Ligophorus pilengas Sarabeev & Balbuena, 2004 (Monogenea: Ancyrocephalidae) is proposed as the senior synonym of L. gussevi Miroshnichenko & Maltsev, 2004

Ligophorus pilengas Sarabeev & Balbuena, 2004 and L. gussevi Miroshnichenko & Maltsev, 2004, both recently described as new species from the so-iuy mullet Mugil soiuy in the Sea of Azov and Black Sea, were compared. A morphometric analysis of both forms strongly suggests that they represent the same species. According to the 'Principle of priority', L. pilengas is proposed as a senior synonym of L. gussevi.     

<Emphasis Type="Italic">Ligophorus llewellyni</Emphasis> n. sp. (Monogenea: Ancyrocephalidae) from the redlip mullet <Emphasis Type="Italic">Liza haematocheilus</Emphasis> (Temminck &; Schlegel) introduced into the Black Sea from the Far East

Ligophorus llewellyni n. sp. (Ancyrocephalidae: Ligophorus Euzet & Suriano, 1977) is described from the gills of Liza haematocheilus (Temminck & Schlegel) introduced into the Black Sea from the Far East. Ligophorus llewellyni is closely related to L. pilengas Sarabeev & Balbuena, 2004, which parasitises the same host species. The two species differ in the morphology of the accessory piece of the copulatory organ and in some of the characters of the haptoral hard-parts. The morphometric variability of L. llewellyni and in its morphologically most similar congeners from the Black Sea is studied. Correlations between 30 morphometric characters of the haptoral hard-parts and the significance of each for species differentiation are examined. It is suggested that only 22 characters are useful as diagnostic criteria permitting the differentiation of morphologically similar species of Ligophorus.     

Ligophorus pilengas Sarabeev & Balbuena, 2004 (Monogenea: Ancyrocephalidae) is proposed as the senior synonym of L. gussevi Miroshnichenko & Maltsev, 2004

Ligophorus pilengas Sarabeev & Balbuena, 2004 and L. gussevi Miroshnichenko & Maltsev, 2004, both recently described as new species from the so-iuy mullet Mugil soiuy in the Sea of Azov and Black Sea, were compared. A morphometric analysis of both forms strongly suggests that they represent the same species. According to the ‘Principle of priority’, L. pilengas is proposed as a senior synonym of L. gussevi.     

Morphometric variability between and within species of Ligophorus Euzet & Suriano, 1977 (Monogenea: Ancyrocephalidae) in the Mediterranean Sea

Monogeneans belonging to the genus Ligophorus Euzet & Suriano, 1977 are parasitic on the gills of several fish species of the family Mugilidae (mullets). At present, 12 species belonging to this genus have been described in the Mediterranean Sea and each host species is infected by a number of Ligophorus species that are not found on other mullets. A multivariate study using principal component and canonical variate analyses was carried out, with the aim of investigating the extent of morphometric variability within and between the 12 Mediterranean species of Ligophorus. Significant intraspecific morphometric variation was detected between two allopatric populations of L. angustus Euzet & Suriano, 1977 parasitic on Chelon labrosus from Sardinia; however, the existence of a new species of Ligophorus needs further evidence. Analysis of interspecific variation confirms the validity of the morphological keys used at present for the identification of Ligophorus at the species level and provides a picture of the relationships between species, indicating that the two species parasitising Mugil cephalus, L. chabaudi Euzet & Suriano, 1977 and L. mugilinus (Hargis, 1955) are the most differentiated. The hypothesis that strict host-parasite ecological relationships may be the result of co-speciation processes is briefly discussed.     

