Molecular Phylogeny of Grey Mullets (Teleostei: Mugilidae) in Greece: Evidence from Sequence Analysis of mtDNA Segments

Mitochondrial DNA sequence analysis has been used to explore genetic differentiation and phylogenetic relationships among five species of the Mugilidae family, Mugil cephalus, Chelon labrosus, Liza aurata, Liza ramada, and Liza saliens. DNA was isolated from samples originating from the Messolongi Lagoon in Greece. Three mtDNA segments (12s rRNA, 16s rRNA, and CO I) were PCR amplified and sequenced. Sequencing analysis revealed that the greatest genetic differentiation was observed between M. cephalus and all the other species studied, while C. labrosus and L. aurata were the closest taxa. Dendrograms obtained by the neighbor-joining method and Bayesian inference analysis exhibited the same topology. According to this topology, M. cephalus is the most distinct species and the remaining taxa are clustered together, with C. labrosus and L. aurata forming a single group. The latter result brings into question the monophyletic origin of the genus Liza.     

Genetic Divergence and Phylogenetic Relationships in Grey Mullets (Teleostei: Mugilidae) Using Allozyme Data

Genetic divergence and phylogenetic relationships among five species of the Mugilidae family (Liza saliens, Liza aurata, Liza ramada, Chelon labrosus, and Mugil cephalus) were investigated, in the present study, on samples taken from Messolongi lagoon in Greece, using allozyme electrophoresis. Ten enzymic systems corresponding to 22 genetic loci were assayed, among which, four were found to be polymorphic in Liza saliens, Liza aurata, and Chelon labrosus, seven in Liza ramada, while only two were polymorphic in Mugil cephalus. Several loci showed different electrophoretic patterns among the species and thus, they can be useful in species taxonomy as diagnostic markers, as well as for further evolutionary studies. It must be underlined that among them the aGPD-2* locus proved to be species-specific, while the other ones can be also used in various combinations for the same purpose. The observed heterozygosity was found to range from 0.020 to 0.051. Allele frequencies of all loci were used to estimate Nei's (1972) genetic distance, which was found to range between 0.249 and 1.171 among the five species studied. UPGMA and NJ trees, obtained by genetic distance matrix methods, as well as, a tree based on the discrete character parsimony analysis were found to exhibit the same topology. Our result show that the three species of the genus Liza are clustered together, Chelon labrosus being closer to the previous clade, while Mugil cephalus being more distinct.     

Phenetic relationships of six species of mullets (<Emphasis Type="Italic">Mugilidae</Emphasis>) from the south Adriatic,as inferred from the study of the visceral and dermal skeleton

This study examines the interspecific variability and phenetic relationships in six southern Adriatic mullet species: Mugil cephalus (Linnaeus, 1758), Chelon labrosus (Risso, 1826), Liza aurata (Risso, 1810), Liza ramada (Risso, 1826), Liza saliens (Risso, 1810), and Oedalechilus labeo (Cuvier, 1829). Twenty-three morphometric characteristics of the visceral skeleton and the dermatocranium of mullets were analyzed. The major distinguishing characters were those of the jaws. M. cephalus was clearly separated from the other species; O. labeo differed to a somewhat smaller extent; and C. labrosus was the least distinguished species. The species of the genus Liza were phenetically most similar, while the other species of mullet differed significantly from them. The greatest distances were between M. cephalus and C. labrosus, as well as between M. cephalus and O. labeo. This text was submitted by the authors in English.     

