Metazoan parasite fauna of grey mullet (Mugilidae) from the coastal waters of Vietnam.

A total of 201 fishes of the family Mugilidae caught off central Vietnam (Nha Trang Bay) were examined for parasites. Eight species of Mugilidae were identified based on cytb and 16S rRNA sequences. Protozoa were not revealed. The metazoan parasites belong to six higher taxa. Myxozoa Grassé, 1970 is represented by six genera, three of which were first found in grey mullets off Vietnam. The specimens of Monogenea Van Beneden, 1858 belong to two genera: Ligophorus Euzet et Suriano, 1977 and Metamicrocotyla Yamaguti, 1953. Twelve species of the former were recorded for the first time both in the coastal waters of Vietnam and in the East Sea. Trematoda Rudolphi, 1808 is represented by the members of the four families, of which only three species, Haplosplanchnus pachysoma (Eysenhardt, 1892), Pseudohaploporus planilizum Atopkin, Besprozvannykh, Ha, Nguyen, Nguyen & Chalenko, 2019 and Spiritestis arabii Nagaty, 1948, were identified. The latter was first recorded in the East Sea. Acanthocephala Rudolphi, 1808 specimens belong to the species Neoechinorhynchus dimorphospinus Amin & Sey, 1996. Crustacea Brünnich, 1772 are represented by five families, of which only the copepod Lernanthropsis mugilii (Shishido, 1898) and the isopod Nerocila orbignyi (Guérin, 1832) were identified. Hirudinea Savigny, 1822 were not determined to genus and species level. A total of 25 species of parasites were found in the Mugilidae fishes examined in Nha Trang Bay. In general, 96 species of parasites are currently documented in these fishes in the East Sea, 55 of which occur off Vietnam. As a result of this study, carried out off central Vietnam, 16 parasite species were recorded in grey mullet for the first time in Vietnamese coastal waters and 15 for the first time in the East Sea. Thus, extending the geographical range of the studies may result in new records of parasites in these fishes.

Helminth Communities of Common Fish Species in the Coastal Zone off Crimea: Species Composition, Diversity, and Structure.

In this paper, we analyzed the diversity and structure of helminth communities of 12 common fish species from the coastal zone of Crimea. A total of 53 helminth species were found. The total number of parasite species per host fish ranged from 3 to 18. Species richness at the infracommunity and component community levels were from 1.4-4.2 to 1.7-7, respectively. The Brillouin index for the infracommunites was 0.1-1, while the Shannon index for the component communities was 0.3-1.2. Component communities demonstrated a bi- or tri-modal distribution of the parasite prevalence and positive correlations between the prevalence and log-transformed abundance indices, thus following the "core-satellite" conception. Overall, the prevalence and abundance index of the dominant parasite in the component communities ranged from 18 to 80% and from 0.6 to 61.5 ind. per fish, respectively. The structure of the helminth component communities demonstrated good accordance with the nestedness mode where the rarest species occurred in the most diverse infracommunities, while the poorest infracommunities were composed of a few dominating species. More than two-thirds of the studied helminth species had an aggregated distribution indicating well-structured and developed communities. Our data provide a basis for further research and may be used for fish resource monitoring and management.

Life-history studies on infrapopulations of Mazocraes alosae (Monogenea) parasitising Alosa immaculata (Actinopterygii) in the northern Black and Azov Seas.

The life-history of Mazocraes alosae Hermann, 1782 on one of its hosts, the Pontic shad Alosa immaculata Bennett, is described for the first time. This anadromous fish, which occurs off the coast of the Crimea and migrates from the Black Sea to the Sea of Azov and into the River Don for spawning, was studied throughout its migration and during all seasons. It is demonstrated that the period of reproduction of this monogenean is significantly longer than that reported for the population in the Caspian Sea, lasting from April to November with a peak in April-May, and continues both in the sea and the river. Experiments showed that water salinity does not limit the development of the eggs or the hatching of the oncomiracidia. Our data reveal that the abundance of M. alosae is not determined by the size or sex of mature fish and that shad of less than two years old can also be infected with this monogenean, although less frequently than older fish. The direction of the migration of A. immaculata, either from the Black Sea to the Sea of Azov and into the rivers or in the opposite direction, does not influence the number of monogeneans present on the host. The main factor affecting the dynamics of the abundance of this monogenean is season, and, as has been indicated previously in the Caspian Sea basin, there is a synchronisation between the parasite's life-history and both the host's spawning behaviour and the duration of its migration.

