Article

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

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Abstract

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 x(1) 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.

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... Here, we used the anemone goby Gobius incognitus (Kovačić and Š anda, 2016) living off natural CO 2 vents to investigate potential behavioural alteration caused by OA. This small territorial benthic fish represents an ideal candidate for in situ OA experiments due to its very limited home range, that ensures the spatial segregation between groups of individuals living in adjacent areas naturally characterized by different pCO 2 conditions. ...
... This small territorial benthic fish represents an ideal candidate for in situ OA experiments due to its very limited home range, that ensures the spatial segregation between groups of individuals living in adjacent areas naturally characterized by different pCO 2 conditions. Moreover, the strict association between G. incognitus and the sea anemone Anemonia viridis (Forsskål, 1775), used as shelter in case of threats (Kovačić and Š anda, 2016;Nagelkerken et al., 2015;Tiralongo et al., 2020), allows to use the sheltering time as a clearly measurable variable linked to the anti-predator behavior. In addition to sheltering, the anemone goby can show alternative antipredator strategies like an escaping ("fleeing") and 'freezing' behaviors as observed in similar species (Larson and McCormick, 2005;McCormick and Larson, 2007). ...
... Gobius incognitus (Kovačić and Š anda, 2016) (Fig. 1a) is widespread in Mediterranean shallow coastal waters, mostly preferring rocky (gravel, cobbles, boulders and bedrock) and sand substrata, specifically when mixed with rocky bottoms (Kovačić and Š anda, 2016;Tiralongo et al., 2020). Its presence from the western to the eastern part of the Mediterranean basin has been often confused with the congeneric Gobius bucchichi (Steindachner, 1870), which is actually only reported from the eastern part of the Adriatic Sea and from the northern Ionian Sea (Albania) (Kovačić and Š anda, 2016). ...
Article
An in situ reciprocal transplant experiment was carried around a volcanic CO2 vent to evaluate the anti-predator responses of an anemone goby species exposed to ambient (∼380 μatm) and high (∼850 μatm) CO2 sites. Overall, the anemone gobies displayed largely unaffected behaviors under high-CO2 conditions suggesting an adaptive potential of Gobius incognitus to ocean acidification (OA) conditions. This is also supported by its 3-fold higher density recorded in the field under high CO2. However, while fish exposed to ambient conditions showed an expected reduction in the swimming activity in the proximity of the predator between the pre- and post-exposure period, no such changes were detected in any of the other treatments where fish experienced acute and long-term high CO2. This may suggest an OA effect on the goby antipredator strategy. Our findings contribute to the ongoing debate over the need for realistic predictions of the impacts of expected increased CO2 concentration on fish, providing evidence from a natural high CO2 system.
... The genus Gobius currently comprises 28 recognized species, which are mainly distributed in the NE Atlantic, the Mediterranean and the Black Sea (Miller, 1986;Patzner et al., 2011;Froese & Pauly, 2019). Notably, two of these species are recent discoveries (Iglésias et al., 2016;Kovačić & Šanda, 2016). Fourteen species are traditionally attributed to the genus Pomatoschistus, including three that were only recognized in the last decade (Miller & Šanda, 2008;Engin & Innal, 2017;Engin & Seyhan, 2017;Froese & Pauly, 2019); however, the genus is not monophyletic according to recent phylogenetic studies (Thacker et al., 2019). ...
... Species of Pomatoschistus mostly occur in coastal habitats along the NE Atlantic, the Baltic Sea, and in some areas of the Mediterranean and the Black Sea (Miller, 1986;Patzner et al., 2011). Identification of the individual species in both genera, Gobius and Pomatoschistus, relies on coloration patterns, the arrangement of the head sensory organs (papillae), and combinations of various other characteristics such as squamation, morphometric and meristic traits; species determination often turns out to be very difficult, especially between closely related species (Miller, 1974(Miller, , 1986(Miller, , 2004Kovačić & Šanda, 2016;Engin & Seyhan, 2017). ...
... But the delimitation of a fossil goby species is often problematic because information on the range of variation in skeletal and otolith traits within a respective species and between related taxa is restricted. Furthermore, the characters of the skeleton may vary little between extant goby species (Miller, 1986;Kovačić & Šanda, 2016). This, together with the loss of delicate characters in fossil skeletal material due to taphonomic processes, can lead to underestimation of the number of fossil species (see Gaudant & Quayle, 1988vs. ...
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Gobies (Gobiidae + Oxudercidae) are among the largest groups of extant marine fishes. Fossils of gobies are abundant since the Miocene, and many species have been reported so far. However, delimitation of fossil goby species is challenging because molecular markers and diagnostic traits such as the disposition of sensory head papillae are lost. This study provides, for the first time, an actualistic framework for the identification of fossil goby species. We focus on characters that can in principle be recognized in fossils, and evaluate their ability to discriminate between extant goby species based on statistical analyses. Using 14 extant species of Gobius and seven species of Pomatoschistus , we conducted otolith morphometry, elliptic Fourier shape analysis of otoliths using the package ‘Momocs’, conventional fish morphometry, and meristic counts. In addition, the otoliths of all species are depicted based on SEM images and briefly described. Otolith Fourier shape analysis proved to be most efficient in discrimination of species within both genera, Gobius and Pomatoschistus . Several characters used in the other approaches also worked well, but the results were variable, and the relative taxonomic significance of particular variables tended to change depending on the species under consideration. We propose otolith shape analysis as a powerful tool to explore ancient goby species diversity when samples with abundant fossil otoliths are present. Overall, the herein presented data will greatly facilitate delimitation of fossil goby species in future studies, and will consequently shed new light on the evolution of goby diversity and biogeography through time.
... This is also valid for the Mediterranean Sea, where, with more than 70 species, Gobiidae is the most specious fish family (PATZNER, 2019). However, data about biology and ecology for most of the species are scarce and dated, and only in relatively recent years some studies have deepened some ecological aspects of some species (HERLER & the Mediterranean Sea, having been observed from the western to the eastern part of the basin; while, the presence of the redescribed G. bucchichi is actually only reported from the eastern part of the Adriatic Sea and from the northern Ionian Sea (Albania) (KOVAČIĆ & ŠANDA, 2016). In consideration of this, most of the previous studies on G. bucchichi probably concern G. incognitus. ...
... Gobius bucchichi was observed associated with the sea anemone Anemonia viridis (Forsskål, 1775), and seeks protection in the tentacles of the sea anemone when it feels threatened (ABEL, 1960). However, on the basis of the study of KOVAČIĆ & ŠANDA (2016), this behaviour could instead concern G. incognitus. This is also valid for the study of PATZNER (1999b) that described the association between juveniles of G. bucchichi and sea urchins, Paracentrotus lividus (Lamarck, 1816) in particular. ...
... Rows of dark spots, forming broken longitudinal lines, are present on body; midlateral longitudinal row with 8-9 larger dark spots, extending from the pectoral axilla to the caudal fin base. Unlike G. bucchichi, in which on the cheeks there are only two rows of elongated dots, without other dots between them at the middle of the cheek itself, in G. incognitus the dots on cheeks are irregularly scattered or, if arranged in rows, additional dots or a third row of dots is present across the middle of the cheek, between the lower row, starting anteriorly at the angle of mouth, and the upper row, that touches ventral margin of the eye (KOVAČIĆ & ŠANDA, 2016). Some data concerning biology and ecology of G. bucchichi were reported from the Gulf of Cadiz (approximatively the westernmost range of the species) and western Mediterranean Sea (SASAL et al., 1996a;SASAL et al., 1996b;BOUCHEREAU & GUELORGET, 1999;PATZNER, 1999b;COMPAIRE et al., 2016). ...
Article
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In this work, we present first data on biological and ecological aspects of Gobius incognitus from specimens from the Ionian Sea (central Mediterranean Sea). In particular, we provided first data on habitat preference, diet and length-weight relationship of G. incognitus on material with confirmed species identity. The species preferred "mixed bottom" (sand and rocks), where it reached relatively high abundances. The diet analysis showed that G. incognitus is a generalist and opportunistic feeder. However, small benthic crustaceans were the most important prey group. The mean total length of specimens was 6.72 ± 1.85 cm and the growth was positive allometric.
... Caudal fin rounded. Scales present on body and on predorsal area, usually visible on photographs, at least dorsally (Kovačić & Šanda 2016). ...
... Rarely, a few dots are visible also in the middle cheek but usually limited to the anterior part of the cheek, very close together, well aligned and not parallel to the lower row. Posterior angle of mouth without a well-defined dark dot (Fig. 33a), so the lower row of dots, which runs along the ventral edge of the cheek, starts with 2 dashes never preceded by a marked dot on the corner of the mouth (Kovačić & Šanda 2016;Renoult et al. 2022). ...
... Infralittoral species, known from 1-8 m depth from all kinds of rocky substrata: gravel, cobbles, boulders and bedrock, but on sand only when it is mixed with a hard substratum, never on pure sand. The substratum was bare or covered with short thallus algae (Kovačić & Šanda 2016). ...
