Identification of a host-associated species complex using molecular and morphometric analyses,with the description of Gyrodactylus rugiensoides n. sp. (Gyrodactylidae,Monogenea)

Gyrodactylus rugiensis was originally described as a parasite occurring on the marine gobies Pomatoschistus minutus and Pomatoschistus microps. In our preliminary survey this species was also frequently found on Pomatoschistus pictus and Pomatoschistus lozanoi. Subsequent molecular analysis of the internal transcribed spacers rDNA region revealed that this parasite actually represents a complex of two apparently cryptic species, one restricted to P. microps and the other shared by P. minutus, P. lozanoi and P. pictus. Morphometric analyses were conducted on 17 features of the opisthaptoral hard parts of specimens collected from all four host species. Standard discriminant analysis showed a clear separation of both genotypes by significant differences in marginal hook and ventral bar features. Statistical classifiers (linear discriminant analysis and nearest neighbours) resulted in an estimated misclassification rate of 4.7 and 3.1%, respectively. Based on molecular, morphological and statistical analyses a new species, Gyrodactylus rugiensoides is described. This species seems to display a lower host-specificity than generally observed for Gyrodactylus species as it infects three sympatric host species.However, seasonal and host-dependent morphometric variation is shown for G. rugiensoides collected on P. pictus. Host-switching and gene flow might be important factors preventing speciation on closely related and sympatric host species. The presence of host associated species complexes in this Gyrodactylus-Pomatoschistus system is also confirmed by the presence of two host-dependent genotypes within G. micropsi found on P. minutus and P. lozanoi, and P. microps, respectively. By comparing host and parasite phylogeny, phylogenetic and ecological factors influencing host-specificity are discussed.     

Age determination in Pomatoschistus mi nut us (Pallas) and Pomatoschistus microps (Kreyer) (Pisces: Gobiidae) from the upper Tagus estuary, Portugal

The use of scale inspection as a means of age-determination in Pomatoschistus microps and Pomatoschistus minutus in the Tagus estuary is confirmed, but annulus formation pattern is different from that described for P. microps in the area of the British Isles (Miller, 1975). In Britain, two annuli are laid down on the scales each year, but in the Tagus, only one annulus is formed. One possible reason for this difference is the fact that in the Tagus the peak breeding season and the lowest water temperatures coincide. Therefore, the two causes of annulus formation coexist at the same time of year.
Population structure in both species is dominated by the 0-group, which appears earlier (April/May) than reported for other areas. Maximum age seems not to exceed 26 months in P. microps and 32 months in P. minutus .     

The food of Pomatoschistus minutus (Pisces, Gobiidae) in Belgian coastal waters, and a comparison with the food of its potential competitor P. lozanoi

Data are presented on the food and gonadosomatic index of Pomatoschistus minutus obtained from the bycatch of a commercial shrimp trawler. P. minutus has a diverse diet including zooplankton and hyperbenthos but mainly epibenthic and benthic animals. During the spawning season males consumed high numbers of Pomatoschistus eggs, presumably after aggressive take–overs of territories. The food of P. minutus is compared to the food of P. lozanoi of the same size classes obtained from the same hauls. In the absence of data on predator and prey abundance the discussion on possible interference competition between the two species remains largely speculative.     

A redescription of Pomatoschistm norvegicus (Collett, 1903) (Teleostei: Gobioidei) based on syntype material

A deeper water species of the Mediterranean-Atlantic gobiid genus Pomatoschistus, P. norvegicus (Collett, 1903), is redescribed from syntype material, including details of the lateral-line system and a summary of geographical distribution. The systematic status of P. norvegicus is briefly discussed with reference to the other nominal species, P. minutus (Pallas) and P. lozanoi (De Buen), of the 'P. minutus' complex.     

