Molecular phylogeny of the three paedomorphic Mediterranean gobies (Perciformes: Gobiidae)

Pelagic gobies are considered of particular zoological interest because the acquisition of a pelagic lifestyle is achieved through the persistence of larval anatomical features. The Mediterranean Sea is inhabited by three goby species (Aphia minuta, Crystallogobius linearis and Pseudaphya ferreri) characterized by paedomorphic traits. Owing to the shared larval morphological features, these species have generally been considered as a monophyletic group. This study aimed at establishing the phylogenetic relationships of these paedomorphic species within the family Gobiidae to ascertain whether the pelagic lifestyle achieved through paedomorphosis is due to an event that took place in a common ancestor or not. For this purpose, we amplified by polymerase chain reaction and sequenced the mitochondrial 12S rDNA (complete sequence) and 16S rDNA (partial sequence) of 15 Mediterranean gobies. The phylogenetic analysis strongly supported the polyphyletic origin of the three paedomorphic gobies, indicating that the heterochronic change leading to the retention of larval features seems to have occurred independently in the ancestors of these species. Thus, the sharing of morphological traits can be considered homoplasious and the classification of these species in the same taxonomic group is rejected on the basis of molecular data.     

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.     

Identification of gobies (Teleostei: Perciformes: Gobiidae) from the North and Baltic Seas combining morphological analysis and DNA barcoding

Gobies are difficult to identify, as they are very similar in appearance. Here, we identified (sub)adult specimens of 12 goby species from the North Sea and the Baltic Sea by carefully analysing meristic characters, coloration patterns, papillae row patterns and morphometric measurements. The results of the morphological identifications were congruent with those obtained with the analysis of COI DNA barcodes; sequences from morphological conspecific specimens were clustered together in clades with bootstrap values ≥ 99%. Mean intra‐ and interspecific distance (uncorrected p) was 0.37 and 18.97%, respectively. A gap between the maximum intraspecific distance and the distance to the nearest neighbour was apparent in every species and ranged from 2.35 to 16.11%. The Barcode Index Number (BIN) analysis performed on the Barcode of Life Data Systems (BOLD) web platform, assigned the DNA barcodes to 12 separate clusters corresponding to sequence‐ and morphology‐based identification. In 25% of the investigated species, the BIN clusters showed taxonomic discordances, as they contained sequences assigned to more than one species. This result demonstrates the importance of accurate morphological species identification at the beginning of the barcoding pipeline. © 2014 The Linnean Society of London     

Molecular cytogenetic analysis of Haemulidae fish (Perciformes): Evidence of evolutionary conservation

Several fish species belonging to the family Haemulidae present a karyotype consisting of 48 acrocentric chromosomes (FN = 48), and apparently similar chromosomal microstructure, especially in genus Haemulon, representing a striking example of intrafamiliar chromosomal conservation. In this study, a more detailed cytogenetic analysis of the species Conodon nobilis and Pomadasys corvinaeformis was performed using C-banding, Ag-NOR, DAPI/CMA₃ staining, in situ digestion by distinct endonucleases and double-FISH to map the 18S and 5S ribosomal genes. Both species showed a similar karyotypic macrostructure with 2n = 48 acrocentric chromosomes and active ribosomal sites at interstitial position on long arms of chromosomal pair 18 and 24 in P. corvinaeformis and C. nobilis, respectively. These sites were the only CMA₃⁺/DAPI^-regions in the karyotype. Digestion with restriction enzymes revealed a low number of digestion sites in the heterochromatic segments of both species. The data indicate some degree of interspecific evolutionary diversification At the microstructural level, incorporated in a general pattern of extensive karyotypic conservatism. Thus, the interspecific reproductive isolation leading to phyletic diversification apparently occurred without the contribution of conspicuous karyotypic changes.     