Article
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Numerous photographs of live fishes posted by anglers and divers on social media and citizen science databases are important sources of information for ichthyological research. However, validating records that extend the known ecology and bathymetric or geographic distribution of species should rely on a rigorous identification process. The family Gobiidae, with their small size, superficial resemblance among species and high species richness are particularly difficult to identify. Therefore, the identification from photographs of live individuals of Mediterranean marine gobies from the continental shelf was studied. A dichotomous identification key is provided based on photographs of live individuals, allowing positive identification of 41 out of the 66 species reviewed in this publication. Then, for all 66 species we provide a brief description of important characters, which can be used for provisional identification for those species that could not be positively identified using the key. Pending further progress in identification of live individuals, we suggest that records extending the known geographic and ecological species distribution be taken into account only if they could be validated using the dichotomous identification key.
... The current status of two morphologically similar gobies, Gobius bucchichi Steindachner, 1870 and G. incognitus Kovačić & Šanda, 2016, in Türkiye is reviewed in the light of previously published and unpublished data sources. The latter species seems to be very common along the Aegean Sea coast, while its northern Levant distribution is represented by scattered records and requires further research. ...
... The incognito goby, Gobius incognitus, was described based on specimens collected in the Adriatic Sea, in the north-western Mediterranean (France, Banyuls-sur-Mer), in the eastern Mediterranean at Crete Island, and in Israel (Kovačić and Šanda 2016). Due to its high similarity with Bucchich's goby (G. ...
... bucchichi), the species remained unrecognized despite its distinct morphological, meristic and genetic characteristics, resulting in a conundrum on their actual occurrences. According to recent studies, the distribution of G. bucchichi appears to be restricted to the East Adriatic (Italy, Slovenia, Croatia) and a few localities in Ionian (Butrint, Albania; Gulf of Arta, Greece), Aegean Sea (Kondyli Beach, Greece) and the Black Sea (Crimea) According to recent studies, while G. incognitus seems to be common and widespread in the Mediterranean Sea coasts, although its status throughout the northern Africa shores is currently questionable (Kovačić and Šanda 2016;Tiralongo and Pillon 2020;Tiralongo et al. 2020a;Renoult et al. 2022). ...
Article
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The current status of two morphologically similar gobies, Gobius bucchichi Steindachner, 1870 and G. incognitus Kovačić & Šanda, 2016, in Türkiye is reviewed in the light of previously published and unpublished data sources. The latter species seems to be very common along the Aegean Sea coast, while its northern Levant distribution is represented by scattered records and requires further research. Gobius bucchichi is currently known only from Saros Bay (northern Aegean Sea) and Marmara Archipelago (Sea of Marmara), but a more comprehensive range should be suspected. This study confirms for the first time the presence of two closely allied gobies in Türkiye based on photographic evidence, filling the information gap to a great extent.
... Paranthias furcifer is known to drift with oil platforms [45]. Some species are native Adriatic species overlooked due to lack of targeted research and were found in the Adriatic Sea using appropriate methods, e.g., Speleogobius llorisi (Kovačić, Ordines and Schliewen, 2016) or overlooked due to their similarity to already known Adriatic fish, e.g., Lepadogaster purpurea [46,47]. The most successful research improvements were cryptobenthic fish sampling and genetic studies revealing cryptic species, both with three species discovered ( Table 3). ...
... Bonnaterre, 1788), or just recently described in the Adriatic Sea (Gobius incognitusKovačić and Šanda, 2016) ...
Article
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The paper presents an analysis of biogeographic and habitat distribution patterns, and the conservation status data of Adriatic fishes, based on the last published checklist and evidence-based critical analyses of species presence. The total number of species recorded in the Adriatic is 449. The Adriatic has 58.8% of Mediterranean species richness, 76.1% of its families, and 87.8% of its orders. Among species discovered in the Adriatic after 2010, twelve species were attributed to biological invasion, mostly Atlantic immigrants or alien species, and ten species were attributed to improved research on the native ichthyofauna of the Adriatic area. About 58% of species are native species of Atlanto-Mediterranean origin, 21% are native species of wider global occurrence, 15% are Mediterranean or Mediterranean and Black Sea endemics and 5% originated outside Mediterranean Sea. The majority of species inhabit the benthic environment (71.9%), while others occur in the pelagic environment (20.7%) or are euryhaline (7.3%). The benthic littoral species are the most numerous Adriatic fishes, representing 40% of all species richness, whereas pelagic fishes are mainly eurybathic or epipelagic; only 3.6% of species are deep pelagic species. A Red Book of marine fishes of the Adriatic Sea is urgently needed to assess their conservation status, covering the entire Adriatic Sea and reviewing all fish species to assess their conservation status.
... Previous molecular phylogenies suggested consistently a close relationship of Zosterisessor ophiocephalus (Pallas 1814) with the species of the genus Gobius (e.g. Penzo et al. 1998;Giovannotti et al. 2007;Neilson and Stepien 2009;Vanhove et al. 2011;Kovačić and Šanda 2016), although the number of studied Gobius species was always limited in the mentioned works. Our study unambiguously identifies Z. ophiocephalus inside the genus Gobius and suggests close relationships of this species with G. bucchichi/G. ...
... Our study unambiguously identifies Z. ophiocephalus inside the genus Gobius and suggests close relationships of this species with G. bucchichi/G. incognitus, concordantly with the previous studies (Penzo et al. 1998;Giovannotti et al. 2007;Vanhove et al. 2011;Kovačić and Šanda 2016). Moreover, our results show that G. ophiocephalus form two distinct, considerably divergent lineages (Fig. 6, Table S2), see later discussion. ...
Article
Gobius xoriguer sp. nov., a new offshore species of goby (Teleostei: Gobiidae: Gobiinae) is described based on three specimens collected in 2010, 2012 and 2018 in the western Mediterranean, off Menorca (Spain), in the Gulf of Lion and off Corsica (France), at 51–104 m depth on coralline algae sea bed. It is easily distinguishable from its Atlantic-Mediterranean congeners by the combination of the following characters: large eyes, ~27-28% of head length; anterior nostril with a small triangular process; 14 soft rays on D2, 13 soft rays on A; enlarged first dorsal fin rays (adult males), with third D1 spine the longest, 23-27 % SL; uppermost P fin rays not free from membrane; long V-shape pelvic fins with vestigial frenum; 50-51 scales on LL; head oculoscapular canal with pores σ, λ, κ, ω, α, β, ρ, ρ1, ρ2, and preopercular canal with pores γ, δ, ε present; row x1 ending anteriorly behind pore β; a groove section between pore ρ and ρ1; suborbital row d discontinuous with large gap below suborbital rows 3 and 4; rows o separated; seven enlarged orange blotches on body side; white dotes on cheek and opercle on an orange background.With a known maximum size of 64 mm TL, it is among the smaller species of Gobius. Bayesian inference and Maximum likelihood phylogenetic tree topologies based on mitochondrial DNA COI sequences (barcoding region), including most Atlantic-Mediterranean Gobius species, support Gobius gasteveni Miller 1974 as the closest relative to Gobius xoriguer sp. nov. These sister species exhibit a high genetic divergence of 9.5% (uncorrected p-distance).
... Small size and morphological similarity of these species may contribute to their diversity being underestimated in field surveys. Such cryptic diversity can contribute to undescribed species being hidden among sympatric populations (Berrebi et al., 2005;Kon et al., 2007;Knebelsberger & Thiel, 2014;Viswambharan et al., 2015;Kovačić & Šanda, 2016). Thus, in the past 10 years, 282 new gobiid species have been described (212 within Gobiinae) suggesting that considerable gobiid diversity is still largely undescribed (Iglesias et al., 2016). ...
... The Gobiidae is the most diverse fish family in the Mediterranean Sea (Miller, 1986) and several new Mediterranean gobiid species have been described over the past few decades (Heymer & Zander, 1992;Miller, 1992;Ahnelt & Patzner, 1995;Kovačić & Miller, 2000;Miller & Šanda, 2008;Kovačić & Šanda, 2016). Systematic and phylogenetic relationships of gobiid species are still unresolved and the sand-goby group is taxonomically difficult to identify (Webb, 1980). ...
Article
The new sand goby species Pomatoschistus nanus (Teleostei: Gobiidae) is described from the northern coast of the Levantine Sea (eastern Mediterranean Sea) based on both morphological and DNA barcoding data. The new species is the smallest fish in the Mediterranean Sea and may be distinguished from congeners by the following features: predorsal area, first dorsal-fin base and breast naked; δ-pore missing; anterior point of the suborbital row b not reaching level of posterior point of suborbital row d; slightly emarginated caudal fin and nape coloration pattern. DNA barcode data clearly discriminate Pomatoschistus spp. in the neighbour-joining tree with an average of 17·7% interspecific K2P distance. The most closely related taxon to P. nanus sp. nov. is Pomatoschistus bathi and the most distantly related is Pomatoschistus tortonesei with 11·9 and 21·9% K2P distances respectively. Morphometric and genetic data are also provided for Pomatoschistus bathi.
... Hg3 from Shafaroud (this study)]. It was already found that between sister species of Ponticola genus there is no recognizable differences in the head lateral-line system (Kovačić & Engin, 2008;Zarei, Esmaeili, Schliewen, et al., 2021b), and even the recent descriptions of new cryptic species among the entire Gobius-lineage struggled to find differences to sister species or to similar congeneric species in the head lateral-line system (e.g., Kovačić & Šanda, 2016). ...