Ligophorus abditus n. sp. (Monogenea: Ancyrocephalidae) and other species of Ligophorus Euzet & Suriano, 1977 infecting the flathead grey mullet Mugil cephalus L. in the Sea of Japan and the Yellow Sea

As a result of the re-examination of museum slides and new material of monogeneans collected from Mugil cephalus L. in the Sea of Japan, the estuary of a river which flows into the Sea of Japan, the Yellow Sea (off Zhifu, at the boundary of the Bohai Sea) and the East China Sea (off the Ryukyu Islands), five species of Ligophorus Euzet & Suriano, 1977 were identified, one of which is new. The known species are L. chabaudi Euzet & Suriano, 1977, L. cheleus Rubtsova, Balbuena & Sarabeev, 2007, L. domnichi Rubtsova, Balbuena & Sarabeev, 2007 and L. pacificus Rubtsova, Balbuena & Sarabeev, 2007, which are reported from the Yellow Sea; in addition, L. domnichi is reported for the first time from the East China Sea. Ligophorus abditus n. sp., from the Sea of Japan, differs from its most similar congeners, L. pacificus and L. domnichi, in the shapes of the dorsal anchors and the accessory piece of the male copulatory organ. A comparison of all of the species of Ligophorus recovered from M. cephalus in the Sea of Japan was carried out using Principal Component Analysis, and their distribution and origin are discussed.     

Kudoa iwatai (Myxosporea: Multivalvulida) in wild and cultured fish in the Red Sea: redescription and molecular phylogeny

Gilt-head sea bream, Sparus aurata L., the Mediterranean's most important mariculture species, has been cultured for the last 30 yr in Eilat (Israeli Red Sea). Kudoa sp. was the first myxosporean parasite reported from this species. In recent years, an increase in prevalence in both land-based and sea-cage facilities in Eilat has been observed. Infections with the same Kudoa species appeared in cultured European sea bass Dicentrarchus labrax (L.) and grey mullet Mugil cephalus in the same farms, as well as in 10 species of wild Red Sea reef fish, indicating that Kudoa sp. is not fastidious with regard to its host. All affected species displayed 1- to 2-mm (up to 5 mm) whitish, spherical, or oval polysporous plasmodia. The parasite established multiple site infections, most commonly in the muscles and intracranial adipose tissue of the brain and eye periphery. Other sites were subcutaneous adipose tissue, nerve axons, mouth, eye, mesenteries, peritoneum, swim bladder, intestinal musculature, heart, pericardium, kidney, and ovary. On the basis of spore morphology, the parasite was identified as Kudoa iwatai Egusa and Shiomitsu, 1983. Ultrastructural features were comparable to those of previously studied Kudoa species. The 18S rDNA from 7 Red Sea isolates was sequenced and compared with the sequence of the same gene from K. iwatai isolated from cultured red sea bream, Pagrus major, in Japan. The phylogenetic position of K. iwatai within the genus was determined using sequence analysis of all related taxa available in GenBank. The 3 isolates of K. iwatai clustered together on a newly formed, highly supported clade. The Red Sea strain of K. iwatai is apparently native to the region. In the absence of records of this Kudoa sp. from the extensive Mediterranean sea bream and sea bass production industries, introduction with its Mediterranean hosts seems unlikely. Therefore, we conclude that K. iwatai is an Indo-Pacific species that, in the Red Sea, has extended its host range to include the allochthonous gilt-head sea bream, European sea bass, and grey mullet.     

The description of Gyrodactylus mulli sp. n. (Monogenea: Gyrodactylidae) from the Black Sea blunt-snouted mullet Mullus barbatus ponticus

A new species Gyrodactylus mulli sp. n. is described from the thorax fins of blunt-snouted mullet Mullus barbatus ponticus by the collections of B. E. Bychowsky made in 1947 near Karadag on the Black Sea. Gyrodactylus mulli sp. n. differs from G. alviga Dmitrieva et Gerasev, 2000 (described from blunt-snouted mullet too) in having another morphotype of the marginal hooks. The new species differs from G. proterorhini Ergens, 1967, which has the identical type of the marginal hooks, in having longer membrane of the ventral connective bar, longer point of the anchors, shorter marginal hooks, and lesser size of the cirrus.     

