Checklist of parasites for Ponto-Caspian gobies (Actinopterygii: Gobiidae) in their native and non-native ranges

This study provides a comprehensive checklist of parasites reported for native and non-native populations of seven Ponto-Caspian goby species, i.e. Babka gymnotrachelus, Neogobius fluviatilis, N. melanostomus, Ponticola gorlap, P. kessleri, Proterorhinus semilunaris and P. semipellucidus, all of which have recently expanded their natural range, either by transoceanic translocation or along the three main European intracontinental invasion corridors. In some cases, invasion success is facilitated by loss of natural enemies, such as predators and parasites; thus, knowledge of a species natural parasite fauna is essential for assessments of enemy release in the host's non-native range. The study is based on published literature and personal unpublished data related to parasites of Ponto-Caspian gobiids. A total of 249 parasite taxa have been reported during 1931—2019, with higher numbers reported from native (214) than non-native (151) ranges. This checklist indicates apparent reduction in native parasites in non-native ranges in all goby species, although parasite colonisation in non-native ranges differs between fish species and invasion corridors. Interestingly, the cumulative number of species reported is still gradually increasing in both ranges. Acquisition of high number of local parasite species also indicates competence of Ponto-Caspian gobies to a wide range of generalist parasites.     

Interspecific variation of acoustic signals in Mediterranean gobies (Perciformes, Gobiidae): comparative analysis and evolutionary outlook

Sound production of 11 Mediterranean goby species, belonging to five different genera, have been comparatively analysed on the basis of the quantitative properties of the acoustic signal emitted by the male in both the reproductive and aggressive context. The results obtained showed that three groups of species can be recognized on the basis of signal similarity: the larger sized species (genus Padogobius and Gobius paganellus ) producing tonal sounds, showing high values of pulse rate and low values of duration; the larger-sized species producing grunt sounds (genus Gobius and Zosterisessor ) with low pulse rate and low duration; and the small-sized species producing grunt sounds (genus Pomatoschistus and Knipowitschia ) with low pulse rate and high duration. The comparison between these results and those found in previous studies suggests congruence between the acoustic affinities among species and that obtained by means of morphological and genetic data. Furthermore, first hypotheses on the evolution of acoustic communication and the associated mechanisms in this fish group are suggested.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 763–778.     

Gobius xoriguer,a new offshore Mediterranean goby (Gobiidae), and phylogenetic relationships within the genus Gobius

Ichthyological Research - Gobius xoriguer sp. nov., a new ofshore species of goby (Teleostei: Gobiidae: Gobiinae) is described based on three specimens collected in 2010, 2012 and 2018 in the...     

The shallow water gobiid assemblage of the Venice Lagoon: abundance, seasonal variation and habitat partitioning

The gobiid assemblage of the Venice Lagoon shallow waters was investigated by means of a semi‐quantitative standardized sampling (using a small beach seine), stratified into five main types of shallow subtidal habitats and conducted on a seasonal basis during 1 year. The degree of overlap in resource utilization among six coexisting goby species was assessed, along both the time axis, by analysing the seasonal variation in abundance and reproductive status (as revealed by the gonado‐somatic index) and the habitat axis, by comparing species abundance across different habitat types and controlling for the effects of some abiotic factors. Smaller species, and especially the marbled goby Pomatoschistus marmoratus , dominated the local assemblage. Although the cycle of shallow water colonization and seasonal variation in total abundance were basically similar, species showed differences in timing of reproduction and recruitment, as well as in habitat preference. The larger species belonging to the genera Gobius and Zosterisessor tended to overlap their habitat use, being more abundant in seagrass habitats than in the unvegetated habitats, whereas the smaller species belonging to the genera Knipowitschia and Pomatoschistus avoided seagrasses, preferring in most cases mud flats and salt marsh creeks. Within these two groups of species some further slight differences in species habitat preference, relationship with abiotic factors and reproductive ecology could be detected. Results are discussed in the light of both ecological mechanisms underlying coexistence of closely related species and the current knowledge of the phylogeny of Mediterranean gobies.     

Reconstruction of phylogenetic relations of Ponto-Caspian gobies (Gobiidae, Perciformes) based on mitochondrial genome variation and some problems of their taxonomy

Phylogenetic relations of Ponto-Caspian gobies in the context of different taxonomic hypotheses have been studied based on the analysis of variations in cytochrome b gene. Evidence for the paraphyly of the genus Neogobius sensu lato, represented by two independent phyletic lineages that should be treated as separate genera Neogobius s. stricto (includes a species group of N. melanostomus-N. fluviatilis-N. caspius) and Ponticola, has been provided. It has been demonstrated that the Racer goby N. gymnotrachelus refers to the latter genus. It is proved that the Grass goby Gobius ophiocephalus belongs to the genus Gobius s. stricto.     