... Furthermore, based on our data, ranges for D2 branched rays (15-16, usually 16 vs. 15-17, usually it has been pointed out that for the sister species or similar congeneric species supported by molecular data, additional effort should be invested in a search for useful diagnostic characters that can distinguish the species (Kovačić et al., 2021;Kovačić & Šanda, 2016). ...
Article
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Freshwater habitats of the Caucasus biodiversity hotspot represent a center of endemism for the gobiid genus Ponticola Iljin, 1927. Hitherto, large-scale molecular studies, owing to restricted taxon and geographical sampling, have failed to give an elaborate picture of diversity and evolutionary history of these species. Here, to contribute to filling this gap, we assessed taxonomic diversity, phylogeography and evolutionary history for the south Caspian populations of Ponticola presently classified as P. iranicus and P. patimari, using an integrative taxonomic approach comprising an entire geographic range sampling, and analyses of mitochondrial DNA haplotypes, the head lateral line system, otolith shape, and meristic and morphometric variation. All freshwater samples of the P. syrman group belong to a monophyletic clade with two main subclades: a small subclade confined to the upper Sefidroud sub-basin including the type locality of P. iranicus and a large subclade with three geographically constrained haplogroups (Hg1, Hg2, and Hg3), comprising the rest of the distribution. Hg1 showed an eastern distribution including the type locality of P. patimari, while Hg2 and Hg3 are sister groups with central and western-central distributions, respectively. The freshwater clade diverged from P. syrman during the Tyurkyanian low stand (~150 m b.s.l. lasting ~0.1 Myr), while the divergence of P. iranicus and P. patimari and radiations within P. patimari took place during the Bakunian high stand (up to 50 m a.s.l. lasting ~378–480 kya). Species delimitation analyses indicated two distinct species, corresponding to each main subclade. Although the otolith shape and lateral line analyses did not reflect with phylogeographic pattern, PCA and DFA plots of meristic and morphometric data showed a clear separation of the two major subclades corresponding to P. iranicus and P. patimari, suggesting the presence of significant morphological variation meriting formal taxonomic recognition. Overall, our findings (i) reveal the presence of two freshwater endemic species in the P. syrman group, and pending further investigation, hypothesize the presence of a third cryptic species; (ii) revise and document a narrow distributional range and low diversity for P. iranicus, in contrast to a wider distributional range and high diversity for P. patimari; (iii) suggest that the climatic oscillations of the Pleistocene were associated with the cladogenesis within the P. syrman group; and (iv) allowed for the recognition of conservation units and proposition of management measures.
... The gobiid fishes, which are important to coastal zones, comprise 98 species, and are the largest fish family in the northeastern Atlantic, the Mediterranean, and the Black Sea (Kovačıć and Patzner 2011;Van Tassell and Kramer 2014;Iglésias et al. 2015;Rothman and Goren 2015;Kovačić and Sanda 2016;. However, distribution of small gobiid fauna of the Mediterranean is relatively unknown (Kovačić 2005). ...
... However, distribution of small gobiid fauna of the Mediterranean is relatively unknown (Kovačić 2005). To date 64 gobiid species have been reported from the Mediterranean and 54 of these species are native to the Mediterranean (Kovačić 2005;Kovačić and Golani 2007;Goren et al. 2009;Salameh et al. 2010;Patzner 2015;Rothman and Goren 2015;Kovačić and Sanda 2016;. Fifty-two of these gobiid species are included in the IUCN Red List of Threatened Species as "Least Concern", one as "Endangered", and one as "Data Deficient" (IUCN 2016, www.iucnredlist.org). ...
Article
In this study gobiid fish species were investigated at 31 stations from the northeastern Aegean Sea. Different habitat types between 0 and 50 m depth were studied by scuba diving with newly modified underwater visual census with ranked prevalence using time-transect. Thirty-three species were determined as belonging to family Gobiidae. Distribution, abundance, and some ecological properties of the species were examined. Considering this, four of the observed species determined as very common, seven as common, ten as locally common, six as rare, and six as very rare in the area. Pseudaphya ferreri, Gammogobius steinitzi, and Corcyrogobius liechtensteini were determined for the first time for the area. Detailed morphological characters of some species are also given.
... The family Gobiidae (sensu van der Laan et al., 2018) is the most species-rich family of fishes in the Mediterranean with more than seventy known species in the area, and with a recent burst of the new species descriptions with seven new species published since 2016 (Kovačić & Šanda 2016;Kovačić et al., , 2017Kovačić et al., , 2018Engin & Innal, 2017;Engin & Seyhan, 2017;Engin et al., 2018). ...
Article
A new genus and species of goby, Gymnesigobius medits sp. nov., is described from the western Mediterranean slope bottoms at the Balearic Islands. The new goby belongs to the Gobius-lineage (Gobiinae). Examination of the single known specimen exhibits a unique combination of morphological characters which could not be fitted to any known genus. Gymnesigobius gen. nov. is morphologically distinguished from all other genera in the Gobius-lineage by the following combination of characters: chin without fold or barbels; mouth terminal with anterior tip above horizontal level of lower eye edge; predorsal area and first dorsal fin base naked; pelvic fin anterior membrane well developed; head with anterior oculoscapular and preopercular canals, posterior oculoscapular canal absent; pores of head canals enlarged, e.g. pores α and ρ larger or of about the same size as interspaces to pore β; six transverse suborbital rows of sensory papillae, four continuous suborbital rows in front of row b, fifth row divided in three parts but in front of row b, sixth row just as superior part above row b and below pore α; longitudinal suborbital row b barely reaching forward to the vertical from posterior edge of eye. A full description of the new genus and species is provided. The new species, collected at a depth of between 344 and 364 m (mean depth of 354 m), is one of very few gobiid species found at bathyal depths.
... Indeed, recently, new species were discovered (e.g. Van Tassell & Kramer 2014, Kova i & Šanda 2016, Engin et al. 2018) while others were recorded in new areas (e.g. Colombo & Langeneck 2013, Tiralongo & Pagano 2015. ...
Article
The present paper reports the first record of the very rare Mediterranean Didogobius schlieweni Miller, 1993 in Greece. Furthermore, this record of D. schlieweni is the first from the Eastern Mediterranean basin. Based on the current record, it seems that D. schlieweni might have a wider distribution than previously thought. A summary of D. schlieweni records is provided.
... In the Mediterranean Sea, this family includes 73 species, thus being the most diverse fish group of the region [11]. Despite this and their relevance for marine ecosystems, an increasing interest in gobies only started recently, with several published studies on their biology and ecology and the description of new species [12][13][14][15][16][17][18]. ...
Article
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Invasive alien species (IAS) are one of the greatest causes of native species extinction. Indeed, they represent a global threat for biodiversity and can also affect the economy and human health. The colonization success of IAS is presumably not only due to their biological and ecological characteristics, but also to the lack of predators and/or parasites in the invaded new areas. In the present work, we demonstrate evidence of predation of the invasive alien crab Percnon gibbesi (H. Milne Edwards, 1853) by the Rock Goby Gobius paganellus Linnaeus, 1758. The diet of G. paganellus was studied analyzing the stomach content of 162 specimens collected in the central Mediterranean Sea. The results obtained from the calculation of the diet indices, namely, frequency of occurrence (%F), percentage weight (%W), percentage abundance (%N), and the Index of Relative Importance (%IRI), showed that small benthic crustaceans were the main prey types. Additionally, these indices and the Levins’ index (Bi) clearly indicated that the invasive crab P. gibbesi was by far the most abundant prey type in the diet of G. paganellus. The relevance of this predator–prey interaction and the role of native species for the biological control of invasive ones are discussed. We also provide a general view on the diet of G. paganellus and other biological and ecological aspects of specimens studied from the central Mediterranean Sea.
... As mentioned by Kovačić and Šanda (2016) and Patzner (2019), G. incognitus shows a less restricted choice of substrate type, occuring in all kinds of substrates. In our study, the species was seen mostly in sandy-rocky bottoms and was also observed together with Anemonia viridis (snakelocks anemone). ...
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The aim of the present study is to assess the spatial distribution and habitat preferences of the cryptobenthic fish assemblages around the Gökçeada Island, where no study on these species were previously conducted. A total of 51 quadrats of 1 m2 were applied to 12 sites at the depths of 0–15 m in June, August and October 2017. The cryptobenthic fishes were studied by anaesthetic sampling. A total of 187 cryptobenthic fish specimens of 23 species were observed and 111 of them were obtained. The results showed that the 0–1 m depth zone is dominated by the blennies and shows greater density than 1–15 depth zone, where gobies are outnumbered. The average abundance of cryptobenthic fish was 3.7 individuals/m2 and Gobius incognitus represented 21% of all individuals. Among the studied species, Parablennius incognitus and P. zvonimiri were only found within 0–1 m, preferring rocky substrates with short thallus algae. Coryphoblennius galerita was found on bare rocky surfaces, mostly in 0–1 m depth interval and Microlipophrys canevae preferred rocky surfaces with short thallus algae covers in as well very shallow waters. While Lepadogaster lepadogaster preferred the depths between 3–5 m, L. candolii was observed in a wider depth range (1 and 6 m).
