Neogobius kessleri appeared in the Middle Danube, where it now seems to thrive. Relatively little is known of the environmental biology and ontogeny of this species in its native and non‐native ranges. In this paper, preliminary results on the external morphology of bighead goby from the Slovak stretch of the Danube are presented within an ontogenetic context. Patterns of relative growth with no apparent changes at small size suggest direct development in bighead goby, although not as profoundly direct as observed in round goby N. melanostomus. Differences in life history between these two closely related species may have important implications for their success in novel environments, favouring the latter in short term (several years) and the former in long term (decades and longer) perspective.     

Historic speciation and recent colonization of Eurasian monkey gobies (Neogobius fluviatilis and N. pallasi) revealed by DNA sequences,microsatellites, and morphology

Aim Hidden diversity within an invasive ‘species’ can mask both invasion pathways and confound management goals. We assessed taxonomic status and population structure of the monkey goby Neogobius fluviatilis across Eurasia, comparing genetic variation across its native and invasive ranges. Location Native populations were analysed within the Black and Caspian Sea basins, including major river drainages (Dnieper, Dniester, Danube, Don and Volga rivers), along with introduced locations within the upper Danube and Vistula river systems. Methods DNA sequences and 10 nuclear microsatellite loci were analysed to test genetic diversity and divergence patterns of native and introduced populations; phylogenetic analysis of mtDNA cytochrome b and nuclear RAG‐1 sequences assessed taxonomic status of Black and Caspian Sea lineages. Multivariate analysis of morphology was used to corroborate phylogenetic patterns. Population genetic structure within each basin was evaluated with mtDNA and microsatellite data using F_ST analogues and Bayesian assignment tests. Results Phylogenetic analysis of mitochondrial and nuclear sequences discerned a pronounced genetic break between monkey gobies in the Black and Caspian Seas, indicating a long‐term species‐level separation dating to c. 3 million years. This pronounced separation further was confirmed from morphological and population genetic divergence. Bayesian inference showed congruent patterns of population structure within the Black Sea basin. Introduced populations in the Danube and Vistula River basins traced to north‐west Black Sea origins, a genetic expansion pattern matching that of other introduced Ponto‐Caspian gobiids. Main conclusions Both genetic and morphological data strongly supported two species of monkey gobies that were formerly identified as subspecies: N. fluviatilis in the Black Sea basin, Don and Volga Rivers, and the Kumo‐Manych Depression, and Neogobius pallasi in the Caspian Sea and Volga River delta. Genetic origins of introduced N. fluviatilis populations indicated a common invasion pathway shared with other introduced Ponto‐Caspian fishes and invertebrates.     

Genetic and biogeographic relationships of the racer goby Neogobius gymnotrachelus (Gobiidae: Teleostei) from introduced and native Eurasian locations

The racer goby Neogobius gymnotrachelus , along with several other neogobiin fishes, has been spreading north and west from its native Ponto-Caspian range for the past two decades via shipping and canals. It has been predicted as a likely future invader of the North American Great Lakes, where it would join its neogobiin relatives – the round and freshwater tubenose gobies. The present study is the first to analyse the population genetic and biogeographic relationships of the racer goby, establishing a baseline to aid interpretation of its future spread patterns and likely donor-recipient population relationships. The mitochondrial DNA cytochrome b gene was sequenced from representative areas of the racer goby's range, including rivers of the northern Black Sea and areas of spread upstream in the Danube River and outside the Ponto-Caspian region to the Vistula River in Poland. Results discerned nine haplotypes, with few shared among drainages of native rivers and most sites housing unique alleles. Racer goby populations significantly diverged among watersheds, supporting historic low migration and little non-anthropogenic gene flow. The Dnieper River was identified as a likely donor source for the Vistula River colonization, where appreciable variability suggests a relatively large number of founding genotypes.     