Ecological plasticity of tubenose goby,a small invader in South Moravian waters

Hydrobiologia - The western tubenose goby Proterorhinus semilunaris, an invasive Ponto-Caspian fish species, has established populations in a wide range of habitat types in the Dyje/Morava river...     

Biogeographic zonation of the Black Sea and Caspian Sea basin based on Mysid fauna (Crustacea: Mysidacea)

A biogeographic analysis of the Black Sea and Caspian Sea basin and adjacent Aegean Sea and the Sea of Marmara was conducted based on the distribution of 55 mysid species. The Black and Caspian Sea basin proper is inhabited by 35 mysid species belonging to 12 genera, among them 26 species and 3 genera are endemic to the region. Ponto-Caspian and Caspian species are predominant. The explored areas were included into two provinces of the Mediterranean subrealm of the East-Atlantic Subtropical realm and three provinces of the Ponto-Caspian realm.     

Abundance and night-time habitat use patterns of Ponto-Caspian gobiid species (Pisces, Gobiidae) in the littoral zone of the River Danube, Hungary

We examined the abundance and meso‐habitat use of gobiid species during both day and night along 43 stretches (500‐m long) of the littoral zone at five locations of the Middle Danube, Hungary, in spring and in summer 2004. Electro‐fishing catch per unit effort sampling revealed significantly higher relative densities at night than during the day. Gobiids occupied all the available habitats encountered during sampling. Habitat‐abundance relationships from night‐time samples revealed that the two most abundant species, round goby Neogobius melanostomus and bighead goby Neogobius kessleri, were found in highest relative density along gravel beach and artificial rip‐rap habitats; these species were in relatively low density in sandy areas. Monkey goby Neogobius fluviatilis was more abundant in natural shorelines than in rip‐rap habitats and dispersed more consistently between sandy and gravel beaches. Tubenose goby Proterorhinus marmoratus demonstrated great plasticity in meso‐habitat use. In conclusion, Ponto‐Caspian gobies occurred rather consistently along the Hungarian section of the Danube, occupying all available habitats examined. Abundance‐habitat relations suggest plasticity in meso‐habitat use but partial segregation between species. Further studies are required at multiple spatial scales to assess how micro‐ and meso‐habitat use varies with fluctuating population densities.     

Comparative energy allocation in two sympatric, closely related gobies: the black goby Gobius niger and the grass goby Zosterisessor ophiocephalus

Seasonal energy allocation of lipid reserves into different body tissues was analysed comparatively in two sympatric, closely related gobies: the grass goby Zosterisessor ophiocephalus and the black goby Gobius niger . Lipid reserves were measured in liver, muscle and ovary and compared between the two species within a given sex and seasonal period (reproductive v . non-reproductive). Furthermore, temporal patterns of lipid reserves were investigated in the two species in relation to gonado-somatic and liver-somatic indices, as well as the relationship between size and lipid content. Results showed that the grass goby allocated more lipid reserves in reproduction while the black goby accumulated more reserves in liver and muscle, at a given size, although the temporal patterns of lipid accumulation and depletion were basically similar. Results are discussed in the light of life-history theories, taking into account both adaptation and evolutionary constraints.     

Discrimination between invasive Ponto‐Caspian gobies using a PCR‐RFLP method

Accurate identification of invaders, and especially their juveniles and eggs, is a difficult task if several morphologically similar species co‐occur. The aim of the study was to develop and test a rapid and cost‐effective procedure for identification of five species of invasive gobies occurring in the middle Danube basin, namely round goby Neogobius melanostomus, bighead goby Ponticola kessleri, monkey goby Neogobius fluviatilis, racer goby Babka gymnotrachelus and tubenose goby Proterorhinus semilunaris. First, a 708 bp fragment of the cytochrome oxidase I gene was amplified and sequenced for representative samples of these five species. Appropriate sequences of the five species available in public databases were used for in silico analysis. A digestion of the amplified fragment with the BfaI enzyme was found to be suitable for the species identification, as it showed unique restriction patterns for each species. The technique was also successfully applied for fish remains from burbot Lota lota stomachs. Thus the technique could be a useful tool in monitoring biological invasions, especially by identifying specimens that could not be determined on the basis of morphological features. The results demonstrate that the PCR‐RFLP method may in some cases be more reliable for species identification than a standard DNA sequencing.     