... These events are not exclusive for the Black Sea -in the Mediterranean Sea haplotypes 4 and Н_5 are widely spread while Н_2 and Н_3 were reported so far only along Croatia coastal areas (Kovacic & Sanda, 2016). ...
Article
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According to their origin the Black Sea gobies can be divided to Ponto-Caspian relicts and Mediterranean immigrants. The increase of species diversity in the Black Sea is a consequence of Mediterranean immigrants. The present study investigated the variability in Cytochrome b sequences isolated from two former Mediterranean immigrant species: Gobius niger and Zosterisessor ophiocephalus. The annotated at NCBI Cyt b sequences of the species were also used to extend the reach of the study. The analyses demonstrated that Black Sea inhabited by four haplotypes of G. niger. Four other haplotypes were found in sequences of Mediterranean black gobies. In the Black Sea haplotype, 6 is predominant and 83.5% of all G. niger representatives belong to it. Three other haplotypes (H_1; H_7 and H_8) were found to form isolated populations. The Tajima D-test indicated that in the Black Sea G. niger in a stage of expansion and significant evolutionary pressure according to data from Maximum Composite Likelihood model of Tamura-Nei, which can explain the accumulation of mutation and appearance of new haplotypes. Unlike G. niger, Z. ophiocephalus populations are shrinking according to Tajima D-test and only one haplotype is still surviving in isolated locations in the Black Sea. These data are in agreement with previous reports of other authors which alarmed that the grass goby is a critically endangered species close to extinction. Key words: Cytochrome b, fishes, Gobiidae, Mediterranean immigrants, Gobius, Zosterisessor
... Nový druh byl objeven ale i v evropském mořském prostředí. V roce 2016 byl popsán hlaváč nazvaný Gobius incognitus(Kovačić & Šanda 2016). Také další fylogenetické a populačně genetic- ké výzkumy prokázaly existenci dosud neznámých evolučních linií, které představují dosud neznámé a nepopsané druhy, například u parem rodu Barbus v Albánii (Marková et al. 2010), jelců rodu Telestes v západním řecku v oblasti jónského úmoří či hořavek rodu Rhodeus z oblasti východního řecka (Bar- táková et al. 2019). ...
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This critical catalogue from the “The Age of Genes” exhibition presents readers with the comprehensive development of molecular phylogenetic and taxonomic research conducted at the National Museum. The reader will thus become acquainted with the Natural History Museum, which is part of the National Museum, and within which modern molecular genetic laboratories were established in 2003 at the Zoological Department and in 2016 at the Entomological Department. To facilitate orientation in this type of research, a general introduction is provided to present in brief a global-scale general advances in the field of genetics as well as a research into the DNA molecule itself, in the form of a historical survey. From Mendel‘s Laws of 1869 to the first real photo of the DNA helix taken by a transmission electron microscope in 2012. In addition, the actual structure of the DNA molecule, its placement, function and inheritance, which are among the basic attributes of this compound, are described in detail. This publication is written in Czech language. Certainly major milestones, and not just for zoology, were Sanger‘s DNA sequencing, i.e. determining the order of the individual components (nucleotides) in the DNA obtained from an organism, and then the Mullis polymerase chain reaction, which serves to simply amplify a selected DNA segment taken from along its entire length. These two molecular genetic methods are now quite routinely used in zoology, and these approaches have opened up a whole new level of knowledge. In general, these methods are used in a major way in research into the evolution of organisms, of their phylogenetic relationships, systematics and taxonomy in general and, of course, in the study of organism populations and their interrelationships. In addition to new, hitherto unresolved questions, these methods have an impact on many already existing results, which have often been transformed into quite different forms. The catalogue presents in essence the whole of the molecular taxonomic and phylogenetic research of the Natural History Museum. It is an up-to-date list of works making use of these methods and for selected studies such approaches are described in more detail for the lay public. The reader will thus learn more about the work of the Natural History Museum, which includes describing dozens of new species and subspecies from the world’s fauna every year. This type of research not only progressively reveals more fragments of the overall diversity of organisms on our planet, but can also be used very successfully in conservation practice. At the same time, research is pointed out that was directly funded by the NAKI II project, which also gave rise to The Age of Genes exhibition and this critical catalogue that accompanies it. The final part of the catalogue consists of individual exhibit photographs with details.
... In total, there are more than 90 marine species of gobies in European seas listed in the last review [38]. However, new species are still being discovered, e.g., [36,37,39,40], and the knowledge about the distribution of many species is still quite limited e.g., [41][42][43]. The information about the population genetic diversity and phylogeography of European marine gobies is very poor too. ...
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Gobies (Gobiiformes: Gobiidae) are the most species-rich family of fishes in general, and the most abundant fish group in the European seas. Nonetheless, our knowledge on many aspects of their biology, including the population genetic diversity, is poor. Although barriers to gene flow are less apparent in the marine environment, the ocean is not a continuous habitat, as has been shown by studies on population genetics of various marine biota. For the first time, European marine goby species which cannot be collected by common fishery techniques were studied. The population genetic structure of two epibenthic species, Gobius geniporus and Gobius cruentatus, from seven localities across their distribution ranges was assessed, using one mitochondrial (cytochrome b) and one nuclear gene (first intron of ribosomal protein gene S7). Our results showed that there is a great diversity of haplotypes of mitochondrial gene cytochrome b in both species at all localities. Global fixation indices (FST) indicated a great differentiation of populations in both studied gobies. Our results did not show a geographic subdivision to individual populations. Instead, the data correspond with the model of migration which allow divergence and recurrent migration from the ancestral population. The estimated migration routes coincide with the main currents in the studied area. This matches well the biology of the studied species, with adults exhibiting only short-distance movements and planktonic larval stages.
... In addition, our results indicate that there is a strong similarity of N. melanostomus sounds with those documented from another vocal con-generic species, i.e. the monkey goby N. fluvitalis (Horvatić et al. 2015). From a comparative perspective, the two Neogobius species, sharing a close genetic and morphological relationship (Miller 2003;Neilson and Stepien 2009;Medvedev et al. 2013;Kovačić and Šanda 2016), were able to produce same sound type, with a strong overlap in the general structure and acoustic behaviour. While this resemblance in acoustic features is not so surprising for con-generic species, more striking is the similarity of acoustic signals for the two Neogobius species and another vocal Mediterranean species, the Arno goby, Padogobius nigricans. ...
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Divergence in acoustic traits between closely related species can be explained by phylogenetic history. In gobies, phylogenies reconstructed with acoustic signals primarily overlap with studies based on morphological or molecular data. Here, sound production of the two Ponto-Caspian gobies, Neogobius melanostomus and Ponticola kessleri, was recorded in controlled conditions and compared to determine the degree of interspecific acoustic variation across benthophilin gobies. Both species produced tonal-like sounds characterized by unique temporal and spectral properties during agonistic and reproductive intraspecific interactions, while the acoustic comparison revealed that the vocalizations of these two species differ in almost every acoustic property. N. melanostomus vocal structure was characterised by short (c. 100 ms), low-frequency (< 100 Hz) tonal sounds repeated at a relatively faster rate, while P. kessleri sounds appeared as a broadband, downward frequency modulated longer calls (c. 450 ms). Some acoustic features (sound rate and duration) proved to be stereotyped and could be considered a species-specific trait that could potentially be utilized to discriminate males or used by females for mate assessment. The tonal sounds appear to have a deeper origin within the Benthophilinae subfamily, as all acoustically investigated species to date have been able to produce this sound type. The recorded vocal repertoire represents a baseline for future comparative acoustic studies among the benthophilin gobies, aiming to gain additional information on the evolution of acoustic communication within Ponto-Caspian gobies and highlighting their importance in reconstructing phylogenetic relationships.
... Within the European gobiid fauna, the intensively studied sand gobies in the genus of Pomatoschistus comprise 13 species (Eschmeyer, Fricke, van der Laan, 2018). Several new gobiid species including sand gobies have been described in the Mediterranean in the past decade (Miller & Šanda, 2008;Kovačić & Šanda, 2016;Kovačić, Ordines, & Schliewen, 2016Engin & Seyhan, 2017;Engin & İnnal, 2017). ...
Article
Minor morphological differences between the Atlantic and Mediterranean populations of Pomatoschistus pictus (Malm, 1865) resulted in the description of two subspecies of this species by Miller (1973): Pomatoschistus pictus pictus and P. p. adriaticus. However, the similarity of morphometric and meristic characteristics led to an ambiguity about their status. Despite high morphological similarities between the Atlantic and Mediterranean populations, we found that the population in the Sea of Marmara (Erdek) differs from the Atlantic population in terms of the frequency of papillae in the occipital rows g and h, the position of the suborbital row b and the colouration of the second dorsal fin. Examination of the DNA sequence of the COI gene showed that the K2P genetic distance between the population in the Atlantic and the Sea of Marmara (Erdek) was 7.9%, while the minimum interspecific distance between any other Pomatoschistus species was determined as 4.3%. We concluded based on the combined genetic and morphological results that the population in the Sea of Marmara represents a species that is distinct from the Atlantic species P. pictus. It is likely that it belongs to Pomatoschistus pictus adriaticus described in the Adriatic Sea and should be given a species rank.