Genetic architecture of the marbled goby Pomatoschistus marmoratus (Perciformes, Gobiidae) in the Mediterranean Sea

The marbled goby Pomatoschistus marmoratus, a species inhabiting coastal Mediterranean lagoons, has been studied by measuring its mitochondrial DNA variation. This analysis revealed a Mediterranean west vs east split and, subsequently, an eastern differentiation among the Libyan-Tunisian Gulf, the Adriatic Sea and the Aegean Sea. The high cohesion between the samples collected in the vast area of western Mediterranean contrasts with the genetic mosaic of the more sub-structured eastern Mediterranean. This western homogeneity can not yet be fully explained even if a human-mediated migratory flow, due to a maritime traffic, has been posited. The pattern in the eastern basin revealed a genetic architecture possibly due to the non-migratory habit of the gobid. Within this perspective, the role of the Mediterranean lagoon habitat should be related to how much it amplifies the effects of historical (e.g. past sea-level changes) and environmental (e.g. present-day hydrographic regime) processes as regards the genetic structure of the inhabiting species.     

Rapid speciation and ecological divergence in the American seven-spined gobies (Gobiidae,Gobiosomatini) inferred from a molecular phylogeny

Abstract.— The American seven-spined gobies (Gobiidae, Gobiosomatini) are highly diverse both in morphology and ecology with many endemics in the Caribbean region. We have reconstructed a molecular phylogeny of 54 Gobio-somatini taxa (65 individuals) based on a 1646-bp region that includes the mitochondrial 12S rRNA, tRNA-Val, and 16S rRNA genes. Our results support the monophyly of the seven-spined gobies and are in agreement with the existence of two major groups within the tribe, the Gobiosoma group and the Microgobius group. However, they reject the monophyly of some of the Gobiosomatini genera. We use the molecular phylogeny to study the dynamics of speciation in the Gobiosomatini by testing for departures from the constant speciation rate model. We observe a burst of speciation in the early evolutionary history of the group and a subsequent slowdown. Our results show a split among clades into coastal-estuarian, deep ocean, and tropical reef habitats. Major habitat shifts account for the early significant acceleration in lineage splitting and speciation rate and the initial divergence of the main Gobiosomatini clades. We found that subsequent diversification is triggered by behavior and niche specializations at least in the reef-associated clades. Overall, our results confirm that the diversity of Gobiosomatini has arisen during episodes of adaptive radiation, and emphasize the importance of ecology in marine speciation.     

Signature of selection on the rhodopsin gene in the marine radiation of American seven-spined gobies (Gobiidae, Gobiosomatini)

In comparison with terrestrial and freshwater ecosystems, information about speciation modes and the role of selection in marine environments is scarce. Recent studies have indicated that spectral adaptation could play an important role in the diversification of marine species flocks. Natural selection influences specific amino acids (AAs) that are involved in the spectral tuning mechanism of visual pigment genes. To study the wider occurrence and the characteristics of spectral adaptation in marine radiations, a reinterpretation of the rhodopsin (RH1) data of American seven-spined gobies (genus Elacatinus; Gobiidae; Teleostei) was carried out. Reanalysis revealed that some AAs, which are well known in the literature as spectral tuning sites, are variable in Elacatinus. Those crucial AA substitutions originated polyphyletically, indicating convergent evolution within the genus Elacatinus. Moreover, statistical tests based on the d(N)/d(S) ratio detected selection in several phylogenetic lineages and at specific AAs. Many of these AAs were previously shown to be under selection in other marine radiations. Therefore, the current phylogenetic approach provided an extended list of AAs that are probably involved in spectral tuning, and which should be validated by mutagenic experiments.     