Erythrocyte profile of circulating blood of Neogobius melanostomus (Pallas, 1814) under conditions of experimental hypothermia

The influence of hypothermia on erythrocyte profile of thermophile teleost species round goby, Neogobius melanostomus (Pallas, 1814), has been studied. Fish were acclimated to temperature 1-2^оС, 15-16^оС and 19-20^оС (control group) and held at given conditions for 5 days. The number of red blood cell precursors (pronormoblasts, basophilic and polychromatophilic normoblasts) in circulating blood has been estimated. Also, the number of abnormal erythrocytes, i.e. cells with micronuclei, nuclei invaginations, red blood cell shades, dacryocytes and cells undergoing amitosis has been determined on smears. The number of immature erythrocytes increased more than two times (p < 0,001) at 1-2^оС. The number of low-differentiated precursors, pronormoblasts and early basophilic normoblasts, increased for the most part. The number of abnormal erythrocytes did not change substantially, The changes in cellular blood composition were accompanied with the increase of plasma lactate concentration, indicating hypoxic state of fish. The results of the present work indicate that hematopoietic tissue remains sensitive to controlling factors at hypothermia, such as hypoxia, and may enhance proliferation and differentiation of erythroid cells.     

Ontogenetic variability in external morphology of round goby Neogobius melanostomus from Middle Danube, Slovakia

In the previous decade, four species of non‐native goby have invaded the middle section of the River Danube. To understand the problems associated with these invasions better, we examined the external morphology of 184 round goby Neogobius melanostomus specimens using both triple regression (distance‐based measurements) and geometrical (coordinates‐based measurements) analysis within an ontogenetical and epigenetical aspect. Using geometrical analysis of distance‐based characters, we found that the external body shape in four size groups of round goby remained essentially unchanged. Such direct development represents a strongly precocial (i.e. specialized) life‐history, whereas earlier maturation in this non‐native population suggests a shift back towards more altricial (i.e. less specialized, more generalized) life history. This combination of altricial–precocial trajectories seems to be typical for invasive round gobies and may be one of key factors for their successful colonization of new environments.     

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.     

Morphological divergence and diagnostic characters of ginger goby <Emphasis Type="Italic">Neogobius eurycephalus</Emphasis> (Kessler, 1874) and surman goby <Emphasis Type="Italic">N. cephalargoides</Emphasis> Pinchuk, 1976 (Perciformes,Gobiidae)

Comparative analysis of morphometric characters of surman goby Neogobius cephalargoides and of two subspecies of ginger goby N. eurycephalus is made. Characteristics liable to size variation and sex dimorphism are revealed. Independence of the investigated species and the diagnostic value of characters previously suggested for classification are confirmed. The obtained data confirm isolation of north-western populations of ginger goby as a separate subspecies N. eurycephalus odessicus and suitability of three meristic characters for differentiation of the subspecies: the number of rays in the second dorsal fin and anal fin and the number of transverse rows of scales.     

Some features of biology of big-headed goby <Emphasis Type="Italic">Neogobius gorlap</Emphasis> (Perciformes,Gobiidae) in waters of the Kuibyshev Reservoir

On the basis of materials of 2003–2005, the size-age composition, dates of reproduction, as well as qualitative and quantitative food composition of big-headed goby Neogobius gorlap in the coastal part of the Kuibyshev Reservoir are considered. It is shown that big-headed goby prefers coastal parts of the reservoir with depths up to 2.5 m and rocky, sandy-slimy grounds; the bulk of its feeding consists of crustaceans.     

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.     

What can morphology tell us about ecology of four invasive goby species?

This study presents a detailed comparative analysis of external morphology of four of the most invasive goby species in Europe (round goby Neogobius melanostomus, bighead goby Ponticola kessleri, monkey goby Neogobius fluviatilis and racer goby Ponticola gymnotrachelus) and interprets some ecological requirements of these species based on their morphological attributes. The results are evaluated within an ontogenetic context, and the morphological differences between the species are discussed in terms of the question: can special external shape adaptations help to assess the invasive potential of each species? The morphometric analyses demonstrate important differences between the four invasive gobies. Neogobius melanostomus appears to have the least specialized external morphology that may favour its invasive success: little specialization to habitat or diet means reduced restraints on overall ecological requirements. The other three species were found to possess some morphological specializations (P. kessleri to large prey, N. fluviatilis to sandy habitats and P. gymnotrachelus to macrophytes), but none of these gobies have managed to colonize such large areas or to reach such overall abundances as N. melanostomus.     