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Pomatoschistus anatoliae sp. n. is described from estuary of Göksu River on the Mediterranean coast of the Anatolia. It is distinguished from its congeners by the suborbital papilla pattern, meristic, and ecological features. DNA barcoding based on COI sequences revealed that there is a high nucleotide sequence divergence to the nearest neighbour. Kimura’s two parameter distances between P. anatoliae sp. n. and other species of Pomatoschistus and Knipowitchia have found to be at least 5.1%. http://www.zoobank.org/urn:lsid:zoobank.org:pub:4592CF50-07B5-4C25-892F-5D009057675B
Article
An updated and evidence-based checklist of Mediterranean Sea fishes is provided. Each of the fish species in the Mediterranean Sea listed here was either listed in the last published checklist of the Mediterranean fishes or in other, reports or new records, and the checklist is critically assessed. Out of the assessed 791 species previously reported from the Mediterranean, the presence of 759 species is confirmed while 32 species are excluded from the new checklist, by lacking evidence of presence or representing obvious taxonomic confusions. The net increase in known Mediterranean fish species richness since the last checklist is 11%. The non-native Mediterranean species now represent 22.1% (168 species) of the known Mediterranean fish diversity. The evidence-based protocol applied here provides a reliable checklist of marine fishes, for which each of the included species has indeed been recorded at least once within the discussed geographic area in the Mediterranean Sea.
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The gobies (Gobiidae) are the most diverse fish family in the Mediterranean Sea. Nevertheless, knowledge on their diversity, taxonomy, and phylogenetic relationships is still inadequate. The phylogenetic analyses reveal two genetically highly distinct clades among specimens identified as Zebrus zebrus . A new species, Zebrus pallaoroi sp. nov., is described based on an integrative approach. The neotype of Zebrus zebrus is designated. Genetic data confirm a pronounced level of divergence between Z. pallaoroi and Z. zebrus , with the mean genetic distance on cytochrome b being 18.1% and 1.07% on rhodopsin. Phylogenetic relationships within the Gobius -lineage were estimated on both markers. Morphologically, Z. pallaoroi is distinguished from the only congener Z. zebrus by having a snout longer than its eye, posterior nostril about 4/5–9/10 of the anterior nostril, eye diameter 4.3−4.7 in head length, ventrolateral head ridges transversally connected on the anterior side by a short transversal ridge, anterior membrane midline depth about 2/3 of the spinous ray, head canal pore α diameter about half of the distance between pore ρ and ρ 1 , suborbital sensory papillae row 5i going downwards to or near the level of row d , the distance between row 5i and row d absent or much smaller than the length of row 5i , and the body with ten to eleven vertical dark brown bands. Zebrus pallaoroi was recorded from the southern Adriatic, northern Ionian, and northern and western Aegean Seas, and is a cryptobenthic fish from very shallow waters.
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Characterizing fish communities must be a priority to safeguard resources and determine critical changes. Here, species richness and the spatial and temporal evolution in the structure of fish assemblages were analysed based on photos taken in underwater free-diving contests. A total of 29 contests held from 2008 to 2015 at four different locations along the northeastern Spanish coast, including a marine protected area were analysed. Contests reward the number of species per participant and photographic quality. Species image frequency from each tournament were standardized to catch image rate. A total of 88 taxa were recorded, including 32 cryptobenthic species, the highest number recorded in the Mediterranean littoral system so far. Cluster analyses yielded four major groups. Catch image rates in the marine protected area were significantly higher for seven species of high commercial interest and for two big labrids of recreational interest, including an endangered species (Labrus viridis). Overall, the study showed that photographic free-diving contest data are a potential tool for determining species richness in littoral systems since contest rules promote competition between participants to obtain maximum fish diversity. We believe that this type of cost-effective data can be applied worldwide as a complementary way of monitoring littoral fish assemblage.
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In fish, species identity can be encoded by sounds, which have been thoroughly investigated in European gobiids (Gobiidae, Gobius lineage). Recent evolutionary studies suggest that deterministic and/or stochastic forces could generate acoustic differences among related animal species, though this has not been investigated in any teleost group to date. In the present comparative study, we analysed the sounds from nine soniferous gobiids and quantitatively assessed their acoustic variability. Our interspecific acoustic study, incorporating for the first time the representative acoustic signals from the majority of soniferous gobiids, suggested that their sounds are truly species-specific (92% of sounds correctly classified into exact species) and each taxon possesses a unique set of spectro-temporal variables. In addition, we reconstructed phylogenetic relationships from a concatenated molecular dataset consisting of multiple molecular markers to track the evolution of acoustic signals in soniferous gobiids. The results of this study indicated that the genus Padogobius is polyphyletic, since P . nigricans was nested within the Ponto-Caspian clade, while the congeneric P . bonelli turned out to be a sister taxon to the remaining investigated soniferous species. Lastly, by extracting the acoustic and genetic distance matrices, sound variability and genetic distance were correlated for the first time to assess whether sound evolution follows a similar phylogenetic pattern. The positive correlation between the sound variability and genetic distance obtained here emphasizes that certain acoustic features from representative sounds could carry the phylogenetic signal in soniferous gobiids. Our study was the first attempt to evaluate the mutual relationship between acoustic variation and genetic divergence in any teleost fish.
Article
Reef ecosystems are characterized by highly heterogenous habitats and functionally diverse fish communities. Few studies have examined how functional diversity differs among habitats within these communities, i.e. species associated with a specific habitat may have similar trophic ecologies meaning that the functional diversity within the community is driven by habitat diversity, or conversely, high functional diversity within each habitat would indicate that resource segregation also occurs at the habitat level. We used stable isotopes ratios of carbon and nitrogen to estimate trophic position, resource use and ontogenetic niche shifts of 15 reef fishes associated with four distinct habitat types (cryptobenthic, epibenthic sand, epibenthic rock and hyperbenthic) on the Croatian coast of the Adriatic Sea. Trophic ecology was quite similar across fish assemblages, but there was strong evidence of niche segregation among fish species within each assemblage showing high functional diversity within each microhabitat. The sampled fish community contained benthic and pelagic resource users, along with multiple intermediate generalists. Consumer stable isotope ratios revealed considerable interspecific variation in resource use amongst fishes within each habitat type. The cryptobenthic fishes were a notable exception to this trend with the narrow range of resource use values, indicating reliance of these species on a single resource. The greatest diversity of trophic positions within a guild was observed in cryptobenthic and rock associated epibenthic fishes. The majority of observed ontogenetic variation in studied fish species reflected an increase in benthic resource use and trophic position. However, the degree of ontogenetic variation in trophic ecology of studied species, if present, was generally low, showing no dramatic change in the ecology of any species. The size structuring among guilds was considerable, with cryptobenthic fishes were the smallest on average and hyperbenthic fishes were largest, despite guilds having similar ranges of trophic positions.
Article
A checklist of 73 gobiid species (Teleostei: Gobiidae) recorded to date from the Mediterranean Sea is established following the evidence approach for checklists. The Mediterranean gobiofauna currently has 62 known native species and 11 alien species. An identification key to gobiid species known from the area is provided. The principles of character selection for the key are discussed.
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The updated checklist of Adriatic Sea fishes with a critical assessment of each species using an evidence approach is provided. Each fish species in Adriatic Sea listed in the last published checklist and those reported in published new records not included in the most recent Adriatic checklist, have been included. Of the total of 466 fish species, the presence in the Adriatic Sea was confirmed for 444 species by at least one positive record of the species in the area, the presence of 10 species is still unconfirmed, and 12 fish species are excluded from the list. An evidence approach protocol is recommended for general use for compiling checklists of marine fishes.
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Gobius kolombatovici sp. nov. is described from eleven specimens collected at the eastern coast of the Island of Krk, in the northern Adriatic Sea. The new species is assigned to Gobius on the basis of agreement with the diagnostic features of the genus, but differs from other species most obviously by the possession of a prominent dark spot on the rear part of the first dorsal fin. Meristic values for the new species are D2 I/13-14, A I/13, P 17-19 and LL 52-57. The species was found on the open substrates, at depths of 22-38 m.
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Two new tropical eastern Atlantic gobies are described, which are obligatorily living in burrows of the axiid shrimp Axiopsis serratifrons. Both new species are placed in Didogobius, which is re-diagnosed. Didogobius amicuscaridis spec. nov. from São Tomé islands and D. wirtzi spec. nov. from the Cape Verde Islands, are similar to each other in overall appearance, but differ by a suite of morphometric characters, meristics and coloration. A comparison of key characters for all known species of the apparently closely related genera Didogobius, Gammogobius and Chromogobius is provided. Definitions for all used 41 morphometric distances including several new ones are represented to serve as a reference for gobioid morphometrics.
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Although much biological research depends upon species diagnoses, taxonomic expertise is collapsing. We are convinced that the sole prospect for a sustainable identification capability lies in the construction of systems that employ DNA sequences as taxon 'barcodes'. We establish that the mitochondrial gene cytochrome c oxidase I (COI) can serve as the core of a global bioidentification system for animals. First, we demonstrate that COI profiles, derived from the low-density sampling of higher taxonomic categories, ordinarily assign newly analysed taxa to the appropriate phylum or order. Second, we demonstrate that species-level assignments can be obtained by creating comprehensive COI profiles. A model COI profile, based upon the analysis of a single individual from each of 200 closely allied species of lepidopterans, was 100% successful in correctly identifying subsequent specimens. When fully developed, a COI identification system will provide a reliable, cost-effective and accessible solution to the current problem of species identification. Its assembly will also generate important new insights into the diversification of life and the rules of molecular evolution.