Molecular phylogeny of species of Ligophorus (Monogenea: Dactylogyridae) and their affinities within the Dactylogyridae

The taxonomic framework of Ligophorus, monogenean specialists of the gills of grey mullets (Mugilidae), is evaluated and its interspecific relationships are assessed for the first time using molecular data. The position of Ligophorus within the paraphyletic Ancyrocephalinae is re-assessed based on newly sequenced species. Furthermore, the relationship between morphometric and genetic interspecific similarities is evaluated. Partial 28S and complete ITS1 rDNA sequences from representatives of 14 of the 16 nominal species of Ligophorus from the Mediterranean, Black and Azov Seas were analysed together with published sequences of members of the Dactylogyridae. The phylogenetic analyses of the Dactylogyridae (i) confirmed the position of Ligophorus within the marine Ancyrocephalinae; (ii) revealed a sister relationship between Ergenstrema and Ligophorus, whose species are all exclusive parasites of grey mullets; and (iii) substantiated the affinities of Ergenstrema with the marine Ancyrocephalinae. The phylogenetic analysis restricted to Ligophorus confirmed the distinct status of the included species. The ITS1 region provided the highest divergence between species and phylograms with the strongest branch support. Both the 28S and ITS1 phylograms revealed two main clades. One included species from hosts with Mediterranean and NE Atlantic distribution and another was formed by species parasitising several Liza spp., including Lz. haematocheilus from the Northwestern Pacific, and Mugil cephalus, which suggests an origin outside the Mediterranean for the latter clade. The phylogenetic evidence presented herein indicated that a combination of host-switching and lineage duplication events accounted for the diversification of this genus in the Mediterranean basin. The agreement between molecular and morphological interspecific similarities observed in Ligophorus supports the validity of morphometric characters used for species identification.     

New species and new records of Ergasilus Nordmann (Copepoda: Ergasilidae) from the gills of grey mullet (Mugilidae)

Four new Ergasilus species are described from five host species of grey mullet, family Mugilidae, held in the fish collections of The Natural History Museum, London. Ergasilus extensus n. sp. was found on Myxus petardi (Castelnau) from Australia, E. sittangenesis n. sp. on Sicamugil hamiltoni (Day) from Burma, E. piriformis n. sp. on Sicamugil cascasia (Hamilton Buchanan) from India, and E. ecuadorensis n. sp. on Mugil curema Valenciennes and M. hospes Jordan & Culver from Ecuador. New records of E. cyanopictus Caravalho, E. magnicornis Yin, E. orientalis Yamaguti and E. rostralis Ho, Jayarajian & Radhakrishnan on grey mullet are also given.     

Descriptions of eight new species of <Emphasis Type="Italic">Ligophorus</Emphasis> Euzet &; Suriano, 1977 (Monogenea: Ancyrocephalidae) from Red Sea mullets

Eight new species of Ligophorus Euzet &; Suriano, 1977 (Monogenea: Ancyrocephalidae) are described from two species of mullets from the Red Sea. Ligophorus bykhowskyi n. sp. and L. zhangi n. sp. from Crenimugil crenilabris (Forsskål) differ from other species of the genus in the structure of the male copulatory organ, which has a simple accessory piece and a wide copulatory tube that arises from a large, single-chambered, expanded base. Ligophorus simpliciformis n. sp., L. bipartitus n. sp., L. campanulatus n. sp., L. mamaevi n. sp., L. lebedevi n. sp. and L. surianoae n. sp. from Liza carinata (Valenciennes) are differentiated on the basis of the morphometrics of the hard parts of the haptor and male copulatory organ. The eight species represent the first records of species directly attributed to Ligophorus from the Red Sea. Measurements of the haptoral hard-parts and the male copulatory organ of the new species are analysed with the aid of Principal Component Analysis. Three morphological types of male copulatory organ, five types of anchor, and two types of ventral and three types of dorsal bars were distinguished among these species. L. bykhowskyi and L. zhangi from C. crenilabris have the same type of male copulatory organ and anchors. Those species from Liza carinata have only one common morphological character, a thick copulatory tube, but have two types of accessory piece, four types of anchors and three types of bars. All species of Ligophorus found on mullets in the Red Sea have an accessory piece without a distal bifurcation and thus differ from most species of this genus from other regions of the world’s oceans.     

