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.     

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.     

New records of non-indigenous gobies (Neogobius spp.) in the Austrian Danube

Beginning in the early 1990s, species of the genus Neogobius rapidly expanded their native range from the Black Sea and the lower reaches of the River Danube upstream to include the middle and upper Danube. In 2002, 83 rip‐rap sites at 25 locations along the Austrian section of the River Danube were sampled by catch per unit effort (CPUE) with electrofishing to assess goby distribution. Bighead goby N. kessleri was found in all locations and at 76% of these sites, it was the third‐most abundant species (17% of CPUE). Densities of Neogobius spp. were highest in industrial harbours (IH), whereas the native species European bullhead Cottus gobio was found primarily in the main channel and tubenose goby Proterorhinus marmoratus in non‐industrial harbours. The history of records and the present distribution of Neogobius spp. suggest that IHs are the main dispersal points.     

Parasite fauna of native and non‐native populations of Neogobius melanostomus (Pallas, 1814) (Gobiidae) in the longitudinal profile of the Danube River

Parasite fauna of round goby Neogobius melanostomus (Pallas, 1814) in the Danube River was investigated in both its native range (two sites in the Bulgarian stretch of the Danube) and non‐native range of distribution (Croatian, Slovak and Austrian stretches) during 2005 and 2006. The aim was to identify possible changes in parasite communities associated with the introduction of a host into the new environment. A total of 29 metazoan parasite species were found to parasitize round goby in the Danube River; twelve of these parasite species were found in both the native and non‐native range of distribution. Introduction of a novel parasite species to the non‐native range via the round goby was not found. Eight parasite species occurred only in the native range and nine species only in the non‐native range of the round goby distribution. Losses of native parasite species in non‐native round goby populations and/or acquiring of novel parasite species in a new environment were not significant. Thirteen parasite taxa were recorded for the first time in round gobies. Three parasite taxa (Diplostomum spp., Pomphorhynchus laevis and Raphidascaris acus) were found in high prevalence and abundance at each sampling site in both the native and non‐native range. Parasite species diversity was assessed for each sampling site and season using three diversity indices (the Shannon, Simpson and Equitability indices), with the highest same‐season values found in a non‐native site in Slovakia (1.38, 0.69 and 0.60, respectively) and the lowest in a native site in Bulgaria (0.28, 0.12 and 0.14, respectively). Species diversity was higher in both non‐native round goby populations (Slovak and Austrian) compared to native Bulgarian populations. However, diversity indices values varied among almost all sampling sites.     

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.     

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 .     

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.     

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     

Occurrence of non‐native fishes in the Danube east of Vienna (Austria) and potential interactions of invasive gobiids with native fishes

Non‐native fish species pose a major threat to local fish populations and aquatic ecosystems in general. Invasive gobies are a particular focus of research, but with partly inconsistent results. While some studies reported severe detrimental impacts on native species, others have concluded less serious or neutral effects. We provide results from a large‐scale, multi‐annual fish monitoring program on the occurrence and abundance of non‐native fishes in the main stem of a free‐flowing section of the Austrian Danube. Special emphasis was placed on identifying positive or negative interactions of invasive gobies with native species. Whereas most non‐native species occurred too sporadically or were too few in number to infer a direct threat on the local fish community, invasive gobies were among the most common fishes throughout all sampling years. Co‐occurrence analyses revealed species‐ and mesohabitat type‐specific associations of gobies with native species, which were primarily positive. Notably, native predators such as asp, burbot, or perch probably benefit from the ubiquitous gobies. Two characteristic fluvial fishes revealed negative associations with invasive gobies, namely barbel (Barbus barbus) and Danube whitefin gudgeon (Romanogobio vladykovi): they appear to avoid habitats occupied by gobies. Accordingly, high abundances of round and bighead goby most likely resulted in population losses of barbel and whitefin gudgeon, respectively. Overall, our results indicate a limited negative impact of non‐native species in the sampling area. This is because only two out of 51 occurring species were found to be adversely affected by gobies, the share of co‐occurrences with native species was high, and other non‐native species were generally rare. Nevertheless, invasions are highly dynamic, and new non‐native species are likely to occur in the Austrian Danube, calling for continued monitoring and awareness.     

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.     

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.     

Comparative morphology of the sagittal otolith in three species of south Caspian gobies

Sagittal otolith shapes were investigated in order to identify three sympatric species of south Caspian gobies (Caspian goby Neogobius caspius, deepwater goby Ponticola bathybius and bighead goby Ponticola gorlap). The sagittal otoliths in P. bathybius have a rectangular shape and are thick, whereas in N. caspius they are relatively round and thin. In P. gorlap, otoliths have an elongated shape and are relatively thick. The noticeable difference among the otoliths of the three species is the presence of one anterior and one posterior projection in the otoliths of N. caspius and P. gorlap. Among shape indices, form factor (irregularity of surface area), circularity, aspect ratio and rectangularity are the foremost that indicate interspecific variability. The canonical discriminant analysis correctly classifies 94·7% of the original group cases. The overall analyses show the relevance of applying otolith shape for interspecific distinction of the three species of Caspian gobies.     