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Between 2003 and 2006, almost 80 localities in all main hydrological systems in Albania were sampled and data on the distribution of loach fishes gathered. The spined loach Cobitis ohridana Karaman, 1928 was found to be a common species in Albania, occurring in most of its river systems, from the Ohrid-Drin-Shkodra system in the east and north to the River Vjosë basin in the south. Cobitis meridionalis Karaman, 1924 occurs in Lake Prespa, while a spined loach with mtDNA of Cobitis sensu stricto origin is present at least in the Ohrid-Drin- Shkodra system. The most common stone loach in Albania was found to be Oxynoemacheilus pindus (Economidis, 2005). It was caught in the basins of the rivers Vjosë, Seman, Shkumbin and Erzen. Barbatula sturanyi (Steindachner, 1892) was recorded in the River Black Drin.
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This work updates the Tunisian fish species inventory (Agnatha, Gnathostoma), analyses the bio-geographic features of this fauna, draws comparisons between Tunisian coasts sectors and makes comments on the recent changes in the Tunisian fish fauna. It is based on (1) the analysis of the ichthyological knowledge available for the Tunisian waters, (2) surveying campaigns (using the INSTM's oceanographic vessel and commercial fishing fleet) and (3) visits to the main landing points. Genera and species of recorded fish were updated. A grouping of the recorded fish species is given according to their geographic distribution into: (1) Cosmopolitan (C); (2) Atlanto-Mediterranean (AM); (3) Endemic (E); (4) Indo-Pacific (IP). The following categorisation of the recorded species is also given: species mainly distributed in cold and temperate waters (AF) and species having affinity for tropical and subtropical waters (AC). According to the data available in 2002, Tunisian fish biodiversity can be summarized as follows: Petromyzontiformes 1 species, Holocephali 1, Elasmobranchii 61, Chondrostei 1, Osteichthyes 263 for a total of 327 valid species belonging to 113 families, 219 genus and 30 orders. Most species (282 of 327) have an Atlantic origin, 16 species are considered as sub-cosmopolitan, 6 are lessepsian, and 23 are endemic to the Mediterranean. Species coming from the Northern Atlantic are mainly found in the northern zone, while the subtropical and saharian species are mainly found in the Gabes Gulf area. These results confirm the tropical character of this region. The species distribution along the Tunisian coasts is unbalanced; 135 species were found everywhere, 270 in the northern sector (from the Algerian border to Kelibia), 170 in the Central sector (from Kelibia to Ras Kapudia) and 247 in the southern sector (from Ras Kapudia to the Libyan border). Some species appear to be restricted to a particular sector; 52 were recorded only in the northern sector, 6 in the central sector and 44 only in the southern sector. A special attention has been paid to the arrival of exotic species. The Tunisian coasts are concerned by this bio-geographic phenomenon with marine invaders originating from the tropical Atlantic and Indo-Pacific areas. The Atlantic new comers are Chaunax suttkusi Caruso, 1949, Seriola fasciata (Bloch, 1793), Seriola carpenteri Mather, 1971, Pisodonophis semicinctus (Richardson, 1848), Solea senegalensis Kaup, 1858, Sphoeroides cutaneus (Günther, 1870), whereas the Indo-Pacific new comers are Parexocoetus mento (Valenciennes, 1846), Pempheris vanicolensis Cuvier and Valenciennes, 1831, Stephanolepsis diaspros (Fraser-Brünner, 1940), Siganus luridus (Rüppell, 1828), Siganus rivulatus Forsskål, 1775, Priacanthus hamrur (Forsskål, 1775).
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Through the study of the phylogeographic structure and demographic history of the common goby, Pomatoschistus microps, the influence of Quaternary climatic changes on the evolutionary history of coastal and marine fishes is investigated. Because of its sedentary life cycle in Mediterranean lagoons, it is also a good model to study more specifically if the formation of lagoons during the Holocene had an impact on population structure and demography. Mitochondrial sequences of Northeastern Atlantic and Western Mediterranean specimens were used for phylogenetic reconstructions as well as divergence time estimates, demographic history and population structure analyses. Pomatoschistus microps was a highly supported monophyletic clade including four lineages. It may have appeared 77,000 yr ago, and the divergence of its lineages likely occured shortly thereafter (between 61,000 and 54,000 yr). Most lineages had polytomic topologies, low nucleotide diversity and demographic analyses providing evidence of population expansion. Each lineage was characterized by a large number of private haplotypes. Most haplotypes found in Mediterranean localities were endemic, and one was dominant. Complex reticulated relationships connecting North European, Atlantic and Mediterranean haplotypes were observed. Moderate to high population structure was underlined. Contrary to previous published studies, no significant differentiation was observed between Atlantic and Mediterranean populations, indicating that the Gibraltar Strait is not a phylogeographic break for P. microps. Indeed, molecular dating combined with the tree topologies, phylogeographic and demographic analyses as well as high haplotype diversity underline a recent and rapid population divergence during the last glacial. However, population structure indicates that differentiation is an ongoing process. From an ancestral population trapped in the Atlantic, this goby colonized first northern Europe and later the Mediterranean Sea. Shared haplotypes could have dispersed in the western Mediterranean basin before the lagoon formation, while most private haplotypes, evidencing a recent isolation, probably diverged in lagoons after their closure.
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Gobioidei is one of the largest suborders of teleost fishes, with nearly 2000 extant species currently recognized. They have a worldwide distribution and show a spectacular variety in morphology, ecology, and behavior. Despite their importance, phylogenetic relationships among many groups of gobioids (including some of the major lineages) still remain poorly understood. In this study, we analyze sequence data of five molecular markers (two mitochondrial and three nuclear) averaging 6000 bp for 222 species of gobioids. Our study is the first to include both multiple nuclear and mitochondrial genes to reconstruct a comprehensive multilocus phylogeny of gobioids encompassing most major lineages representing the overall diversity of one of the most speciose vertebrate lineages. Two separate datasets are produced and used to specifically address the phylogenetic placement of Rhyacichthyidae and Odontobutidae, and the phylogenetic relationships among gobioid lineages. Our results strongly support that the initial split in the gobioid tree separated a clade containing Rhyacichthyidae + Odontobutidae as the sister group of all other lineages. The family Eleotrididae branches off the gobioid tree after the Rhyacichthyidae + Odontobutidae clade, followed by the Butidae as sister to the Gobiidae. Additionally, several major monophyletic groups are confidently identified within the two major Gobiidae subclades, the gobiine-like gobiids and the gobionelline-like gobiids. Robustness of the phylogenetic trees inferred here is significantly higher than that of previous studies, hence our results provide the most compelling molecular phylogenetic hypothesis of Gobioidei thus far. For the first time, we provide a comprehensive sampling of European gobies that traditionally have been divided into "transverse" and "sand gobies". We show that the European gobies cluster in three distinct lineages, the Pomatoschistus-, Aphia-, and Gobius-lineages. The former resolved within the gobionelline-like gobiids and the latter two within the gobiine-like gobiids. These findings have significant implications for our understanding of the phylogeographic origin of European gobies in the light of the closure of the Paratethys. A rogue taxon analysis identified Kraemeria as an unstable taxon decreasing support at the base of the gobiine-like gobiids. Removal of this rogue taxon significantly increased phylogenetic resolution in that part of the tree and revealed additional insights into early bursts of cladogenesis of the gobiine-like gobiids.
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Samples of the widely distributed sand goby Pomatoschistusminutus have been investigated genetically from ten localities in the north-eastern Atlantic, North Sea, western Mediterranean and Adriatic Sea. Levels of genetic diversity and differentiation were assessed with starch (SGE) and cellulose acetate (CAGE) gel electrophoresis for 13 enzyme systems. Genetic differentiation between spatial samples points to a reduction or even absence of gene flow between the Adriatic and the other samples, including the western Mediterranean Sea (pair-wise FST=0·37 and 0·32 for SGE and CAGE respectively). The sample from the Adriatic Sea was clearly differentiated from the other samples at the lactate dehydrogenase loci LDH-A* (SGE and CAGE) and LDH-C* (CAGE). Values for genetic differentiation between Venetian and other sand gobies were of the same order of magnitude as between P.minutus and its closest relative P.lozanoi, suggesting allopatric speciation in the lagoon of Venice. At locations outside the Adriatic Sea, the sand goby has the typical features of a marine fish with a high level of gene flow and a low degree of genetic differentiation.