Phylogenetic relationships of nine mullet species (Mugilidae) in the Mediterranean Sea

Phylogenetic relationships among four genera and nine species (Mugil cephalus, Mugil soiuy, Liza ramada, Liza aurata, Liza abu, Liza saliens, Liza carinata, Chelon labrosus, Oedalechilus labeo) of the Mediterranean mullets (Mugilidae) were investigated by means of allozyme electrophoresis using a seven-enzyme system (CK*, GAPDH*, G3PDH*, IDHP*, ME*, MDH*, PGM*) comprising eleven putative loci. The highest genetic divergence was 1.299, detected between M. cephalus and L. aurata and the lowest (0.280) was found between L. carinata and L. saliens. The amount of genetic divergence between the genera Chelon and Oedalechilus did not appear to be high (0.285). In a UPGMA tree, all nine species were grouped in two main branchings. In the first branch, C. labrosus and O. labeo clustered as closest taxa and were sister group to L. ramada. The other four Liza species produced two sub-branching in this group; L. carinata branched together with L. saliens, and L. aurata branched together with L. abu. In the second branch the two species of the genus Mugil, M. cephalus and M. soiuy, clustered together and were clearly isolated from the other three genera.     

Length–weight relationships for 22 fish species of the Mar Menor coastal lagoon (western Mediterranean Sea)

Length–weight relationships were estimated for 22 species of the Mar Menor coastal lagoon (south‐eastern Spain): Lipophrys dalmatinus, Salaria pavo, Callionymus pusillus, Aphanius iberus, Engraulis encrasicolus, Gobius cobitis, G. niger, G. paganellus, Pomatoschistus marmoratus, Symphodus cinereus, Chelon labrosus, Liza aurata, L. ramado, L. saliens, Mugil cephalus, Diplodus puntazzo, D. sargus sargus, Lithognathus mormyrus, Sarpa salpa, Sparus aurata, Hippocampus guttulatus and Syngnathus abaster. The captures were made in shallow areas of the lagoon during 2002–2004. Significant length–weight relationships were found for all species.     

Spatial and temporal variations and assemblage structure of fish species in Beymelek Lagoon,Turkey

Spatial and temporal variations and assemblage structure of fish species were investigated in Beymelek Lagoon, on the south‐western Mediterranean coast of Turkey. A total of 3956 fish, mainly juveniles, from 24 species representing 11 families were sampled by gillnets and trammel nets from February 2006 to January 2007. Twelve of the species were marine straddlers, 11 were marine‐estuarine‐dependent and one was catadromous. Numerical contribution of marine‐estuarine‐dependent species to the total fish abundance was 86.2%, while marine straddlers contributed only 13.8%. Despite the higher number of species, the numerical abundance of marine straddlers was lower than the abundance of marine‐estuarine‐dependent species. The contribution of catadromous species by number was only 0.02%. The assemblage was taxonomically dominated by Sparidae (seven species, 51.3%), Mugilidae (five species, 36.0%), and Clupeidae (one species, 10.3%). Among sparids, Sparus aurata, Diplodus sargus and Lithognathus mormyrus contributed 38.0, 7.2 and 4.6% to the total catch, respectively. The most abundant mugilid species was Liza saliens with 25.2%, followed by Chelon labrosus 5.2%, and Liza aurata 4.0%. Contribution by the other mugilid species to total catch was quite low. The Clupeidae (10.3%) were represented by Sardinella maderensis with 9.1% of the total catch. While the upper reaches of Beymelek Lagoon were dominated by Sparus aurata and Liza saliens, dominant in the lower reaches were Sardinella maderensis, Sparus aurata, Diplodus sargus, Lithognathus mormyrus and Liza saliens. Sparids were generally caught from mid‐summer to mid‐winter while mugilids were caught throughout the year. Clupeids occurred mainly from autumn to spring.     