Four new species of Gyrodactylus von Nordmann, 1832 (Monogenea,Gyrodactylidae) on gobiid fishes: combined DNA and morphological analyses

Four Gyrodactylus species parasitising four closely related gobiid species in European coastal waters were studied and compared with G. arcuatus Bychowsky sensu Bychowsky & Poljansky (1953) from Gasterosteus aculatus . These were G. gondae n. sp. from Pomatoschistus minutus and P. lozanoi , G. flavescensis n. sp. from Gobiusculus flavescens , G. arcuatoides n. sp. from P. minutus and G. branchialis n. sp. from P. microps. Combined molecular and morphological analyses, as well as morphometric and statistical methods, were used. The ssrRNA V4 region and the complete ITS rDNA region were sequenced. Genetically the four new species are clearly distinct from G. arcuatus . From a morphological point of view, the haptoral hard parts of G. gondae n. sp., G. flavescensis n. sp. and G. arcuatoides n. sp. are related to those of G. arcuatus, while these parts of G. branchialis n. sp. are different, but related to those of G. quadratidigitus Longshaw, Pursglove & Shinn, 2003. For the latter two species, a new species group is formed. The V4 and ITS sequence analyses, however, indicate a close relationship between G. branchialis and the three G. arcuatus-like species.     

First report of Gyrodactylus spp. (Platyhelminthes: Monogenea) in the western Mediterranean Sea: molecular and morphological descriptions

The Gyrodactylus spp. fauna on species of gobies, Pomatoschistus, Gobiusculus, and Knipowitschia (Gobiidae: Teleostei), from the western Mediterranean and Adriatic seas is strikingly similar to that found in the Baltic Sea and eastern Atlantic Ocean, both in morphology and in internal transcribed spacer (ITS) rRNA. The fauna consisted of Gyrodactylus branchialis, G. ostendicus, G. gondae, G. rugiensis, G. rugiensoides, and G. arcuatus. No new species have been found. A morphometric comparison between G. branchialis from the Mediterranean and Adriatic seas and its type locality in Ostend (Belgium) showed significant differences in ventral bar and marginal hook features. The morphometric variation was lower in G. rugiensis, whereas no significant differences were found in G. ostendicus. Gyrodactylus branchialis and G. ostendicus collected on P. microps were slightly different in the ITS rDNA (-0.6%) compared with specimens on the closely related P. marmoratus, probably reflecting ongoing speciation. A hybrid zone was identified in the Vaccarès lagoon complex (France) where both host species are sympatric. There was no clear geographic or host-related pattern in the variation found in the ITS2 rDNA in G. arcuatus sampled from P. microps, G. flavescens, Pungitius pungitius, K. panizzae, and its original host Gasterosteus aculeatus (2 polymorphic sites). Of all studied species, only G. arcuatus, G. rugiensis, and G. rugiensoides showed minor intraspecific variation in the ITS rDNA. Hence, the physical separation by a shoreline of more than 10,000 km is hardly reflected in the parasite ITS rDNA.     

Occurrence of vertebral deformities in Gobiidae (Pisces) from the Tagus estuary

In four species of gobiid fish caught in Tagus estuary during 1996, the incidence of skeletal deformities ranged from 2 to 5%. Differences between sampling stations or between the two most abundant species Pomatoschistus minutus and Gobius niger were statistically not significant (p > 0.05). The type of deformity depended upon species. P. minutus and P. microps showed bone nodules at the inferior side of the abdominal vertebrae, whilst G. niger and G. paganellus had shortened caudal vertebrae (achondroplasia) and deformed ural plates. The possible role of pollution or other conditions in the Tagus estuary is discussed.     