Xenoligophoroides cobitis (Ergens, 1963) n. g., n. comb. (Monogenea: Ancyrocephalidae), a parasite of Gobius cobitis Pallas (Perciformes: Gobiidae) from the Mediterranean and Black seas.

Based on an integrative taxonomic approach, combining morphological characters and partial sequences of the nuclear 28S rRNA gene, a new genus and combination for the species Xenoligophoroides cobitis (Ergens, 1963) is proposed, to accommodate ancyrocephalid monogeneans, parasites on the gills of Gobius cobitis Pallas (Gobiidae) from the western Mediterranean Sea and the northern Black Sea. A morphological comparison of newly collected material with the descriptions of Ancyrocephalus cobitis Ergens, 1963 and Haliotrema cupensis Sasal, Pages & Euzet, 1998, recently synonymised and named as Haliotrema cobitis (Ergens, 1963), confirms their similarity and belonging to the same species. However, characters of this species, as the vas deferens not looping the caecal branch, the bilobed base of the male copulatory organ and the marginal hooks with an upright thumb, do not correspond to the diagnosis of Haliotrema Johnston & Tiegs, 1922. Morphologically, this species is close to members of Ligophorus Euzet & Suriano, 1977 and Kriboetrema Sarabeev, Rubtsova, Yang & Balbuena, 2013, but differs from the former in the accessory piece articulated with MCO and two prostatic reservoirs, and from the latter in the uncoiled MCO with bilobed base and the dextral vaginal pore. Moreover, all species of Ligophorus and Kriboetrema are parasites of grey mullets. A 28S rDNA-based phylogenetic analysis of sequences derived from specimens of X. cobitis from the Mediterranean and Black seas, along with sequences from several closely related genera of the Ancyrocephalidae, suggested the occurrence of a new taxonomic unit, which corresponded to the X. cobitis specimens. This finding supports the establishment of the new genus for the ancyrocephalid parasites on the gills of G. cobitis from the western Mediterranean Sea and the northern Black Sea.

Musculoskeletal and nervous systems of the attachment organ in three species of Diplectanum (Monogenea: Dactylogyroidea).

The taxonomy of Diplectanum Diesing, 1858, a genus of monopisthocotylean monogeneans, remains unsettled and needs to be revised based on new morphological criteria. Recent studies in monopisthocotyleans have shown that the muscle arrangement in the posterior attachment organ (haptor) differs between congeneric species and can be used as an additional criterion in genus-level taxonomy. To explore the possibility of using the haptoral musculature and nervous system in the taxonomy of Diplectanum, we conducted a detailed confocal-microscopy study of three species of Diplectanum (D. aculeatum Parona et Perugia, 1889, D. sciaenae van Beneden et Hesse, 1863 and D. similis Bychowsky, 1957) with phalloidin staining for muscle and indirect immunostaining for 5HT and FMRFamide. A further goal was to clarify the functional mechanics of the haptor and the role of its essential components (squamodiscs and anchors) in attachment to the host. The system of connecting bars and gaffing anchors was found to have a complex musculature consisting of 23 muscles in D. aculeatum and D. sciaenae, and 21 muscles in D. similis. The squamodiscs were shown to be operated by several groups of muscles attached primarily to the area termed the squamodisc fulcrum. Most of the haptoral musculature is identical in D. aculeatum and D. sciaenae and these species differ only in the presence of a muscle sheath around the tissue strand between the squamodiscs in D. sciaenae and in the different patterns of superficial squamodisc muscles. Diplectanum similis shows more significant differences from the other two species: besides lacking two of the haptoral muscles, it also differs in the shapes and arrangement of several other muscles. The nervous system of all three species conforms to the general pattern typical for the Dactylogyroidea and shows little variation between species.