Analysis of mitochondrial DNA: Taxonomic and phylogenetic relationships in two fish taxa (Pisces: Mugilidae and Cyprinidae)

To solve some systematic questions as well as to study genetic variability and evolutionary relationships in two groups of fish belonging to the Mugilid (Mugilidae) and Cyprinid (Cyprinidae) families, we have used restriction fragment length polymorphism analysis of mitochondrial DNA (mtDNA) fragments amplified in polymerase chain reaction. The analysis of three mtDNA fragments of 7220 bp total length of six Mugilid species has shown that Mediterranean Liza aurata, L. ramada, L. saliens, and Chelon labrosus form a common cluster, L. aurata and C. labrosus being the closest relatives, whereas L. haematocheilus (syn. C. haematocheilus) of the Sea of Japan forms a sister group to the Mediterranean cluster. It was found that Chelon and Liza genera are paraphyletic, and therefore their division into two genera is unnatural and they should be synonymized. According to priority, Liza species should be ascribed to Chelon genus. Mugil cephalus is the most distant compared to the rest of the species studied. The level of genetic divergence between allopatric samples of M. cephalus from the Sea of Japan and the Mediterranean Sea has proved to be very high—4.5% of nucleotide substitutions. The analysis of four mtDNA fragments of 9340 bp total length of six Cyprinid species has shown that L. waleckii is the most genetically distant. Pseudaspius leptocephalus is a sister group to Tribolodon species. All Tribolodon species form a common cluster with T. sachalinensis as a root. The remaining species form two branches, one of which includes T. nakamurai and T. brandtii, another one combines T. hakonensis and a new form of Tribolodon revealed that is close to T. hakonensis by its mtDNA (2.4% of nucleotide substitutions). This new form might be an independent species. Published in Russian in Biokhimiya, 2007, Vol. 72, No. 12, pp. 1651–1658.     

Mitochondrial DNA divergence and phylogenetic relationships in mullets (Pisces: Mugilidae) of the Sea of Japan and the Sea of Azov revealed by PCR-RFLP-analysis

Three Mugilid species: Mugil cephalus (Linnaeus, 1758) and Liza haematocheila (Temminck et Schlegel, 1845; syn. Mugil soiuy, M. haematocheilus, L. soiuy, Chelon haematocheilus) from the Sea of Japan, as well as M. cephalus and Liza aurata (Risso, 1810) from the Sea of Azov were investigated on the basis of PCR-RFLP analysis of mitochondrial DNA (mtDNA) fragments, which included 12S/16S rRNA, and ND3/ND4L/ND4 genes. Among 61 individuals of three Mugilid species thirteen different haplotypes were detected. Eight and thirteen restriction endonucleases were found to be species-specific in 12S/16SrRNA and ND3/ND4L/ND4 respectively. This method may be useful for species identification. M. cephalus showed the largest genetic divergence while L. haematocheila and L. aurata were closely related and clustered together. The level of mtDNA differentiation between the two M. cephalus samples from the Sea of Japan and the Sea of Azov, i.e., nucleotide substitutions of approximately 3%, appeared to be relatively high.     

Morphological and morphometric changes of sagittae otoliths related to fish growth in three Mugilidae species

The aim of this study was to analyze morphologic and morphometric changes of sagittae otoliths throughout the growth of three mullets: Mugil liza, Mugil cephalus and Liza ramada. Fish were collected seasonally between October 2011 and April 2014, and three methods were used: regression analysis of traditional biometric characters; morphological analysis for group identification; and morphometrical analysis of shape indices among groups (circularity, rectangularity, aspect ratio and percentage occupied by the sulcus). In all species, dependence of standard length on otolith length and height were best described as power functions. Two morphological groups were identified for M. liza and M. cephalus, while three for L. ramada. Morphological changes were supported by morphometry differences only for the first two species. Smaller size specimens of M. liza had more rectangular otoliths with more percentage covered by the sulcus while bigger fish tended to have a more circular shape. For M. cephalus smaller size fish had a more elliptical shape. The observed changes may be reflecting life history changes, related to habitat or dietary shifting.     