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.     

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.     

Food Resource Partitioning between Two Invasive Gobiid Species (Pisces,Gobiidae) in the Littoral Zone of the River Danube,Hungary

At present, the invasive round and bighead gobies are the most abundant benthic fish species in the near shore zone of the Middle Danube. We compared their diet seasonally in natural and artificial habitats and contrasted it with the food supply. The composition of the macroinvertebrate community was determined mainly by seasonal changes, whereas habitat type had smaller effect. Round gobies followed these changes flexibly. They consumed mainly chironomids in the spring, whilst amphipods and molluscs in the summer and autumn. Bighead gobies relied on amphipods in each season and in both habitats, and consumed fish, too, including round goby (intraguild predation). Diet overlap was determined by the morphological differences of the species allowing a varying degree of differentiation according to the seasonally changing food supply. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

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.     

The sampling efficiency of electrofishing for Neogobius species in a riprap habitat: a field experiment

Even though electrofishing is commonly used to sample Neogobius spp. and other swimbladder‐lacking benthic fishes, its efficiency is considered poor especially in habitats with abundant interstitial spaces. To determine the efficiency of electrofishing Neogobius spp. and quantitatively estimate sampling bias in a riprap (shot rock used to armor shorelines against water erosion) mesohabitat, riprap fragments were set up in a natural riverine environment. The experimental setting enabled us to collect all fish remaining in the riprap fragments after these areas had been electrofished. The sampling efficiency of electrofishing Neogobius spp. (dominated by Neogobius melanostomus) varied between 17.6 and 47.4% (mean 29.7%), while percids (possessing a well‐developed swim bladder) were collected with 74.6% efficiency. Fish size had no effect on the probability of capturing Neogobius spp. by electrofishing. Within Neogobius spp., N. melanostomus was less susceptible to electrofishing than Neogobius gymnotrachelus (23.7% and 50.1%, respectively). Decreased electrofishing efficiency in areas of rocky substrate should be considered in estimates of total abundance of Neogobius spp., especially if they are to be compared with catches of other species possessing swim bladders.     

Life-history traits of invasive bighead goby Neogobius kessleri (Günther, 1861) from the middle Danube River, with a reflection on which goby species may win the competition

Life history traits of an invasive population of bighead goby Neogobius kesslerei (Günther, 1861) from the middle Danube, including absolute and relative fecundity, egg size, number of spawning batches and size at first maturation, were examined and evaluated within an epigenetic context. Ripe bighead goby females attained 42.8–142.5 mm L _S, with absolute fecundities ranging from 669 to 5646 eggs (mean 2109 eggs), and relative fecundities of 61.6–174.0 eggs g⁻¹ body weight (mean 119.6 eggs). Egg diameters varied between 0.04 mm and 1.70 mm (mean = 0.57 mm). In the pre-spawning period there was no clear size distinction in eggs (0.12–1.45 mm; mean = 0.52 mm) in 34.1% of females; whereas in 65.9% of females, two egg size groups were distinguished: group I diameters of 0.06–0.85 mm (mean = 0.43 mm), and group II diameters of 0.55–1.70 mm (mean = 1.17 mm). Females with size-group II eggs at the beginning of the reproductive season were assumed to be ready to spawn and the others to be subsequent spawners. Bighead goby appears to be altricial compared to the round goby, although in both species a shift from highly precocial towards a less precocial life history was observed. These differences, affected by epigenetic mechanisms and resulting in alternative ontogenies, may have important implications for a species' potential success in novel environments, favouring the round goby over short time periods (several years) and bighead goby over longer periods of time (decades and longer).     

New records of endoparasitic helminths in alien invasive fishes from the Carpathian region

Six non-native invasive fish species from the Slovak part of the Tisa River basin, namely Carassius gibelio, Pseudorasbora parva, Ameiurus melas, A. nebulosus, Lepomis gibbosus, Perccottus glenii, and three gobiid species from the Danube River, namely Neogobius melanostomus, N. fluviatilis and N. kessleri, were investigated for endohelminth parasites. The expanding Asian cestode Nippotaenia mogurndae (syn. Amurotaenia perccotti) (Nippotaeniidea) has been introduced to Europe with its invasive host P. glenii. Pumpkinseed, Lepomis gibbosus, is a new definitive host of Proteocephalus percae and it is reported as the second intermediate host of the bothriocephalidean cestode Triaenophorus nodulosus in Slovakia.     

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.