Isolation and characterization of microsatellites in Neogobius kessleri (Perciformes,Gobiidae) and cross‐species amplification within the family Gobiidae

Fourteen polymorphic microsatellites were isolated from Neogobius kessleri, a benthic fish of Ponto‐Caspian origin which has been recently introduced into the Middle and Upper Danube River. Number of alleles and heterozygosity per locus in a sample of 32 fish individuals ranged from two to four and from 0.13 to 0.75, respectively. These primers will be useful in determining the population structure of N. kessleri. In addition, successful cross‐amplification was obtained for four related species, N. melanostomus, N. fluviatilis, N. gymnotrachelus and Proterorhinus marmoratus. These microsatellite loci may be useful for the evaluation of the origin of non‐native goby populations.     

Ontogenetic variability in the external morphology of monkey goby, Neogobius fluviatilis (Pallas, 1814) and its relevance to invasion potential

In the previous decade, four species of non-native gobies have invaded the middle section of the river Danube and its tributaries. An effective tool for understanding biological invasions is the evaluation of various biological traits (morphological, life history, ontogenetic) within an epigenetic context. The present study examines the external morphology of monkey goby Neogobius fluviatilis (Pallas, 1814) from the mouth of the River Hron, the morphological differences among three goby species (monkey, bighead and round) and the relevance of these differences for invasive potential. Monkey goby reach their definite phenotype very early in their ontogeny and thus represent a strongly precocial (specialized) species with direct development. The morphological differences between monkey and two other goby species also reflect its strong specialization for sandy substrata and smaller prey types. Thus, monkey goby are not expected to spread to new areas as fast as the round and bighead gobies, and their distribution is likely to be limited to sandy and/or sandy-gravel substrata. If this assumption is correct, then the potential adverse impact of monkey goby on native fauna or even ecosystem is likely to be less than that of the bighead and round gobies. Handling editor: K. Martens     

Invasion Genetics of Ponto-Caspian Gobies in the Great Lakes: A ‘Cryptic’ Species, Absence of Founder Effects, and Comparative Risk Analysis

Genetic variability and structure of nonindigenous vs native populations are compared for the Eurasian round goby Neogobius melanostomus and the tubenose goby Proterorhinus marmoratus, which both invaded Lake St. Clair of the North American Great Lakes about 1990. The round goby spread rapidly to all of the Great Lakes and the tubenose goby largely has been restricted to Lake St. Clair, with some recent range extension into western Lake Erie. Risk analyses may indicate whether genetic variability of colonizers is predictive of their relative invasive and establishment successes. The present investigation examined DNA sequence variation across the left domain of the mitochondrial DNA cytochrome b gene in round and tubenose gobies from Eurasian and Great Lakes locations. We also sequenced six additional Neogobius species (including the monkey N.␣fluviatilis, racer N. gymnotrachelus, and bigheadN. kessleri gobies that have been ‘on the move’ in Europe) and the knout goby Mesogobius batrachocephalus from the Black Sea in order to develop diagnostic genetic characters to identify them in case of future and/or undetected invasions and to delineate their phylogenetic relationships. Results show that a diverse number of haplotypes characterize round and tubenose goby populations from both North America and Eurasian sites, fitting a risk analysis prediction of high genetic variability in their successful introductions. Phylogenetic results indicate that the current genus Neogobius is paraphyletic and that the subgenusApollonia thus should be elevated to the level of genus, containingApollonia (N.) melanostomus (the round goby) andA. (N.) fluviatilis (the monkey goby). In addition, there appear to be two separate species of Proterorhinus marmoratus, a marine P. marmoratus Pallas 1814 in the Black Sea (matching the original type locality), and a ‘cryptic’ freshwater species in the Danube and Dnieper Rivers and probably other Eurasian freshwater habitats, as well as invasive in the Great Lakes. We suggest resurrecting the name P. semilunaris Heckel 1837 for the freshwater species (a taxon that was originally described from rivers draining into the Aegean Sea and the Danube River, but was later placed in synonymy with P. marmoratus). An erratum to this article is available at .