Saturnius minutus n. sp. and S. dimitrovi n. sp. (Digenea: Hemiuridae) from Mugil cephalus L. (Teleostei: Mugilidae), with a multivariate morphological analysis of the Mediterranean species of Saturnius Manter, 1969

Three species of the bunocotyline genus Saturnius Manter, 1969 are described from the stomach lining of mugilid fishes of the Mediterranean and Black Seas. Two of the species are new: S. minutus n. sp. occurs in Mugil cephalus off the Mediterranean coast of Spain; and S. dimitrovi n. sp., a parasite of M. cephalus off the Bulgarian Black Sea coast and the Spanish Mediterranean coast, was originally described as S. papernai by Dimitrov et al. (1998). In addition, S. papernai Overstreet, 1977 is redescribed from M. cephalus off the Spanish Mediterranean coast and from Liza aurata and L. saliens off the Bulgarian Black Sea coast. The three species are distinguished morphometrically using univariate and multivariate analyses. These results were verified using Linear Discriminant Analysis which correctly allocated all specimens to their species designations based on morphology (i.e. 100% successful classification rate) and assigned almost all specimens to the correct population (locality). The following variables were selected for optimal separation between samples: the length of the forebody, ventral sucker and posterior testis, the length and width of the posteriormost pseudosegment, and the width of the muscular flange at ventral sucker level.     

L'Intestin chez les Mugilidae (Poissons; Téléostéens) a différentes étapes de leur croissance I. Aspects morphologiques et histologiques

The intestine of Mugilidae (Pisces; Teleostei) at different growth stages. I. Morphological and histological aspects A morphological and histological study of the intestine of Mugilidae (Mugil cephalus, Liza aurata, Liza ramada and Liza saliens) has been carried out from the larval to the adult stages. The allometric relationship between length of intestine and standard length of fish are represented by regression lines correlated with species and kind of diet. S.E.M. observations on the intestinal mucosa surface show finger-like villi, as in Mammals. Histologically, the mucosa structure appears similar to that in others Teleosteans. The gobblet cells are scarce in the larval stages but very numerous in the juvenile stages; furthermore, they are more abundant in the posterior than the anterior part of the intestine (respectively 25 and 10%). Change in the feeding habits observed in young stages has no effect on the features of the intestine but notably increases its relative length.     

Effects of leaping grey mullet Liza saliens (Osteichthyes, Mugilidae) in the macrophyte beds of oligohaline Mediterranean coastal lagoons

Some mugilid fish are known to enhance small phytoplankton in freshwater macrophyte-free environments due to zooplankton depletion. This suggests that they may have negative effects on natural macrophyte beds of freshwater and oligohaline lagoons due to phytoplantkon enhancement. To test this hypothesis, we compared the ecosystems of control enclosures that contained no fish with those of enclosures stocked with Liza saliens at two different densities. The occurrence of L. saliens at a density of 321±92.42 kg ha^–1 reduced cladoceran density, depleted epiphytic chironomid larvae, enhanced mayfly nymphs and cyclopoid copepods and reduced the organic matter content of sediment, all in comparison with control enclosures. At a density of 673±42.04 kg ha^–1, L. saliens reduced total zooplankton density, depleted epiphytic and sediment dwelling chironomid larvae and enhanced mayfly nymphs. The organic matter contents of sediment was not affected. These results showed that L. saliens was very effective in reducing zooplankton density even when macrophyte biomass was high. However, these effects do not affect phytoplankton density, probably because zooplankton was dominated by species with low filter-feeding rates and macrophytes depleted nutrients.     

Metals (Hg,Pb, Cu,and Zn) Bioaccumulation in Sediment,Fish, and Human Scalp Hair: A Case Study from the City of Mersin Along the Southern Coast of Turkey