Fish communities along an oxygen-poor salinity gradient (Zeeschelde Estuary, Belgium)

Fish living in the tidal fresh and brackish water reaches of the Zeeschelde Estuary were studied in samples collected from the cooling-water inlets of three power stations. Between July 1994 and June 1995, 42 different species were recorded including 26 marine migrants, 14 freshwater species and two diadromous species. Species number as well as fish abundance were correlated significantly with salinity and oxygen concentration. The community structure was analysed with a correlation biplot based on principal component analysis of the root-root transformed fish abundance. Four separated communities could be distinguished since the first principal component expressed a salinity and the second a temperature gradient. During summer and fall Pomatoschistus microps, P. minutus, P. lozanoi and Syngnathus rostellatus were most abundant in the brackish-water reach. At this time, freshwater species seemed to avoid the freshwater area. During winter, Sprattus sprattus, Clupea harengus and Dicentrarchus labrax were the dominant species of the brackish-water zone while the freshwater reaches were dominated by Gasterosteus aculeatus . Migrating fish such as Pleuronectes flesus, Lampetra fluviatilis, Anguilla anguilla and Osmerus eperlanus were restricted to the brackish environment.     

Small-scale clinal variation, genetic diversity and environmental heterogeneity in the marine gobies Pomatoschistus minutus and P. lozanoi (Gobiidae, Teleostei)

Genetic variation was assayed at 14 allozyme loci in estuarine, coastal and offshore samples of lozano's goby, Pomatoschistus lozanoi and the sand goby, P. minutus. Samples were taken from locations on the Belgian Continental Shelf and in the Schelde estuary with a range of environmental heterogeneity. We evaluate whether any differences in (1) the degree of genetic variation and (2) allele frequencies at the various loci exist within samples occurring in various habitats on the BCS and in the Schelde estuary. No significant differences in levels of genetic diversity were recorded between estuarine, coastal and offshore samples in either species. A temporally stable clinal gradient in allele frequencies at the two-allele locus GPI-A(*) was observed in P. lozanoi, differentiating the samples in an estuarine, coastal and offshore group. We suggest that these differences might be maintained by balancing selection at locus GPI-A(*).     

Dynamics of Kudoa camarguensis (myxosporean) infection in two gobiid species,Pomatoschistus microps and P. minutus (Teleostei: Pisces), in the Rhĵne River delta,France

The occurrence of the myxosporean parasite Kudoa camarguensis was surveyed monthly during 1997 in a brackish-water lagoon of the Rh?ne River delta (France). K. camarguensis was found on its typical host, Pomatoschistus microps, and on an additional host, P. minutus. Prevalence and mean abundance were higher in the typical host than in the additional host due to differences in the temporal occupancy of the lagoon by the 2 species. The temporal occurrence of this myxosporean parasite is discussed in relation to the migratory habits of P. minutus and the sedentary habits of P. microps.     

Prespawning sound production in mediterranean sand-gobies

The emission of breeding vocalizations among sand-gobies Pomatoschistus marmoratus , P. canestrini and Knipowitschia panizzae are described. The sounds of these gobies are pulse trains emitted by the male during mating interactions in the nest (prespawning sounds).     

Parasites as indicator of fish behaviour

In the present study two species of gobies, Pomatoschistus microps and P. minutus from southwest Wales were studied and appearance of some unusual parasite of gobies, were used as an indicator for fish behaviour. The occurrence of pleurocercoids of Ligula intestinalis in the body cavity of these fish and Proteocephalus sp., excysted metacercariae and adult of Timoniella spp., excysted metacercariae of Bucephalus baeri , the excysted metacercariae of Podocotyle atomon and progenetic adult of Bucephaloides gracilescens in their gut indicate host anadromous, cannibalistic and scavenging behaviour.     

Sexually selected nest-building--Pomatoschistus minutus males build smaller nest-openings in the presence of sneaker males

Both natural selection and sexual selection may act on nest-building. We tested experimentally how different regimes of egg-predation and male-male competition influence nest-building before mating, using the marine fish sand goby, Pomatoschistus minutus. Males with sneaker males present built the smallest nest-openings, smaller than males held alone or with Pomatoschistus microps males (which may predate eggs and compete over nest-sites but not compete over fertilizations). Males with visual access to other nest-building males tended also to build smaller openings than males held alone or with P. microps. Males with egg-predators present built nests with openings not differing significantly from any other treatment. Our results indicate that the small nest-openings found in the sneaker male treatment are sexually selected through protection against sneaking or by female choice. Across treatments, time span before a male started to build his nest also explained variation in nest-opening width; males starting late built larger nest-openings.     