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The study focused on two gobiid taxa from the northern Adriatic Sea. External features and coloration suggest identification of one of these with Gobius auratus Risso, 181027. Risso A Ichthyologie de Nice, ou Histoire Naturélle des Poissons du Département des Alpes Maritimes: Gobius auratus sp. n. 1810 Paris Schoell 160 161 View all references. This species is characterized by a deeply emarginated pelvic disc and meristic features of typically second dorsal fin rays I/14, anal rays I/13 and scales in lateral series 45. The live coloration of the northern Adriatic population shows a yellow basic coloration but with distinct longitudinal lines of red dots and, therefore, its colour significantly differs from the uniformly yellow coloration supposed to be typical for this species. A redescription of this putative G. auratus Risso, 181027. Risso A Ichthyologie de Nice, ou Histoire Naturélle des Poissons du Département des Alpes Maritimes: Gobius auratus sp. n. 1810 Paris Schoell 160 161 View all references is carried out to extend the morphological characteristics of the species to cover also the northern Adriatic population. Two colour morphs are described and morphometrics as well as details on the lateral line system of the species are newly included in the species description. The second Adriatic taxon was assigned to G. fallax Sarato, 1889. Both taxa from the northern Adriatic were compared to G. xanthocephalus Heymer and Zander, 199212. Heymer , A and Zander , CD . 1992. Le statut de Gobius auratus Risso, 1810 et description de Gobius xanthocephalus n.sp. de la Méditerranée (Teleostei, Gobiidae).. Zoologische Jahrbücher Abteilung für Systematik, Ökologie und Geographie der Tiere, 119: 291–314. View all references from the western Mediterranean and Atlantic indicating their clear distinction. The northern Adriatic specimens of G. auratus show some similarities with the western Mediterranean and Atlantic G. xanthocephalus concerning life coloration, but differ in a series of features such as certain morphometrics (head and pelvic disc longer in the former, fifth pelvic ray, relative to fourth, longer in the latter), meristics (second dorsal rays mostly I/14 in the former and I/15 in the latter, anal rays mostly I/13 versus I/14, scales in lateral series about 45 versus 48). Gobius fallax is distinguishable by a different coloration pattern and shows the lowest values in mean fin and scale meristics. Phylogenetic analysis of DNA sequences of the first section of the control region revealed that all individuals of both colour morphs of G. auratus and of G. fallax form a single cluster of closely related haplotypes which are not sorted according to the species, suggesting their recent origin. Only G. xanthocephalus is, in agreement to morphology, also genetically distinct and represents a separate clade. The existence of a Gobius auratus species complex is therefore confirmed.
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A checklist of the marine fish fauna of Turkey is presented for the first time. Examination of the previous studies revealed the presence of 448 species in 133 families. A total of 45 species, that has a doubtful presence at Turkish seas, was briefly discussed. The ichthyofaunal similarities of seas surrounding Turkey were compared, and the Black Sea and the Sea of Marmara were found to be 55.6% similar, whereas, the proportion computed for the Aegean and the Mediterranean Sea was 83.2%.
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Cryptic taxa present unique difficulties in the description of biological diversity, which DNA sequencing approaches often readily resolve. The tubenose goby Proterorhinus, along with other Ponto-Caspian fauna, has undergone recent Eurasian range expansion, as well as colonized the North American Great Lakes in 1990. We analysed mitochondrial (cytochrome b and cytochrome c oxidase subunit I) and nuclear (recombination activating gene 1; RAG1) DNA sequences and morphological characters from exotic Great Lakes as well as introduced and native Eurasian populations of Proterorhinus marmoratus (Pallas) sensu lato to assess their species identity and biogeographic patterns. The results obtained show marked genetic and morphological divergence that indicates species-level separation between fresh water and marine/brackish lineages, dating back approximately 3.82–4.30 million years. In addition, freshwater lineages within the Black and Caspian Sea basins show significant genetic and morphological differentiation, corresponding to an estimated 0.92–1.03 million years. We describe new evidence to support at least three separate species: the original P. marmoratus in marine and estuarine habitats within the Black Sea, a freshwater species in the Black Sea basin that was introduced to the North American Great Lakes, and another freshwater species inhabiting the Caspian Sea/Volga River basin. The freshwater tubenose goby in the Black Sea basin originally was described as Proterorhinus semilunaris (Heckel), and this is confirmed to be a valid taxon. The Caspian basin taxon may correspond with Proterorhinus semipellucidus (Kessler), a putative freshwater species in the Caspian basin that was originally described from a single specimen.
Article
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Endemic Ponto-Caspian gobies include a flock of ∼24 “neogobiin” species (containing the nominal genera and subgenera Apollonia, Babka, Neogobius, Mesogobius, Ponticola, and Proterorhinus; Teleostei: Gobiidae), of which a large proportion (5 species; ∼21%) recently escaped to invade other freshwater Eurasian systems and the North American Great Lakes. We provide its first comprehensive phylogenetic and biogeographic analysis based on 4709 bp sequences from two mitochondrial and two nuclear genes with maximum parsimony, likelihood, and Bayesian approaches. We additionally compare its relationships with the tadpole gobies (Benthophilus and Caspiosoma), which comprise a related endemic Ponto-Caspian gobiid group; along with a variety of postulated relatives and outgroups. Results of all phylogenetic approaches are highly congruent and provide very strong support for recognizing the subfamily Benthophilinae; which encompasses both the “neogobiins” and tadpole gobies, and genetically diverges from other Gobiidae subfamilies—including (non-monophyletic) Gobiinae and Gobinellinae. Benthophilinae contains three tribes: Neogobiini (Neogobius, which is synonymized here with Apollonia; containing the type species N. fluviatilis, along with N. melanostomus and N. caspius), Ponticolini (containing the genera Mesogobius, Proterorhinus, Babka, and Ponticola—elevating the latter two from subgenera and removing them from the formerly paraphyletic Neogobius), and Benthophilini (tadpole gobies). Within Ponticolini, Proterorhinus and Mesogobius comprise the sister clade of the Ponticola and Babka clade. Further work is needed to clarify the interrelationships of the tadpole gobies. Invasiveness is widespread in freshwater and euryhaline taxa of Neogobius, Proterorhinus, Babka, and Ponticola; but not in marine species, Mesogobius, or tadpole gobies.
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The program MRBAYES performs Bayesian inference of phylogeny using a variant of Markov chain Monte Carlo. Availability: MRBAYES, including the source code, documentation, sample data files, and an executable, is available at http://brahms.biology.rochester.edu/software.html. Contact: johnh{at}brahms.biology.rochester.edu
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Comparative analysis of molecular sequence data is essential for reconstructing the evolutionary histories of species and inferring the nature and extent of selective forces shaping the evolution of genes and species. Here, we announce the release of Molecular Evolutionary Genetics Analysis version 5 (MEGA5), which is a user-friendly software for mining online databases, building sequence alignments and phylogenetic trees, and using methods of evolutionary bioinformatics in basic biology, biomedicine, and evolution. The newest addition in MEGA5 is a collection of maximum likelihood (ML) analyses for inferring evolutionary trees, selecting best-fit substitution models (nucleotide or amino acid), inferring ancestral states and sequences (along with probabilities), and estimating evolutionary rates site-by-site. In computer simulation analyses, ML tree inference algorithms in MEGA5 compared favorably with other software packages in terms of computational efficiency and the accuracy of the estimates of phylogenetic trees, substitution parameters, and rate variation among sites. The MEGA user interface has now been enhanced to be activity driven to make it easier for the use of both beginners and experienced scientists. This version of MEGA is intended for the Windows platform, and it has been configured for effective use on Mac OS X and Linux desktops. It is available free of charge from http://www.megasoftware.net.
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The Mediterranean Sea is an elongated semi-enclosed basin divided in two sub-basins between which the Siculo-Tunisian Strait (STS) is the demarcation of their frontier. The STS has a shallow sill that tends to accentuate the hydrodynamic and ecological differences between the two Mediterranean sub-basins, thus constituting a well-defined biogeographical boundary. Due to the restricted geographical range of Pomatoschistus tortonesei, our sampling was concentrated around the STS, known to represent a breakpoint to gene flow for various species which demonstrated different dispersal abilities. Genetic differentiation between the western (W-MED) and the eastern (E-MED) Mediterranean has been observed.
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Although much biological research depends upon species diagnoses, taxonomic expertise is collapsing. We are convinced that the sole prospect for a sustainable identification capability lies in the construction of systems that employ DNA sequences as taxon 'barcodes'. We establish that the mitochondrial gene cytochrome c oxidase I (COI) can serve as the core of a global bioidentification system for animals. First, we demonstrate that COI profiles, derived from the low-density sampling of higher taxonomic categories, ordinarily assign newly analysed taxa to the appropriate phylum or order. Second, we demonstrate that species-level assignments can be obtained by creating comprehensive COI profiles. A model COI profile, based upon the analysis of a single individual from each of 200 closely allied species of lepidopterans, was 100% successful in correctly identifying subsequent specimens. When fully developed, a COI identification system will provide a reliable, cost-effective and accessible solution to the current problem of species identification. Its assembly will also generate important new insights into the diversification of life and the rules of molecular evolution.