This study investigates mercury, lead, copper, and zinc concentrations in six most frequently consumed fish species (120 samples), sediments (20 samples) taken from Karaduvar Fishing Area where fish species live, and Mersin Port as a contrary region, and human scalp hair for people regularly consuming these fish species (50 samples) and non-fish-eaters (15 samples) in Mersin, Turkey. On taking living environment into account, the fish groups include pelagic species of Liza saliens, Liza aurata, and demersal species of Merluccius merluccius, Mullus barbatus, Upeneus moluccensis, and Solea solea. Total Hg (THg) was found to accumulate in muscle tissues at the lowest concentration (0.01 μg/g) in L. saliens and at the highest (2.66 μg/g) in S. solea. Pb was only detected at high concentrations of 1.86 μg/g in M. barbatus and of 2.16 μg/g in M. merluccius. Cu and Zn concentrations were below the detection limit within all fish species. In the sediment samples, Pb and Cu concentrations were persistently below their effect range–median (ERM) value, whereas this limiting value only maintained for 15% of THg concentrations. On the other hand, the effect range–low (ERL) of sediment exceeded at Pb in 15% of samples and Cu in 25% of samples. Zn remained below the detection limit for sediment samples. The metal concentrations at scalp hairs of regular consumers of these fish groups and non-fish eaters vary from the range 0.40–3.28 to 0.14–1.02 μg/g for THg, 11.16–107.84 to 8.00–22.38 μg/g for Pb, and 151.67–645.35 to 144.92–343.50 μg/g for Zn. An important finding of the present study is the significant adverse impact of sedimentary heavy metal bioaccumulation to human through the consumption of demersal fishes in the city of Mersin along the southern coast of Turkey.     

A review of changes in the fish assemblages of Levantine inland and marine ecosystems following the introduction of non-native fishes

The arrival of non‐native fishes in the Levant Basin began in the late 19th century. Whereas the presence of most of the 40 non‐native freshwater fishes stem from intentional introductions, either for aquaculture or pest control, the 62 species of non‐native marine fishes arrived by natural dispersal via the Suez Canal. Of the non‐native freshwater species, five have established successful breeding populations (mosquitofish Gambusia affinis, common carp Cyprinus carpio, crucian carp Carassius carassius, swordtail Xiphophorus hellerii and rainbow trout Oncorhynchus mykiss), and seven are regularly stocked in natural habitats (thinlip mullet Liza ramada, flathead mullet Mugil cephalus, European eel Anguilla anguilla, grass carp Ctenopharyngodon idella, Asian silver carp Hypophthalmichthys molitrix, bighead carp Aristichthys nobilis, black carp Mylopharyngodon piceus). Some non‐native species appear to have out‐competed native species. Gambusia affinis may have caused the extirpation of two native cyprinid fishes from the Qishon River basin (Levant silver carp Hemigrammocapoeta nana and common garra Garra rufa) and the southern Dead Sea (endemic Sodom's garra G. ghoerensis). The opening of the Suez Canal in 1869 allowed entry into the eastern Mediterranean of Indo‐Pacific and Erythrean biota, with the latter now dominating the community structure (50–90% of fish biomass) and function (altered native food web) of the Levantine littoral and infra‐littoral zones. The process has accelerated in recent years concurrent with a warming trend of the seawater. Record numbers of newly discovered non‐native species is leading to the creation of a human‐assisted Erythrean biotic province in the eastern Mediterranean.     

Use of fish parasite species richness indices in analyzing anthropogenically impacted coastal marine ecosystems

The diversity of fish parasite life history strategies makes these species sensitive bioindicators of aquatic ecosystem health. While monoxenous (single-host) species may persist in highly perturbed, extreme environments, this is not necessarily true for heteroxenous (multiple-host) species. As many parasites possess complex life cycles and are transmitted through a chain of host species, their dependency on the latter to complete their life cycles renders them sensitive to perturbed environments. In the present study, parasite communities of grey mullet Liza aurata and Liza ramada (Mugilidae) were investigated at two Mediterranean coastal sites in northern Israel: the highly polluted Kishon Harbor (KH) and the relatively unspoiled reference site, Ma'agan Michael (MM). Both are estuarine sites in which grey mullet are one of the most common fish species. The results indicate that fish at the polluted site had significantly less trematode metacercariae than fish at the reference site. Heteroxenous gut helminths were completely absent at the polluted sampling site. Consequently, KH fish displayed lower mean parasite species richness. At the same time, KH fish mean monoxenous parasite richness was higher, although the prevalence of different monoxenous taxa was variable. Copepods had an increased prevalence while monogenean prevalence was significantly reduced at the polluted site. This variability may be attributed to the differential susceptibility of the parasites to the toxicity of different pollutants, their concentration, the exposure time and possible synergistic effects. In this study, we used the cumulative species curve model that extrapolates "true" species richness of a given habitat as a function of increasing sample size. We considered the heteroxenous and monoxenous species separately for each site, and comparison of curves yielded significant results. It is proposed to employ this approach, originally developed for estimating the "true" parasite species richness for a given habitat, in the characterization of communities of differentially impacted coastal marine ecosystems. Communicated by H. von Westernhagen, A. Diamant     