Phylogeography and origin of freshwater populations of tubenose gobies of genus Proterorhinus (Gobiidae: Pisces) in Ponto-Caspian Basin

Representatives of the genus Proterorhinus (tubenose gobies) from the water bodies of the Ponto-Caspian Basin were examined for sequence polymorphism of the mitochondrial DNA fragment containing the cytochrome b (Cyt-b) gene. A total of ten haplotypes were discovered, which formed two groups. Thus, the data obtained indicated the existence of only two taxonomically valid phylogenetic lineages, represented by (1) marine and brackish-water populations of the Black Sea and (2) freshwater populations of the whole Ponto-Caspian Basin, along with the brackish-water population of the Caspian Sea. Based on an analysis of the tubenose goby haplotype distribution patterns, the colonization patterns of this group of fishes (phylogeography) in the freshwater drainages of Ponto-Caspian Basin are examined. It was established that the basin of the ancient Caspian Sea was the major donor area for the formation of the freshwater and brackish-water populations. In the Ponto-Caspian Basin, two centers of the tubenose gobies speciation and distribution are distinguished. One of these centers is associated with the modern northwestern part of the Black Sea, and another one is confined to the Caspian Sea. It is suggested that the modern colonization of the Volga River by tubenose goby occurred from the Caspian Sea.     

Phylogeography of Ponto-Caspian crustaceans: a benthic-planktonic comparison

The Black, Azov, Caspian and Aral Seas, remnants of the intracontinental Paratethys basin, are home to a spectacular diversity of crustaceans. This study examines the past history of the Ponto-Caspian fauna through comparative phylogeographical studies on both benthic and planktonic taxa, based on an examination of nucleotide diversity in the mitochondrial, cytochrome c oxidase subunit 1 (COI) gene. The COI data reveal a striking example of phylogeographical concordance. All species analysed, three amphipods and three cladocerans, are characterized by two monophyletic clades corresponding to the Black and Caspian regions. However, this phylogeographical partition is, on average, four times deeper for the benthic amphipods than for the planktonic cladocerans. Based on standard molecular clocks, the Black and Caspian lineages of benthic crustaceans diverged at varied intervals from 1 to 8 million years ago. By contrast, planktonic lineages are more recent with their divergence occurring in the last million years. Levels of intraspecific polymorphisms are variable and generally lower in planktonic than benthic taxa. The mechanisms responsible for the high diversity of crustaceans in the Ponto-Caspian region are discussed on the basis of these results.     

Comparative phylogeography of Ponto-Caspian mysid crustaceans: isolation and exchange among dynamic inland sea basins

The distributions of many endemic Ponto-Caspian brackish-water taxa are subdivided among the Black, Azov and Caspian Sea basins and further among river estuaries. Of the two alternative views to explain the distributions, the relict school has claimed Tertiary fragmentation of the once contiguous range by emerging geographical and salinity barriers, whereas the immigration view has suggested recolonization of the westerly populations from the Caspian Sea after extirpation during Late Pleistocene environmental perturbations. A study of mitochondrial (COI) phylogeography of seven mysid crustacean taxa from the genera Limnomysis and Paramysis showed that both scenarios can be valid for different species. Four taxa had distinct lineages related to the major basin subdivision, but the lineage distributions and depths of divergence were not concordant. The data do not support a hypothesis of Late Miocene (10-5 Myr) vicariance; rather, range subdivisions and dispersal from and to the Caspian Sea seem to have occurred at different times throughout the Pleistocene. For example, in Paramysis lacustris each basin had an endemic clade 2-5% diverged from the others, whereas Paramysis kessleri from the southern Caspian and the western Black Sea were nearly identical. Species-specific ecological characteristics such as vagility and salinity tolerance seem to have played important roles in shaping the phylogeographic patterns. The mitochondrial data also suggested recent, human-mediated cryptic invasions of P. lacustris and Limnomysis benedeni from the Caspian to the Sea of Azov basin via the Volga-Don canal. Cryptic species-level subdivisions were recorded in populations attributed to Paramysis baeri, and possibly in P. lacustris.     