High-frequency calling in Eneopterinae crickets (Orthoptera, Grylloidea, Eneopteridae): adaptive radiation revealed by phylogenetic analysis

The dominant frequency (Fd) of the cricket calling song commonly ranges from 2 to 8 kHz because of physical constraints due to small size and stridulum functioning. However some Eneopterinae crickets are known to call with Fds of 10–15 kHz, and one species ( Eneoptera guyanensis ) produces a modulated call with both a low and a high Fd. We studied Fd evolution with respect to phylogeny in the whole Eneopterinae subfamily to reconstruct its pattern of transformation. The phylogenetic pattern resulting from the analysis showed that Fd is relatively stable through the whole clade: the Fd ancestral state (3–7.9 kHz) has been modified only once, with the occurrence of a high Fd in the clade [ Cardiodactylus ( Lebinthus–Agnotecous )]: high Fd replaced low Fd, resulting in high-frequency calling songs. In E. guyanensis , the pattern of frequency change is different: a high Fd component has been added to the low ancestral Fd, resulting in frequency modulation. Investigation of cladogenesis rate indicated that the onset of high Fd in [ Cardiodactylus ( Lebinthus–Agnotecous )] was accompanied by a high cladogenesis rate, supporting a hypothesis of adaptive radiation for high-frequency calling (phylogeny criterion of adaptation). High frequencies are particularly problematic for long-range communication, especially for forest-living species, because of the increase of excess attenuation with frequency. The effectiveness of high-frequency calling is discussed in the clade [ Cardiodactylus ( Lebinthus–Agnotecous )] in relation to the behavioural ecology of the species.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 577–584.     

The morphology and ultrastructure of the peripheral olfactory organ in newly metamorphosed coral-dwelling gobies, Paragobiodon xanthosomus Bleeker (Gobiidae, Teleostei)

We examined the peripheral olfactory organ in newly metamorphosed coral-dwelling gobies, Paragobiodon xanthosomus (SL=5.8mm+/-0.8mm, N=15), by the aid of electron microscopy (scanning and transmission) and light microscopy. Two bilateral olfactory placodes were present in each fish. They were oval-shaped and located medio-ventrally, one in each of the olfactory chambers. Each placode had a continuous cover of cilia. The placode epithelium contained three different types of olfactory receptor neurons: ciliated, microvillous and crypt cells. The latter type was rare. Following a pelagic larval phase, P. xanthosomus settle to the reef and form an obligate association with one species of coral, Seriatopora hystrix. Their well-developed olfactory organs likely enable larvae of P. xanthosomus to detect chemical cues that assist in navigating towards and selecting appropriate coral habitat at settlement. Our findings support past studies showing that the peripheral olfactory organ develops early in coral reef fishes.     

Sounds produced by the cichlid fish Metriaclima zebra allow reliable estimation of size and provide information on individual identity

Sounds produced by male cichlids Metriaclima zebra during aggressive interactions were recorded to conduct a detailed analysis and to search for potential individual acoustic signatures. Fish from two different size groups (small and large individuals) were analysed. The two groups were significantly different for all acoustic variables considered; six of seven features demonstrated a significant interindividual variability and most of them were correlated with the size of the emitter. A cross-validated and permuted discriminant function analysis (pDFA) separated the two groups and correctly classified around 50% of the sounds to the correct individuals. Acoustic features that best distinguished among males were the instantaneous frequency of sounds and the modulation of pulse amplitude. These results suggest that acoustic signals could bear information about individual identity. The long-term stability of this signature is likely to be weak since the signature of a growing individual may change over time.     

Phylogeny and evolution of Indo-Pacific shrimp-associated gobies (Gobiiformes: Gobiidae)