Haptoral neuromusculature in two species of Dactylogyrus Diesing, 1850 (Monogenea: Dactylogyridae).

The taxonomy of Dactylogyrus Diesing, 1850 (Monogenea: Dactylogyridae), like that of most monopisthocotyleans, relies heavily on the morphology of sclerites of the posterior attachment organ (haptor). However, the associated neuromusculature is essentially unknown and, therefore, the aim of this study was to use confocal microscopy to examine the haptoral neuromusculature in two species of Dactylogyrus: D. amphibothrium Wagener, 1857 and D. crucifer Wagener, 1857. The monogeneans were stained with phalloidin for muscle and with antibodies for FMRFamide and 5HT and confocal reflectance microscopy was used to visualise the sclerites (i.e. anchors, marginal hooks and bars). Both species had a similar architecture of the anchoral musculature, with a pair of extrinsic muscles, two interconnecting muscles and muscles attached to the haptoral wall, anchoral openings and a connecting bar. The musculature of most marginal hooks consisted of retractors and protractors inserted on the proximal ends of the hooks. The two species differed significantly in the musculature of the accessory ventral bar: D. crucifer had a four-rayed bar with an elaborate musculature associated with the marginal hooks and D. amphibothrium had a rod-shaped bar with simple musculature. Patterns of neurotransmitter immunoreactivity (IR) in the haptor were similar in both species: RFamide-IR cells were clustered in a pair of pre-anchoral ganglia interconnected by a neurite bundle, 5HT-IR cells formed a ventral loop projecting neurites to the marginal hooks. The functional roles of haptoral muscles and patterns of neurotransmitters are discussed. In D. crucifer, the confocal reflectance microscopy revealed an additional sclerite (accessory dorsal bar) that had never been described previously in this or any other species of Dactylogyrus, suggesting that the reflected-light technique might be useful in identifying sclerites undetectable by conventional methods.

The first record of Gyrodactylus corleonis Paladini, Cable, Fioravanti, Faria & Shinn, 2010 (Monogenea: Gyrodactylidae) from the wild.

Specimens of Gyrodactylus corleonis Paladini, Cable, Fioravanti, Faria & Shinn, 2010 (Monogenea: Gyrodactylidae) were collected from the body and fins of Gobius cobitis Pallas in the western Mediterranean Sea off northwestern Sardinia. This is the first finding of this species in the wild and also represents a new host record. A morphological comparison of the new material with the type description of G. corleonis is presented. Since G. corleonis was found in the wild on representatives of the Gobiidae, a comparison with species of Gyrodactylus von Nordmann, 1832 parasitising gobies is included. The occurrence of G. corleonis on different host species and its phylogenetic relationships with gyrodactylids from sand gobies are discussed.

Lamellodiscus aff. euzeti Diamanka, Boudaya, Toguebaye & Pariselle, 2011 (Monogenea: Diplectanidae) from the gills of Cheimerius nufar (Valenciennes) (Pisces: Sparidae) collected in the Arabian Sea, with comments on the distribution, specificity and historical biogeography of Lamellodiscus spp.