Far Eastern mullet Mugil soiuy Basilewsky (Mugilidae, Mugiliformes): the genetic structure of populations and its change under acclimatization

The introduction of Far Eastern mullet (pilengas) in the Azov Sea in the 1970s-1980s has resulted in the formation of a self-reproducing commercial population. We have carried out a comparative population-genetic analysis of the mullet from the native (Primorye, the Sea of Japan basin) and the new (The Azov Sea basin) ranges. Genetic characteristics of three Primorye and three Azov local samples were studied using electrophoretic analysis of 15 enzymes encoded by 21 gene loci. In the Azov mullet, the initial heterozygosity characteristic of the donor population was preserved while the genotype and the allele compositions changed; the changes included a 1.9-fold reduction in the percentage of polymorphic loci and 1.5-fold reduction in the mean number of alleles per locus. The genetic differences between the Azov and the Primorye sample groups were highly significant. In the native range, no genetic differentiation among the mullet samples from different areas was found (Gst = 0.42%), whereas in the Azov Sea basin, the samples from spatially isolated populations (ecological groups) exhibited genetic differences (Gst = 1.38). The genetic divergence of the subpopulations and the excess of heterozygotes at some loci in the Azov mullet suggest selection processes that formed genetically divergent groups associated with the areas of different salinity in the new range. The salinity level is assumed to be the most probable factor of local differentiating selection during fast adaptation and naturalization of the introduced mullet.     

Gorgonians (Anthozoa: Octocorallia) of the northwestern Sea of Japan

Three species of gorgonians (Primnoa pacifica, Calcigorgia japonica, and Thouarella superba) are reported as new records for the northwestern Sea of Japan; one of them (Calcigorgia japonica sp. n.) has been described as a new species. Variations of colony morphology in some species are described; the taxonomical value of specific characters is discussed. The geographical range of Primnoa pacifica in the northwestern Pacific is shown to be wider than was considered earlier. Variations of sclerite morphology (number of marginal plates, presence of additional basal plates, and number of marginal spines on the basal plate) are regarded as intracolonial or intraspecific variability. The distribution range of Thouarella superba described earlier from the Aleutian Islands (Nutting, 1912) was shown to extend into the low-boreal part of the boreal Pacific. An examined specimen of Paragorgia sp. differed from all known species and appeared similar only to P. regalis Nutting, 1912 reported from the eastern coast of Japan. The most distinctive characters of Calcigorgia japonica sp. n. were the shape of polyp, with a thin lower part looking like folded podia (in fixed specimens) and the presence of two types of clubs scattered among the sclerites of the polyp.     

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.     

Far Eastern Mullet Mugil soiuy Basilewsky (Mugilidae,Mugiliformes): The Genetic Structure of Populations and Its Change under Acclimatization

The introduction of Far Eastern mullet (pilengas) in the Azov Sea in the 1970s–1980s has resulted in the formation of a self-reproducing commercial population. We have carried out a comparative population-genetic analysis of the mullet from the native (Primorye, the Sea of Japan basin) and the new (The Azov Sea basin) ranges. Genetic characteristics of three Primorye and three Azov local samples were studied using electrophoretic analysis of 15 enzymes encoded by 21 gene loci. In the Azov mullet, the initial heterozygosity characteristic of the donor population was preserved while the genotype and the allele compositions changed; the changes included a 1.9-fold reduction in the percentage of polymorphic loci and 1.5-fold reduction in the mean number of alleles per locus. The genetic differences between the Azov and the Primorye sample groups were highly significant. In the native range, no genetic differentiation among the mullet samples from different areas was found (G _st = 0.42%), whereas in the Azov Sea basin, the samples from spatially isolated populations (ecological groups) exhibited genetic differences (G _st = 1.38%). The genetic divergence of the subpopulations and the excess of heterozygotes at some loci in the Azov mullet suggest selection processes that formed genetically divergent groups associated with the areas of different salinity in the new range. The salinity level is assumed to be the most probable factor of local differentiating selection during fast adaptation and naturalization of the introduced mullet.     