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.     

Current and historic gene flow of the sand goby Pomatoschistus minutus on the European Continental Shelf and in the Mediterranean Sea

Phylogeographical patterns of the sand goby Pomatoschistus minutus (Gobiidae, Teleostei) were studied by means of sequence and single-stranded conformational polymorphism analysis of a 283-bp fragment of the cytochrome b locus of the mtDNA. A total of 228 individuals sampled at 13 sites throughout the species's distributional range revealed a moderate level of diversity and a low but significant level of overall genetic differentiation at all but one site. The goby sample from the Adriatic Sea differed in sequence by approximately 10% from the Atlantic P. minutus and is thought to belong to a cryptic species of the genus Pomatoschistus . Limited genetic differentiation with a weak pattern of isolation-by-distance was recorded throughout the distributional range of the typical P. minutus . Phylogeographical analysis suggested a contiguous range expansion in the Atlantic and Baltic basins during the Eemian and evidence for a glacial refugium in the southern North Sea during the Weichselian. In P. minutus from the western Mediterranean Sea a high number of endemic haplotypes as well as the most common Atlantic haplotype were recorded in appreciable frequencies. This might be explained by secondary contact between different mitochondrial lineages, which evolved in allopatry.   © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 561–576.     

The endurance of endemism: the Mediterranean freshwater gobies and their prospects for survival

Gobies (Teleostei: Gobiidae), spawning in fresh waters of the Mediterranean catchment area, are listed, and their diagnostic features and relationships summarized. The species, by zoogeographic category, include (i) Mediterranean: Padogobius martensii (Günther, 1861) and P. nigricans (Canestrini, 1867) from the northern Adriatic and west-central Italy, (ii) West Balkanian: Economidichthys pygmaeus (Holly, 1929) and E. trichonis Economidis & Miller, 1990, and (iii) Ponto-Caspian: Prorerorhinus marmoratus (Pallas, 1811) and Knipowitschia caucasica (Kawrajsky, in Berg, 1916), from the northern Aegean area but otherwise widespread in the Ponto-Caspian region, and the endemic Mediterranean K. punctatissima (Canestrini, 1864) (north-eastern Italy) and K. thessala (Vinciguerra, 1921) (Thessaly). Scenarios for differentiation following the late Miocene Messinian salinity crisis are outlined. The vulnerability of these species is discussed, and their present status reviewed. Populations no longer exist at the precise type localities of E. pygmaeus (Lefkas) and K. thessala (Hasabali spring, Pinios river system). Special features of scientific and cultural interest are noted, and the biological capability for species recovery is assessed.     

Vacuolated prolactin cells in the pituitary gland of several marine teleosts

A morphometric study of prolactin cell ultrastructure in the pituitary gland of the Corkwing wrasse, Crenilabrus melops L., showed that cytoplasmic vacuoles, which accounted for 25% of the cell volume, were associated with signs of decreased secretory activity. The Golgi apparatus, mitochondria and rough endoplasmic reticulum, all contributed relatively little to the total cell volume, and there was no sign of secretory-granule release by exocytosis. All vacuoles were intracellular and membrane-bound, and probably derived from rough endoplasmic reticulum, Golgi apparatus and the nuclear envelope. It is thought that smaller vacuoles coalesce to form larger ones. The secretory granules were small and sparse, and this could account for the chromophobia of prolactin cells in light-microscopy preparations. Similar vacuoles were reported in the prolactin cells of the gobiid fish, Chaparrudo flavescens, Pomatoschistus pictus, Pomatoschistus minutus and Pomatoschistus microps . The vacuoles in Chuparrudo were of similar ultrastructure to those in Crenilabrus .     

