Distribution,abundance and condition of invasive Ponto‐Caspian gobies Ponticola kessleri (Günther, 1861), Neogobius fluviatilis (Pallas, 1814), and Neogobius melanostomus (Pallas, 1814) in the Sava River basin,Croatia

Distributions, population densities, invasive potentials and adverse impacts of invasive Ponto‐Caspian gobies on native fauna in the Sava River basin remain unknown, whereby 23 locations in the Sava basin were sampled during a three‐year period (2011–2013). Among the five Ponto‐Caspian goby species previously reported in Croatia, only Neogobius fluviatilis (291), Neogobius melanostomus (177) and Ponticola kessleri (21) specimens were collected. Proterorhinus semilunaris and Babka gymnotrachelus were not found. N. fluviatilis was dominant among the invasive gobies at the majority of locations and almost omnipresent (apart from the upper Sava reaches) in the Sava basin. N. melanostomus was found only in the navigable Sava reaches, but its distribution range still seems to be expanding. The status of P. kessleri invasion remains unclear, as it was only found in 2011 at three locations in the lower Sava reaches. No significant (P > 0.05) increase in average catch‐per‐unit‐effort (CPUE) values was observed during the three‐year period. There is some evidence that a decline in Gobio gobio populations might be expected in response to increasing N. fluviatilis population densities. The findings suggest that further range expansions by the Ponto‐Caspian gobies can be expected in Croatia and the implementation of measures to limit their spreading should be considered.     

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.     

Migration and isolation during the turbulent Ponto‐Caspian Pleistocene create high diversity in the crustacean Paramysis lacustris

The Ponto‐Caspian brackish‐water fauna inhabits estuaries and rivers of the Black, Azov and Caspian seas and is fragmented by higher salinity waters and a major interbasin watershed. The fauna is known for the high levels of endemism, complex zoogeographic histories, and as a recent source of successful invasive species. It remains debated whether the Black and Azov Sea brackish‐water populations survived unfavourable Pleistocene conditions in multiple separate refugia or whether the two seas were (repeatedly) recolonized from the Caspian. Using microsatellite and mtDNA markers, we demonstrate deep among‐ and within‐basin subdivisions in a widespread Ponto‐Caspian mysid crustacean Paramysis lacustris. Five genetic clusters were identified, but their relationships did not reflect the geography of the region. The Azov cluster was the most distinct (4–5% COI divergence), despite its geographic position in the corridor between Black and Caspian seas, and may represent a new species. In the northern Black Sea area, the Dnieper cluster was closer to the Caspian cluster than to the neighbouring Danube–Dniester–Bug populations, suggesting separate colonizations of the Black Sea. Overall, the data implied a predominant gene flow from the east to the Black Sea and highlight the importance of Caspian Sea transgressions in facilitating dispersal. Yet, the presence of distinct lineages in the Black Sea points to the persistence of isolated refugial populations that have gained diagnostic differences under presumably high mutation rates and large population sizes. The unfavourable Pleistocene periods in the Black Sea therefore appear to have promoted diversification of the brackish‐water lineages, rather than extirpated them.     

Phylogeography and taxonomic status of trout and salmon from the Ponto‐Caspian drainages,with inferences on European Brown Trout evolution and taxonomy

Current taxonomy of western Eurasian trout leaves a number of questions open; it is not clear to what extent some species are distinct genetically and morphologically. The purpose of this paper was to explore phylogeography and species boundaries in freshwater and anadromous trout from the drainages of the Black and the Caspian Seas (Ponto‐Caspian). We studied morphology and mitochondrial phylogeny, combining samples from the western Caucasus within the potential range of five nominal species of trout that are thought to inhabit this region, and using the sequences available from GenBank. Our results suggest that the genetic diversity of trout in the Ponto‐Caspian region is best explained with the fragmentation of catchments. (1) All trout species from Ponto‐Caspian belong to the same mitochondrial clade, separated from the other trout since the Pleistocene; (2) the southeastern Black Sea area is the most likely place of diversification of this clade, which is closely related to the clades from Anatolia; (3) The species from the Black Sea and the Caspian Sea drainages are monophyletic; (4) except for the basal lineage of the Ponto‐Caspian clade, Salmo rizeensis, all the lineages produce anadromous forms; (5) genetic diversification within the Ponto‐Caspian clade is related to Pleistocene glacial waves; (6) the described morphological differences between the species are not fully diagnostic, and some earlier described differences depend on body size; the differences between freshwater and marine forms exceed those between the different lineages. We suggest a conservative taxonomic approach, using the names S. rizeensis and Salmo labrax for trout from the Black Sea basin and Salmo caspius and Salmo ciscaucasicus for the fish from the Caspian basin.     

Relative abundance of Ponto‐Caspian gobiids in the lower Vistula River (Poland) 3‐ to 4 years after first appearance

In the years 2000–2001, specimens of two Ponto‐Caspian gobiids (racer goby Neogobius gymnotrachelus and monkey goby Apollonia fluviatilis) were recorded for the first time in the lower Vistula River, which is a part of the central inland corridor used by Ponto‐Caspian species to migrate in Europe. In 2004, seine net surveys were made in the river course to examine relative abundance of the non‐indigenous gobiids in the local small‐fish community. Altogether 4420 specimens with total lengths ranging from 10 to 340 mm (mostly <100 mm) belonging to 18 species were recorded in 70 hauls. Monkey goby was one of the subdominant species (18.1% of total number of fish captured; 64.3% frequency of occurrence in hauls) together with bleak Alburnus alburnus (21.0, 54.3), roach Rutilus rutilus (18.2, 67.1) and three‐spined stickleback Gasterosteus aculaetus (17.3, 37.1). Racer goby was less abundant but quite common (2.5, 32.9), similar to common bream/white bream Abramis brama/Blicca bjoerkna (6.4, 47.1), Eurasian perch Perca fluviatilis (5.7, 57.1), dace Leuciscus leuciscus (4.2, 40.0) and European bitterling Rhodeus sericeus amarus (3.4, 27.1). The study shows that Ponto‐Caspian gobiids dispersed successfully in the lower Vistula, becoming species common to its nearshore zone 3‐ to 4 years after their first appearance.     