Policlithrum ponticum sp. n. (Monogenea: Gyrodactylidae: Polyclithrinae) from Mugil cephalus from the Black Sea and problems of suprageneric systematics of the gydrodactylids

Polyclithrum ponticum sp. n. is described and P. mugilini Rogers, 1967 is redescribed. Both monogenean species are parasites of Mugil cephalus in the Black Sea. The new species differs from P. mugilini, P. alberti and P. boegeri by the lesser size of anchors, while it is distinguished from P. corallense by the larger size of these structures. P. ponticum sp. n. differs from all formerly described species by the greater length of dorsal connective bar. In both species from the Black Sea, "ear-like" structures situated near the external roots of anchors are described for the first time. It is suggested, that these structures take part in longitudinal, two-lobe folding of the haptor. The process of opening the haptor is probably performed by the additional bars of the haptor (bars 2 and 3 after: Rogers, 1967), joined to each other and with the anchors. The fifth pair of additional bars (Ernst e. a., 2000) derives from the "beard" of ventral connective bar and is united with its basal part. The sixth pair of additional bars (Ernst e. a., 2000) is considered as a typical "ribs" of the haptor, and therefore the "ribs" are represented by three pairs. Differences between marginal hooks of P. ponticum sp. n. and P. mugilini are insignificant, that probably depends on the presence of "ribs" of the haptor. Based on the subdivision of marginal hooks into two groups, the presence of additional supporting structure in the haptor, and the presence of the seminal receptacle, it is suggested that the subfamily Polyclithrinae Rogers, 1967 should include the genera Polyclithrum Rogers, 1967, Swingleus Rogers, 1969, Macrogyrodactylus Mamlberg, 1959, and probably Fundulotrema Hargis, 1955. Based on such characters as the lack of the anchors, the presence of suckers in the haptor, and ovipositing of eggs, it seems to be expedient to use the following taxa in systematics of gyrodactylids: Isancistrinae Fuhrmann, 1928 (genera Isancistrum, Anacanthocotyle); Gyrdicotylinae Vercammen-Grandjean, 1960 (Gyrdicotyle) and Ooegyrodactylinae Harris, 1983 (genera Phanerothecium, Ooegyrodactylus, Nothogyrodactylus, Hyperopletes).     

The osteology of the golden grey mullet Liza aurata (Teleostei: Mugiliformes: Mugilidae) including interactive three-dimensional reconstructions

Grey mullets are remarkably characterized by their overall uniform external morphology. Identifying species as well as positioning the Mugiliformes in a phylogenetic context is rather difficult. Most recently they were placed in the newly erected Ovalentaria, but more detailed relationships to potential sister taxa were not resolved. Studying the internal morphology, especially the osteology, might provide new insights into the evolution of the Mugiliformes as well as help clarify their phylogenetic position within the Ovalentaria. A detailed osteology of the golden grey mullet Liza aurata is presented. The use of cleared and stained specimens allowed for a complete examination of bony and cartilaginous structures, and a 3D reconstruction from a μCT data set provided additional information on the positional relationships of the bones. Following this, the data obtained were compared with different mugilid species, particularly with the flathead grey mullet Mugil cephalus. Several differences between these species could be identified, such as the position of the basisphenoid, the shape of the hyomandibular and the composition of the branchial arches. These characters might help in understanding the evolutionary changes happening within the mugiliforms and will provide the basis to study this taxon in detail, finally allowing the reconstruction of the body plan of grey mullets.