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The phylogeographical patterns of a small marine fish, the common goby, Pomatoschistus microps, were assessed at 12 sites along the northeastern Atlantic coasts and the western Mediterranean Sea. A combination of two genetic markers was employed: cellulose acetate allozyme electrophoresis (CAGE) and sequence analysis of a 289 bp fragment of the mitochondrial locus cytochrome b. Both markers were congruent in revealing significant differences between samples (global FST = 0.247 for the allozymes and PhiST = 0.437 for the mitochondrial DNA data) and a pattern of isolation-by-distance. Phylogeographical analyses yielded a shallow branching structure with four groups. Three of those were confined to the Atlantic basin and showed a star-like pattern. The fourth group contained a central haplotype occurring at the edges of the species' distribution, accompanied by a few more rare variants, which were restricted to the Mediterranean Sea. A genetic break was observed around the British Isles, with distinct haplotypes dominating at either side of the English Channel. A significantly negative correlation between the degree of genetic diversity and latitude was recorded both for mitochondrial DNA (mtDNA) and allozymes in the Atlantic basin. Gene flow analysis suggested that recolonization of the North Sea and the coasts of western Scotland and Ireland may have taken place from a glacial refugium in the Southern Bight of the North Sea. These results are discussed in the perspective of possible postglacial migration routes of marine fish along the northeastern Atlantic coasts.
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jModelTest is a new program for the statistical selection of models of nucleotide substitution based on “Phyml” (Guindon and Gascuel 2003. A simple, fast, and accurate algorithm to estimate large phylogenies by maximum likelihood. Syst Biol. 52:696–704.). It implements 5 different selection strategies, including “hierarchical and dynamical likelihood ratio tests,” the “Akaike information criterion,” the “Bayesian information criterion,” and a “decision-theoretic performance-based” approach. This program also calculates the relative importance and model-averaged estimates of substitution parameters, including a model-averaged estimate of the phylogeny. jModelTest is written in Java and runs under Mac OSX, Windows, and Unix systems with a Java Runtime Environment installed. The program, including documentation, can be freely downloaded from the software section at http://darwin.uvigo.es.
Chapter
The present review covers all currently known marine and brackishwater gobies in the eastern part of the amphiatlantic Temperate North Atlantic realm sensu Spalding et al. (2007) i.e. in its four eastern provinces: the Northern European Seas, the Lusitanian, the Mediterranean and the Black Sea. The data includes African coast to the Cape Vert peninsula in Senegal, but not the Cape Verde islands. In Briggs’ (1974) classic biogeographic division of continental shelfs, temperate waters of the North-Eastern Atlantic and the Mediterranean were divided in cold-temperate Eastern Atlantic Boreal region and the warm-temperate Mediterranean-Atlantic region (Fig. 2.2.1A). The last one being further divided in the province Lusitania, including the Mediterranean, and the Black Sea and Caspian Sea as separate provinces. Spalding et al. (2007) had similarly positioned boundaries in Atlantic between cold-temperate Northern European Seas and warm-temperate Lusitania as Briggs (1974).
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A preliminary check-list of the common bony fishes of the Benghazi coasts of the Mediterranean was prepared. It appeared that the marine fish fauna at this area is dominated by the family Sparidae (10.95%), followed by Serranidae (6.97%) and finally Carangidae (4.98%). Among the listed families, there are 39 families represented by a single species and 13 families represented by 2 species. The total number of species is 201 belonging to 71 families and 15 orders. The present work reported 42 fish species belonging to 24 families as new records for the Libyan coastal waters.
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A checklist of the Mediterranean marine ichthyofauna of Israel is presented. A total of 402 species belonging to 130 families, 33 orders and 3 classes are listed. The checklist includes all synonyms for each species and citations of all known publications, listed in chronological order, concerning occurrence in Israeli waters of each species. Citations are coded to indicate the focus of each publication.
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Gobius kolombatovici sp. nov. is described from eleven specimens collected at the eastern coast of the Island of Krk, in the northern Adriatic Sea. The new species is assigned to Gobius on the basis of agreement with the diagnostic features of the genus, but differs from other species most obviously by the possession of a prominent dark spot on the rear part of the first dorsal fin. Meristic values for the new species are D2 I/13-14, A I/13, P 17-19 and LL 52-57. The species was found on the open substrates, at depths of 22-38 m.
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Habitat fragmentation is a major force that will influence the evolution of a species and its distribution range. Pomatoschistus minutus, the sand goby, has a North Atlantic–Mediterranean distribution and shows various level of habitat fragmentation along its geographic repartition. The use of mitochondrial sequences of the cytochrome b (cyt b) gene and two co-dominant sets of nuclear markers (introns and microsatellites) allowed us to describe the relationships between P. minutus populations belonging to several different geographical regions of Europe and to assess the structure of populations inhabiting the Golfe du Lion, along the French Mediterranean coast. The present study confirms that the taxon located in the Adriatic Sea (Venice) should be considered as a distinct species, separated approximately 1.75 Mya. The comparison of P. minutus between the Atlantic and western Mediterranean coasts using polymorphic co-dominant markers revealed that they belong to two demographically independent units, and thus could be considered as well as distinct species, more recently separated (0.3 Mya). The Pleistocene glaciations seem therefore to have played an important role in the diversification of this complex. Finally, at a regional scale in the Golfe du Lion, P. minutus appears to form a single huge homogeneous population. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102, 175–198.
Article
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.
Article
The pattern of genetic variability of two species of Mediterranean gobiids was compared, with reference to their different life history traits (Aphia minuta paedomorphic and pelagic; Gobius niger metamorphosed and benthic). The aim was to evaluate how different life histories can affect the genetic structure in these marine teleosts. The study was carried out on populations of both species sampled in the western Mediterranean and in the Adriatic Sea. Seven restriction endonucleases were used for the RFLP analysis of a mitochondrial DNA segment comprising the NADH dehydrogenase subunits 3, 4L and 4. The results highlighted two different patterns of genetic variation, a weak genetic structure in A. minuta and population subdivision in G. niger. These observations may be explained not only in terms of the different dispersal capabilities of these species, but also considering that A. minuta is an abbreviate iteroparous spawner while G. niger is a protracted iteroparous spawner. Because abbreviate iteroparity is a reproductive strategy selected in stable environments with high resource availability, Pliocene and Pleistocene climate oscillations may have represented factors that negatively influenced the reproductive success of A. minuta, producing demographic fluctuations and bottlenecks, as suggested by the mismatch distribution analysis. The weak genetic structure of A. minuta populations seems to be therefore due to a more recent re-colonization of the Mediterranean basin after a severe population decline, rather than to the high vagility of this pelagic goby.
Article
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.
Article
The phylogenetic relationships among 26 species of the subgenus Luciobarbus were examined through comparison of the complete sequence of the mitochondrial genes ATPase 6 and 8 and cytochrome b. The monophyletic condition of the Luciobarbus subgenus was confirmed by several data treatment methods. Findings indicate a closer relationship among species inhabiting Caucasian, Greek, and North African areas than between the latter and those of the Iberian region. The isolation of these main clades is postulated to have occurred after the Messinian salinity crisis (5.5 MY ago) when the Iberian Peninsula broke away from the African continent. Based on a calibrated molecular clock proposed herein, the subsequent splitting of this subgenus corresponds to the isolation of the other three areas (Caucasian, Greek, and North African) about 4.5 MY ago, which would have interrupted any possible gene flow between lineages. However, the present data indicate that the North African Kabilies Mountain area maintained its contact with the south Iberian Peninsula for a longer period.
Article
The increase in the number of large data sets and the complexity of current probabilistic sequence evolution models necessitates fast and reliable phylogeny reconstruction methods. We describe a new approach, based on the maximum- likelihood principle, which clearly satisfies these requirements. The core of this method is a simple hill-climbing algorithm that adjusts tree topology and branch lengths simultaneously. This algorithm starts from an initial tree built by a fast distance-based method and modifies this tree to improve its likelihood at each iteration. Due to this simultaneous adjustment of the topology and branch lengths, only a few iterations are sufficient to reach an optimum. We used extensive and realistic computer simulations to show that the topological accuracy of this new method is at least as high as that of the existing maximum-likelihood programs and much higher than the performance of distance-based and parsimony approaches. The reduction of computing time is dramatic in comparison with other maximum-likelihood packages, while the likelihood maximization ability tends to be higher. For example, only 12 min were required on a standard personal computer to analyze a data set consisting of 500 rbcL sequences with 1,428 base pairs from plant plastids, thus reaching a speed of the same order as some popular distance-based and parsimony algorithms. This new method is implemented in the PHYML program, which is freely available on our web page: http://www.lirmm.fr/w3ifa/MAAS/.
Article
The sand goby Pomatoschistus minutus is a major component of marine shelf and estuarine food webs and an important study organism in behavioural research. Yet, despite the sand goby's significance, its past and present patterns of migration and gene flow are poorly understood. Here we use the mtDNA control region and parts of the flanking tRNA genes of 63 fish from six localities in the Adriatic (Eastern Mediterranean), Western Mediterranean, Atlantic, and North Sea to investigate the phylogeography of this gobiid. Phylogenetic analyses and population genetics statistics reveal the existence of an Evolutionarily Significant Unit, sensu Moritz (1994), in the Adriatic and another in the Western Mediterranean, Atlantic, and North Sea. A possible biogeographical scenario for the separation of the ancestral population is that sand gobies in the Adriatic and Western Mediterranean split between 10,000 and 5000 years ago when due to the rise in sea temperature they migrated northwards and were bisected by the Italian peninsula. A testable prediction of this scenario is that sand gobies from the Western Mediterranean, Adriatic, and Aegean form three reciprocally monophyletic groups which are the descendants of a three-way diversification event.
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