Despite the ubiquity of obligate mutualisms on coral reef ecosystems, little is known about the evolution of many participating species. The shrimp gobies, known primarily from the coral reef habitats of the Indo-Pacific, are small benthic fishes that participate in a remarkable mutualism with alpheid shrimp. In this mutualism, the shrimp build and maintain a burrow that is guarded by the goby, and the shrimp and goby engage in an intricate tactile communication system. The mutualism is obligate for most shrimp gobies as participating species are highly vulnerable to predation when separated from a shrimp partner. We use phylogenetic analysis of nuclear and mitochondrial DNA sequence data to infer evolutionary relationships among shrimp gobies, and between shrimp gobies and their non-mutualistic gobiid relatives. We show that the mutualist shrimp association has arisen twice among gobies, once in a clade composed of Amblyeleotris, Ctenogobiops, and Vanderhorstia, and a second time in a clade including Cryptocentrus, Mahidolia, Tomiamichthys and Stonogobiops. We then compare the evolution of traits within each shrimp goby clade and consider their intrarelationships. We document cryptic diversity among shrimp gobies, with three distinct clades delineated among Mahidolia mysticina specimens captured at the same locality, paired with the same shrimp species. Mahidolia is placed as sister to the Cryptocentrus species Cryptocentrus cinctus; both exhibit pronounced dichromatism, occurring in both brown and yellow (xanthic) forms. We additionally clarify species identities within Amblyeleotris, confirming that widespread similar species Amblyeleotris fasciata, Amblyeleotris steinitzi and Amblyeleotris wheeleri are all distinct. We hypothesize that the flexibility of gobiid gobies and alpheid shrimp to interact with mutualist partners, as well as the apparently highly beneficial nature of mutualism between them, has contributed to the dual evolution of shrimp-association among Indo-Pacific gobies.     

Four new species of dwarfgobies (Teleostei: Gobiidae: Eviota) from the Austral,Gambier, Marquesas and Society Archipelagos,French Polynesia

Four new species of Eviota (Teleostei: Gobiidae) are described from French Polynesia. Eviota hinanoae occurs from the Gambier, Austral and Society Archipelagos and differs from its closest relatives primarily in sensory pore pattern, having cuplike male urogenital papilla, and in lacking prominent dark pigmentation on the base of the pectoral fin. Three other new species, E. dorsimaculata, E. lacrimosa and E. deminuta, are known only from the Marquesas Islands and are distinguished from congeners by differences in pigmentation, sensory pore pattern, urogenital papilla shape, the number of rays in the dorsal and anal fins, and the length of the fifth pelvic-fin ray. Eviota deminuta represents one of the smallest known species of Eviota and features reductions in several morphological characters, including the loss of all head pores. Three of the new species are included in a phylogenetic analysis based on mitochondrial cytochrome c oxidase-I, along with 21 additional species of Eviota.     

Feeding ecology of the American freshwater goby Ctenogobius shufeldti (Gobiidae, Perciformes) in a sub-tropical estuary

The feeding ecology of the American freshwater goby Ctenogobius shufeldti in a low salinity salt-marsh habitat in the Paranaguá Bay estuarine complex (Brazil) was assessed through the gut analysis of 632 individuals. The effects of a set of abiotic factors (type of sediment, salinity, temperature and estuarine reach), season and body size on dietary composition were analysed. Seasonal and size-related changes in feeding strategy, feeding intensity and trophic level were assessed. The effects of gape and body size on prey size use were also analysed. The results showed that C. shufeldti is a typical omnivorous, generalized benthic predator of low trophic levels throughout the seasons and size classes, feeding on 56 dietary items; tanaids, chlorophyte algae, ostracods, gastropods, detritus and benthic diatoms made up the bulk of its diet. The tanaid Kalliapseudes schubarti was the main prey item in both numerical and volumetric terms. The gut fullness was persistently high across the seasons. As expected for a typical generalized, opportunistic omnivorous feeder: (1) seasonal and spatial-temporal variability of abiotic factors had a significant effect on diet structure, (2) season accounted for most of the dietary variation and (3) diet composition and the size of prey consumed did not vary across the size classes.     