Ancient divisions, recent expansions: phylogeography and population genetics of the round goby Apollonia melanostoma

During the past two decades, the round goby Apollonia melanostoma (= Neogobius melanostomus ) has expanded its range via shipping transport and canals, extending north and west from the Ponto-Caspian region of Eurasia and to the North American Great Lakes. Exotic populations of the round goby have been very successful in the Baltic Sea and the Great Lakes regions, exerting significant ecological changes. Our study evaluates the population genetic and biogeographical structure of the round goby across its native and nonindigenous ranges, in light of geological history and its expansion pathways. We analyzed seven new nuclear microsatellite loci and mitochondrial DNA cytochrome b gene sequences from 432 individuals in 22 locations. Population structure was tested using F _ST-analogs, phylogenetic trees, clustering diagrams, Bayesian assignment tests and nested clade analyses. Results show that native populations in the Black vs. the Caspian Sea basins diverge by 1.4% and c. 350 000 years, corresponding to closure of their prior connections and supporting the taxonomic separation of the Black Sea A. m. melanostoma from the Caspian Sea A. m. affinis . Their within-basin populations diverge by ~0.4% and 100 000 years. Nonindigenous populations in the Baltic Sea and Danube and Dnieper Rivers trace to separate northern Black Sea origins, whereas the upper Volga River system houses mixed populations of A. m. melanostoma and A. m. affinis . Native populations average twice the genetic diversity of most exotic sites; however, sites in the Volga River system have high diversity due to mixing of the two taxa. Our results highlight how vicariance and anthropogenic disturbances have shaped a rapidly expanding species' genetic heritage.     

Paleoclimatic history and vicariant speciation in the "sand goby" group (Gobiidae, Teleostei)

Vicariant and climatic cycling speciation hypotheses of the 'sand gobies' belonging to the genera Pomatoschistus, Gobiusculus, Knipowitschia, and Economidichthys are tested using molecular phylogenies constructed of nuclear DNA (ITS1 locus) and mitochondrial DNA (12S and 16S fragments). These gobies are among the most abundant in the Eastern Atlantic-Mediterranean region, and play an important role in the ecosystem. Considerable ITS1 length differences, primarily due to the presence of several tandem repeats, were found between species and even within individuals. Therefore, phylogenetic analyses focused on fragments of the 12S and 16S mtDNA region that have been sequenced for 16 goby taxa. The 'sand gobies' clustered as a monophyletic group as proposed on morphological grounds. However, G. flavescens, E. pygmaeus, and K. punctatissima clustered within the Pomatoschistus species, pointing to a paraphyletic origin of these genera. Furthermore, the genetic divergence between P. minutus from the Adriatic Sea versus the Atlantic-Mediterranean region was as high as the divergence within the P. minutus complex, suggesting that P. minutus from the Adriatic Sea should be considered as a distinct species. The "star" phylogeny might suggest that these gobies evolved in a very short time period, possibly linked to the drastic alterations in the Mediterranean Sea during and immediately after the Messinian salinity crisis at the end of the Miocene. The freshwater life-style appeared monophyletic; equating its origin with the salinity crisis resulted in a molecular clock estimate of 1.4% divergence per million years. The last common ancestor probably occupied sandy bottoms and a coastal niche while several species subsequently adapted to new habitats (pelagic, freshwater or stenohaline). The origin of the shallowest clades dated back to the glacial cycling during the Pleistocene epoch.     

Isozyme criteria in the testing of phyletic relationships between species of Gobius and related eastern Atlantic-Mediterranean genera (Teleostei: Gobüdae)

Horizontal starch gel electrophoresis was employed to investigate allozyme variation at 19 isozyme loci, in nine closely-related Atlantic-Mediterranean gobiid fish species, to assess genetic relationships, and to compare this with a phyletic hypothesis based on morphological apo-morphies. The species examined were Gobius niger, G. auratus, G. cruentatus, G. paganellus, Mauligobius maderensis, Zosterisessor ophiocephalus, Thorogobius ephippiatus, Padogobius martensii and P. nigricans . Various phenetic and cladistic analyses were performed on isozyme and morphological data. The phenetic and cladistic results from morphological data, and the cladistic results from isozyme data, were largely comparable, with P. martensü and P. nigricans forming a sister group to all the other taxa, species of Cobius forming a crown group within the latter, and stem lines formed by Mauligobius, Zosterisessor and Thorogobius , respectively. The isozymic estimates of genetic distance differed from these chiefly in the relative positions of Zosterisessor and G. auratus , which appear less similar to other species of Gohius. G. auratus shows many alleles which are unique within the genus Gobius ; possible explanations for this are put forward. The position of nigricans as congeneric with Padogobius martensü is confirmed by both phenetic and cladistic analyses of isozyme data.     