Trophic shift in young‐of‐the‐year Mugilidae during salt‐marsh colonization

This study investigated the trophic shift of young‐of‐the‐year (YOY) thinlip grey mullet Liza ramada and golden grey mullet Liza aurata during their recruitment in a salt marsh located on the European Atlantic Ocean coast. Stable‐isotope signatures (δ¹³C and δ¹⁵N) of the fishes followed a pattern, having enrichments in ¹³C and ¹⁵N with increasing fork length (L_F): δ¹³C in fishes < 30 mm ranged from ?19.5 to ?15.0‰, whereas in fishes > 30 mm δ¹³C ranged from ?15.8 to ?12.7‰, closer to the level in salt‐marsh food resources. Large differences between the δ¹⁵N values of mugilids and those of food sources (6·0‰ on average) showed that YOY are secondary consumers, similar to older individuals, when feeding in the salt marsh. YOY mugilids shift from browsing on pelagic prey to grazing on benthic resources from the salt marsh before reaching 30 mm L_F. The results highlight the role of European salt marshes as nurseries for juvenile mugilids.     

Intertidal pools as alternative nursery habitats for coastal fishes

Fish were sampled monthly in four tidal pools, for two years, on the west Portuguese coast. Species diversity of transient fish was higher than that found in previous studies, in other parts of the world. The transient fish population comprised six species: the white seabream, Diplodus sargus, sand smelt, Atherina spp., the thinlip grey mullet, Liza ramada, the Baillon's wrasse, Symphodus bailloni, the zebra seabream, Diplodus cervinus and the European pilchard, Sardina pilchardus. Abundance varied seasonally, yearly, and among pools, with peak numbers in spring and summer. The most abundant species in all pools, both as larvae and juveniles, was D. sargus. Diplodus sargus and Atherina spp. were present in most pools, from spring to autumn, while rare species were present mostly in the spring-summer period. Smaller mean sizes of larvae and juveniles were observed at the beginning of spring of 2011 (March–April) and at the end of spring/beginning of summer of 2012 (May–June). Mean size of larvae and juveniles often showed a continuous increase from spring to autumn in both years. The highest density peaks were due to the high number of post-larvae entering the pools in spring. In most pools, the overall condition (Fulton's K) of D. sargus increased throughout the year, in both years. The species richness, the high densities of early stages, and their continuous growth observed in tidal pools strongly emphasize the importance of these environments for larvae and juveniles of several transient marine fishes.     

A review of length–weight relationships of fish from the Segura River basin (SE Iberian Peninsula)

We present the relationships between fork length and total weight for 14 fish species from the Segura River basin (southeastern Spain): Barbus sclateri, Chondrostoma polylepis, Gobio lozanoi, Aphanius iberus, Micropterus salmoides, Lepomis gibbosus, Carassius auratus auratus, Cyprinus carpio carpio, Sander lucioperca, Gambusia holbrooki, Liza ramado, Mugil cephalus, Atherina boyeri and Pomatoschistus marmoratus. Significant length–weight relationships were found for all species. The values of the exponent b of the length–weight relationships ranged from 3.82 for Sander lucioperca to 2.59 for Micropterus salmoides.