Abundance and spatial overlap of gobies (Gobiidae) in Lough Hyne,Ireland

Synopsis The densities of gobies (Gobiidae, Pisces) were counted by scuba divers on 13 sublittoral transects in Lough Hyne, southwest Ireland. The characters used for underwater identification are described. Densities ranged up to 6.5 m^–2 for Pomatoschistus pictus (Malm), 4.0 m^–2 for Gobiusculus flavescens (Fabricius), 0.9 m^–2 for Gobius niger L., 0.3 m^–2 for G. cruentatus Gmelin, and 0.3 m^–2 for Thorogobius ephippiatus (Lowe), while G. paganellus L., G. couchi Miller & El-Tawil and Lesueurigobius friesii Malm were rare on the transects. Gobius cruentatus and T. ephippiatus were largely confined to holes under rocks, while P. pictus and G. niger were most abundant on more open terrain. G. flavescens was hyperbenthic and largely confined to water less than 5 m depth. Significantly less P. pictus, G. flavescens and G. niger were recorded at night, and sleepingG. flavescens were observed. A working model of the spatial overlap of gobies in Lough Hyne is presented, and the possible roles of diet, predation and temporal activity patterns in interspecific competition are discussed.     

Patterns of gonad structure in hermaphroditic gobies (Teleostei Gobiidae)

Synopsis Hermaphroditism has been reported for a small number of gobiid fishes, but the extent of this sexual pattern within the family is not known. Gonad structure was examined in one or more species from twenty-one gobiid genera. No evidence of hermaphroditism was found in the species selected from 14 genera. Laboratory studies supported the conclusion of gonochorism for the examined species in four of them:Asterropteryx, Bathygobius, Gnatholepis, andPsilogobius. Currently, the absence of precursive testicular tissues associated with the ovary in females, in conjunction with no retained ovarian features in the testes of males, appear to be reliable indicators of a gonochoristic sexual pattern in gobiid fishes. Evidence for hermaphroditism was observed in seven genera:Eviota, Trimma, Fusigobius, Lophogobius, Priolepis, Gobiodon, andParagobiodon. Protogyny was experimentally confirmed inE. epiphanes, and the gonad structure in another nine of ten species ofEviota suggested either protogyny or protogynous tendencies. With the exception ofGobiodon andParagobiodon, which exhibited similar gonadal structure, ovarian and testicular structure varied considerably among the hermaphroditic genera examined, both with regard to the configuration and to the degree of development of ovarian and testicular tissues, or testicular tissue precursors. Findings of this study indicate that hermaphroditic gonad structure will prove to be a useful trait in determining evolutionary relationships within the Gobiidae.     

Isolation and characterization of microsatellites in Zosterisessor ophiocephalus (Perciformes,Gobiidae)

We isolated and characterized 10 polymorphic microsatellite loci from two partial genomic libraries of Zosterisessor ophiocephalus (Perciformes, Gobiidae), enriched for AC and for multiple motifs. Variability was tested on specimens from the Venice lagoon revealing a high degree of genetic variation. The loci will be useful for monitoring the biodiversity of the species at the genetic level and for parentage assessment purposes.     

Interspecific variation of ontogeny and skull shape among porpoises (Phocoenidae)

All extant members of Phocoenidae (porpoises) have been characterized as pedomorphic based on skeletal characters. To investigate the ontogenetic background for pedomorphosis and assess interspecific differences in ontogeny among phocoenids, samples of the six extant species were compared in terms of development of both epiphyseal and cranial suture fusion. Across all species, full maturity of the vertebral column was rare. Vertebral epiphyseal development did not progress so far in most Phocoena phocoena as in Phocoenoides dalli and Phocoena dioptrica. P. phocoena, Phocoena spinipinnis, Ph. dalli, and P. dioptrica, for which large series were available, were further compared in terms of ontogeny of cranial shape by three‐dimensional geometric morphometrics. Ph. dalli and P. dioptrica generally showed further development of cranial sutures than the other species. Postnatal skull shape development was similar for all species studied; the majority of interspecific shape differences are present at parturition. Smaller species had a higher rate of shape development relative to growth in size than Ph. dalli and P. dioptrica, but they still showed less allometric development due to less postnatal growth. Interspecific shape differences indicate phylogenetic relationships similar to that proposed based on morphology or convergent evolution of the two pelagic species, Ph. dalli and P. dioptrica, under the scenarios suggested by recent molecular studies. A shape trend coinciding with habitat preference was detected; in species with pelagic preference the position and orientation of the foramen magnum aligned the skull with the vertebral column; the rostrum showed less ventral inclination, and the facial region was larger and more concave in lateral aspect. J. Morphol., 2011. © 2010 Wiley‐Liss, Inc.     