Variability of GnRH secretion in two goby species with socially controlled alternative male mating tactics

Male reproductive phenotypic plasticity related to environmental-social conditions is common among teleost fish. In several species, males adopt different mating tactics depending on their size, monopolizing mates when larger, while parasitizing dominant male spawns when smaller. Males performing alternative mating tactics are often characterized by a strong dimorphism in both primary and secondary reproductive traits. According to studies on sex-changing species and on species where only one male morph is reproductively active, male alternative phenotypes are expected to vary also in gonadotropin-releasing hormone (GnRH) neurons in forebrain preoptic area (POA). Here, we compared the intra- and inter-sexual variations in number and size of GnRH neurons, along with gonads and male accessory structure investment, in two goby species, the grass goby, Zosterisessor ophiocephalus, and the black goby, Gobius niger, characterized by male alternative mating phenotypes. In both species, older and larger males defend nests, court and perform parental care, while younger and smaller ones try to sneak territorial male spawning. We found that grass goby and black goby have different patterns of GnRH expression. Grass goby presents a clear intra-sexual dimorphism in GnRH expression, related to the occurrence of alternative mating tactics, while in the black goby, only inter-sexual differences are observed. The inter- and intra-specific variability in the GnRH neurons in these two goby species is discussed in light of the differences in migratory behavior, nest type, and mating system.     

The legacy of a vanished sea: a high level of diversification within a European freshwater amphipod species complex driven by 15 My of Paratethys regression

The formation of continental Europe in the Neogene was due to the regression of the Tethys Ocean and of the Paratethys Sea. The dynamic geology of the area and repetitious transitions between marine and freshwater conditions presented opportunities for the colonization of newly emerging hydrological networks and diversification of aquatic biota. Implementing mitochondrial and nuclear markers in conjunction with a large‐scale sampling strategy, we investigated the impact of this spatiotemporal framework on the evolutionary history of a freshwater crustacean morphospecies. The Gammarus balcanicus species complex is widely distributed in the area previously occupied by the Paratethys Sea. Our results revealed its high diversification and polyphyly in relation to a number of other morphospecies. The distribution of the studied amphipod is generally characterized by very high local endemism and divergence. The Bayesian time‐calibrated reconstruction of phylogeny and geographical distribution of ancestral nodes indicates that this species complex started to diversify in the Early Miocene in the central Balkans, partially in the shallow epicontinental sea. It is possible that there were several episodes of inland water colonization by local brackish water lineages. Subsequent diversification within clades and spread to new areas could have been induced by Alpine orogeny in the Miocene/Pliocene and, finally, by Pleistocene glaciations. The present distribution of clades, in many cases, still reflects Miocene palaeogeography of the area. Our results point out that investigations of the historical aspect of cryptic diversity in other taxa may help in a general understanding of the origins of freshwater invertebrate fauna of Europe.     

Has the invasive round goby caused new links in Baltic food webs?

The Ponto-Caspian round goby (Neogobius melanostomus, Pallas 1814) most probably was established in the Gulf of Gdańsk, Baltic Sea, in the late 1980’s and has since become one of the dominant species in the region. In this study we assess the role of round gobies as prey for two important fish species in the Gulf of Gdańsk, cod (Gadus morhua) and perch (Perca fluviatilis). We compared their present diet with stomach analyses from the area prior the round goby establishment, as well as with diet analysis from Baltic regions where round gobies are absent. There were large differences in the diet between cods from the Gulf of Gdańsk 2003–2006 compared to cods in earlier studies (1977–1981) from the Southern Baltic Sea. There were also large differences in cod and perch diets from areas with and without round goby. Presently, round goby constitutes the most important prey for medium sized cods in Gulf of Gdańsk, and perch from the same area almost exclusively feed on gobiids. Stomach analysis, trophic level estimates, and stable isotope analyses all indicated that cod and perch in Gulf of Gdańsk after the round goby establishment belonged to a similar trophic level. Beside round goby, no mussel feeding fish contributed much to the diet of cod or at all to the diet of perch. Thus, it is likely that round gobies constitute a new energetic pathway from mussels to top predators. However, due to the short time elapsed after round goby establishment, we can only speculate on the species future impacts on Baltic food webs.