Specimens of Lamellodiscus Johnston & Tiegs, 1922 (Monogenea: Diplectanidae) were collected from the gills of Cheimerius nufar (Valenciennes) (Sparidae) in the Arabian Sea. All of these parasites belonged to one and the same species, which is morphologically very close to L. euzeti Diamanka, Boudaya, Toguebaye & Pariselle, 2011. A different host, distant locality and small morphological differences compared with the original description of L. euzeti acted as a stimulus for a detailed redescription. The specimens from the Arabian Sea differ slightly in the details of the male copulatory organ (MCO) from the type-specimens of L. euzeti, which were re-examined, and from the respective drawings in its original description. Such differences include a longer inner process of the large element of the accessory piece associated with the proximal part of the copulatory tube, a longer point on the small element of the accessory piece associated with the distal part of the copulatory tube, and the presence of a smooth or slightly folded inner margin of this element rather than structures resembling spines which occur in the type-specimens of L. euzeti. Therefore, the present specimens infecting C. nufar in the Indo-Pacific may represent a different, but morphologically very similar species to the Atlantic form L. euzeti; consequently, they are recognised here as Lamellodiscus aff. euzeti. This form belongs to the 'ignoratus s. str.' subgroup of the genus. The composition of this subgroup is redefined to comprise 17 species, including L. corallinus Paperna, 1965 but excluding L. acanthopagri Roubal, 1981, and the morphology of the MCO of representatives of this group is clarified. A link between the diversity of Lamellodiscus species and the ancestral origin of present-day sparid species in the Tethys Sea is suggested. It is shown that Lamellodiscus spp. exhibit rather high levels of specificity to their hosts, since half of them parasitise only a single host species and c.90% infect closely related host species. Comparison of the levels of host-specificity of the species of this genus with other narrowly specific genera of the Dactylogyridea revealed that their estimations are comparable. The possibility of intra-host speciation within Lamellodiscus is discussed. It is shown that a co-evolutionary model is more discernible if it includes data on the occurrence of morphologically similar species from different regions and host taxa.

Microcotyle omanae n. sp. (Monogenea: Microcotylidae), a parasite of Cheimerius nufar (Valenciennes) (Sparidae) from the Arabian Sea.

Microcotyle omanae n. sp. (Monogenea: Microcotylidae) is described from the gills of Cheimerius nufar (Valenciennes) (Sparidae) from the Arabian Sea. The new species closely resembles Microcotyle arripis Sandars, 1945, M. helotes Sandars, 1944, M. caudata Goto, 1984 and M. sebastis Goto, 1984, which have also been found in the Indo-Pacific. Microcotyle omanae n. sp. differs from M. arripis, M. helotes and M. caudata by its greater number of testes, from M. arripis, M. helotes by its greater length of the genital atrium, length/width ratio of the genital atrium and length of the eggs, and from M. helotes also in greater width of the clamps, from M. caudata and M. sebastis in its greater number of clamps and additionally from M. sebastis by its smaller genital atrial spines and clamps and by the ratio between length and width of the genital atrium. Moreover, the mature specimens of the new species have greater average body length than all above mentioned species. Correlations between 15 morphometric characters and body length are analysed in the new species, and their significance for species differentiation is 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.

Descriptions of eight new species of Ligophorus 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.

Redescription of Ligophorus mediterraneus Sarabeev, Balbuena & Euzet, 2005 (Monogenea: Ancyrocephalidae) with some methodological notes.

A redescription of Ligophorus mediterraneus Sarabeev, Balbuena & Euzet, 2005, based on original material from the Black and Mediterranean Seas, is presented and new diagnostic characters for its recognition are proposed. The unlikely wide range of variation in the angle between the shaft and point of the anchors, reported for this species and for some others in the genus, is analysed, and the structure of the ventral bar in Ligophorus spp. is described and its taxonomic significance discussed.

Redescriptions of Ligophorus cephali Rubtsova, Balbuena, Sarabeev, Blasco-Costa & Euzet, 2006 and L. chabaudi Euzet & Suriano, 1977 (Monogenea: Ancyrocephalidae), with notes on the functional morphology of the copulatory organ.

Redescriptions of Ligophorus cephali Rubtsova, Balbuena, Sarabeev, Blasco-Costa & Euzet, 2006 and L. chabaudi Euzet & Suriano, 1977 based on original material from the Black Sea, the Mediterranean Sea and the Sea of Japan are presented. A comparison of samples of these two species from different regions was carried out with the aid of principal components analysis. The occurrence of L. chabaudi on Mugil cephalus in the Sea of Japan was confirmed. The functional morphology of the male copulatory organ was examined, and the use of the shape of this structure in the taxonomy of Ligophorus Euzet & Suriano, 1977 is discussed.

Ligophorus llewellyni n. sp. (Monogenea: Ancyrocephalidae) from the redlip mullet Liza haematocheilus (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.