Mitochondrial DNA variation and the evolutionary history of the Mediterranean species of Cicada L. (Hemiptera,Cicadoidea)

Cicadas are widely distributed in the Mediterranean area and are mainly distinguished by male acoustic signals, which act as specific mate recognition systems. Within the genus Cicada L. several species have diverged in their calling songs without showing external morphological differences, so acoustic recordings and genetic studies are particularly useful for systematic, biogeography and evolutionary studies. This study assesses sequence variation in closely related species of Cicada from the Mediterranean area, using domain III of the 12S rRNA mitochondrial gene in order to determine the phylogenetic relationships and the evolutionary history of this group, as well as the population structure of the two most common species, C. orni and C. barbara. Five distinct haplogroups were identified, C. orni, C. barbara, C. mordoganensis, C. cretensis and C. lodosi, each corresponding to a distinct evolutionary group. C. barbara was the most divergent species within this group, while C. orni and C. mordoganensis were the most similar. The population structure and demographic parameters of the species were not completely resolved. However, there is evidence for the separation of the C. orni Greek populations from the rest of Europe and also for demographic expansions probably related to Pleistocene climate changes. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 266–288.     

A new species of Gobius (Perciformes: Gobiidae) from the Mediterranean Sea and the redescription of Gobius bucchichi

A new species of the gobiid genus Gobius (Gobiidae, Perciformes), Gobius incognitus sp. nov. is described from the Mediterranean Sea, and its most morphologically similar species Gobius bucchichi is redescribed. The new species is distinguished from its congeners by: scales in lateral series 51–59; predorsal scales 25–35; opercle scaled in adults with 10–16 scales present; pectoral fin with ray count 18–20 and free tips on upper rays well developed and on the first ray longer than two thirds of the entire ray length; pelvic disc complete and with well‐developed anterior membrane without lateral lobes; anterior oculoscapular canal with pore α at rear of orbit; oculoscapular row x¹ not extending forwards to pore β; suborbital row d discontinuous with large gap below suborbital rows 3 and 4; eye diameter 1·08–1·32 in snout length; by pigment rows on cheek and pigmentation on pectoral‐fin base.     

Differentiation and adaptive radiation of amphibious gobies (Gobiidae: Oxudercinae) in semi-terrestrial habitats

During several surveys made in the region of the lower Fly River and delta, Papua New Guinea, nine species of oxudercine gobies (Gobiidae: Oxudercinae) were recorded: Boleophthalmus caeruleomaculatus, Oxuderces wirzi, Periophthalmodon freycineti, Periophthalmus darwini, Periophthalmus novaeguineaensis, Periophthalmus takita, Periophthalmus weberi, Scartelaos histophorus and Zappa confluentus. An exploratory multivariate analysis of their habitat conditions discriminated five guilds, differentially distributed in habitats with different quantities of environmental water and three guilds corresponding to different levels of salinity. A partial correspondence between phylogenetic and ecological categories suggested the presence of parallel adaptive radiations within different genera. In particular, the species found in the most terrestrial habitats (P. weberi) was also found in the widest range of conditions, suggesting that colonization of extreme semi-terrestrial and freshwater habitats by this species was facilitated by eurytypy. It is proposed that these findings provide insight into convergent adaptations for the vertebrate eco-evolutionary transition from sea to land.