Systematics of freshwater gobies from Greece (Teleostei: Gobiidae)

Five species of gobiid fish are recognized in the fresh waters of Greece. Most distinctive is the south-western Hellenic (West Balkanian component) endemic genus Economidichthys Bianco, Bullock, Miller & Roubal, 1987, with the type-species E. pygmaeus (Holly, 1929) from Lefkas, Epirus and Lake Trichonis, and E. trichonis sp. nov., from Lake Trichonis, the latter probably the smallest freshwater European teleost at maturity (female, 18·0 mm) and maximum size (male, 30·0 mm). In tributaries of the northern Aegean basin (north-eastern Hellenic freshwater fauna), two species of the Ponto-Caspian genus Knipowitschia can be identified, the widespread Kn. caucasica (Kawrajsky in Berg, 1916) (coastal plain of Macedonia and Thrace) and the endemic Kn. ihessala (Vinciguerra, 1921) (Pinios river system, Thessaly). The larger Ponto-Caspian tubenose goby, Prolerorhinus marmoratus (Pallas, 1811), occurs in the Evros (= Maritsa) (Thrace) and Strymon (Macedonia) systems. Only the genera Economidichthys and Knipowitschia are considered in this paper, being redefined in terms of osteology and head lateral-line system, and the species described in detail, with special reference to meristic variation and squamation, and, in Economidichthys , the unique perianal organ of this genus. The relationships of Economidichthys and Kn. thessala are discussed and possible scenarios for their evolution are suggested in the light of Miocene to Recent events.     

Gyrodactylus longidactylus n. sp., a monogenean from Pomatoschistus lozanoi (de Buen) (Gobiidae) from the North Sea

Gyrodactylus longidactylus n. sp., a gyrodactylid monogenean parasitising the gills of Pomatoschistus lozanoi (de Buen, 1923) in the North Sea is described. It is a species with rather small anchors which are only connected to a ventral bar which lacks a ventral bar membrane. The marginal hooks have long handles, which are always longer than the total length of the anchor. The pharynx has long pharyngeal processes. Measurements, drawings of the opisthaptoral hard parts, penis and pharynx, and SEM micrographs of this monogenean are presented. G. longidactylus n. sp. is the first monogenean species described from P. lozanoi.     

Twenty years of invasion: a review of round goby Neogobius melanostomus biology, spread and ecological implications

The round goby Neogobius melanostomus is one of the most wide-ranging invasive fish on earth, with substantial introduced populations within the Laurentian Great Lakes watershed, the Baltic Sea and several major European rivers. Rapid expansion and deleterious ecosystem effects have motivated extensive research on this species; here this research is synthesized. Maps of the global distribution are provided and the invasion history of N. melanostomus, which spread more rapidly at first in North America, but has undergone substantial expansion over the past decade in the Baltic Sea, is summarized. Meta-analyses comparing their size at age, diet, competitors and predators in North American and European ecosystems are provided. Size at age is region specific, with saline habitats typically supporting larger and faster growing individuals than fresh water. Neogobius melanostomus prey differs substantially between regions, demonstrating a capacity to adapt to locally abundant food sources. Neogobius melanostomus comprise at least 50% of the diet of eight taxa in at least one site or life stage; in total, 16 predator taxa are documented from the Laurentian Great Lakes v. five from Eurasia. Invasive N. melanostomus are the only common forage fish to heavily exploit mussels in the Laurentian Great Lakes and the Baltic Sea, facilitating the transfer of energy from mussels to higher trophic levels in both systems. Neogobius melanostomus morphology, life history, reproduction, habitat preferences, environmental tolerances, parasites, environmental effects, sampling strategies and management are also discussed. Neogobius melanostomus inhabit a wide range of temperate freshwater and brackish-water ecosystems and will probably continue to spread via ballast water, accidental bait release and natural dispersal worldwide. Climate change will probably enhance N. melanostomus expansion by elevating water temperatures closer to its energetic optimum of 26° C. Future research needs are presented; most pressing are evaluating the economic effects of N. melanostomus invasion, determining long-term population level effects of egg predation on game-fish recruitment and comparing several variables (density, ecological effects morphology and life history) among invaded ecosystems. This review provides a central reference as researchers continue studying N. melanostomus, often as examples for advancing basic ecology and invasion biology.