Diet partitioning in summer of two syntopic neogobiids from two different habitats of the lower Vistula River, Poland

Monkey goby Neogobius fluviatilis and racer goby N. gymnotrachelus, both native to the Ponto‐Caspian region, were discovered in Polish rivers during the mid‐1990s and currently they are invading the River Vistula. Preliminary studies were carried out on the diet of both species at two sites in the lower River Vistula, one lotic (main channel of the river downstream of the W?oc?awek river reservoir) and the other lentic (left bank, middle section of the W?oc?awek Reservoir). Significant differences were noted in diet between the two species and within species according to habitat, in particular the types of chironomid taxa taken. Over a single diel cycle, monkey goby were captured between 07:00 and 19:00 hours only, feeding more intensively on chironomid larvae than the racer goby, which was captured at most times of day. The wide diet plasticity of these non‐native fish species favours their expansion.     

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.     

Microsatellite loci for Ponto-Caspian gobies: markers for assessing exotic invasions

We developed and tested eight polymorphic microsatellite loci for Ponto‐Caspian ‘neogobiin’ gobies, many of which are invasive in Eurasia and North America, whose study will aid understanding of the population genetics underlying their success. We tested samples from one to two locations from 12 taxa in the recently revised genera Babka, Benthophilus, Mesogobius, Neogobius = Apollonia, Ponticola and Proterorhinus; including the bighead, Caspian, knout, monkey, racer, round, tadpole and tubenose gobies; and taxa from introduced vs. native populations, those diverging between fresh and marine waters, and those differentiated between the Black and Caspian Seas. Populations conformed to Hardy–Weinberg equilibrium expectations, averaging five to 15 alleles per locus and 0.11 to 0.67 mean heterozygosity. Allelic variation significantly differentiated among all taxa and populations.     

Some remarks on parasitic infections of the invasive Neogobius spp. (Pisces) in the Hungarian reaches of the Danube River, with a description of Goussia szekelyi sp. n. (Apicomplexa: Eimeriidae)

During a survey of the parasite fauna Neogobius spp., which only recently invaded Hungarian reaches of the Danube River upstream from Budapest, it was found that Neogobius melanostomus and N. fluviatilis had become common species. Neogobius kessleri was less frequent and the previously widespread Proterorhinus marmoratus was drastically reduced. The parasite fauna of the newcomer fish species was characterized by ubiquitous parasites of a wide host range, such as the trematode Nicolla skrjabini, the metacercariae of Metagonimus yokogawai and Apatemon cobitidis, the larval stages of the acanthocephalan Pomphorhynchus laevis and the nematode Rhaphidascaris acus, the glochidia of an Anodonta sp. (Mollusca) and the ciliophoran Ichthyophthirius multifiliis. The specific parasites of Neogobius spp. were represented by three intestinal coccidia: Eimeria daviesae, Goussia kessleri, and a new species described here as Goussia szekelyi sp. n. This survey showed that the rapid invasion of Ponto‐Caspian gobies in the northern Danube caused important changes in the fish fauna composition of the Danube River and enriched the parasite fauna of Hungarian fishes.     

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.     

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.     

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 .     

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.     

Invasion phylogeography of the Ponto-Caspian crustacean Limnomysis benedeni dispersing across Europe

Aim  Limnomysis benedeni Czerniavsky, 1882 is a mysid crustacean native to the Ponto-Caspian (Black and Caspian Sea) rivers and estuaries, and has recently spread across Europe through intentional and unintentional introductions. We explored the structuring of genetic variation in native and non-native populations with an aim to trace the sources of the invasions, and to infer whether the spread has occurred through a single or multiple invasion waves.
Location  Native estuaries in the Ponto-Caspian basin (Volga, Don, Dnieper, Dniester, Danube) and the recently colonized range along the Danube–Rhine river systems and Lithuania.
Methods  A fragment of the mitochondrial COI gene was sequenced to assess genetic affinities and diversity in native and recently established populations.
Results  The genetic diversity in the native regions is organized into several strongly diverged haplotype groups or lineages, partly allopatric, partly sympatric. All these lineages have also spread beyond the native range. Even the recent rapid dispersal across Europe along the Danube–Rhine system towards the North Sea basin involved several lineages from the Danube delta sector. The structuring of genetic diversity among invaded sites suggests multiple invasion events to the Danube–Rhine drainage. This contrasts with data from some other Ponto-Caspian species, where a single haplotype seems to have occupied most invaded areas. There is no evidence that intentionally stocked reservoirs in the Baltic Sea basin would have contributed to further unintentional spread of L. benedeni.
Main conclusions  Limnomysis benedeni is spreading across Europe using the southern invasion corridor. The invasion most likely involved several waves from differentiated sources in the native Danube delta area.     

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     

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.     

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.     

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.     

Feeding preferences of an invasive Ponto‐Caspian goby for native and non‐native gammarid prey

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