Helminths of three species of White Sea fishes.

Parasitic fauna of the White Sea cod, Gadus morhua marisalbi; the navaga, Eleginus nawaga; and the shorthorn sculpin, Myoxocephalus scorpius, in the White Sea was repeatedly studied, but no large-scale parasitological surveys have been made in the recent three decades. To fill this gap, we conducted a survey of the helminths of these three fish species at the White Sea Biological Station (Karelia, Russia) of the Lomonosov Moscow State University in August 2021. The navaga (50 specimens studied) was found to be infected with 13 species of helminths; the White Sea cod (50 specimens), with 12 species; and the shorthorn sculpin (21 specimens), with 13 species. Plerocercoids of Diphyllobothrium schistochilus and third-stage juveniles of Pseudoterranova bulbosa were recorded in the White Sea for the first time. The helminth infracommunities of the navaga and the White Sea cod were closer in structure to each other than to those of the shorthorn sculpin. In general, the levels of helminth infection of the White Sea cod, the navaga, and shorthorn sculpin have been consistently high over 85 years of observations in the White Sea, but long-term trends in the abundance of some helminth species were multidirectional.

Parasites of Perccottus glenii Dybowski, 1877 (Actinopterygii: Odontobutidae) in the native and the introduced host range: Abundance-occupancy and abundance-variance relationships.

The Chinese sleeper Perccottus glenii Dybowski, 1877 is an invasive fish species rapidly expanding in Siberia and Europe. Its native range encompasses the Far East region of Russia, northeastern China and northern North Korea. We studied species composition, prevalence, mean abundance and variance of mean abundance of macroparasites of the Chinese sleeper in the native and the introduced range. The species composition of the parasite component communities differed considerably in the native and the introduced range. The frequency distributions of prevalence, mean abundance and variance of mean abundance of the parasites did not demonstrate any significant differentiation between the two parts of the host range. However, an analysis of the abundance-occupancy and the abundance-variance relationships revealed that the parasite component communities in the two parts of the host range were quite distinct. In the native range, prevalence increased faster and variance increased more slowly with the increasing abundance of the parasites than in the introduced range. These features are mostly associated with considerably increased prevalence, abundance and aggregation of the host-specific cestode Nippotaenia mogurndae in recipient water bodies as compared with the native habitats.

New record of Tylodelphys metacercariae (Diplostomidae) from Perccottus glenii (Odontobutidae) and their phylogenetic assessment.

Metacercariae of Tylodelphys sp. were found in the abdominal cavity of the Chinese sleeper (Perccottus glenii) collected in Liaoning Province and Inner Mongolia Autonomous Region of China. The sequences of the mitochondrial cox1 gene and ribosomal ITS1-5.8S rDNA-ITS2 region were obtained and used for molecular identification and phylogenetic assessment of this parasite species. Results of phylogenetic analyses based on ITS and cox1 markers showed that the metacercariae of Tylodelphys sp. ex P. glenii from China were conspecific with specimens of Tylodelphys sp. collected by Sokolov et al. (2013) from the same fish-host species captured earlier in West Siberia, Russia. The examined Tylodelphys sp. ex. P. glenii is the only member of the genus whose metacercariae parasitise the abdominal cavity of fish in northern Eurasia. Tylodelphys sp. ex P. glenii clustered with T. darbyi, T. immer, T. podicipina, and Tylodelphys sp. of Soldánová et al., 2017 based on mitochondrial DNA markers, and with T. darbyi, T. immer, T. kuerepus, and T. schreuringi using nuclear DNA markers.

Is Gymnophallus Odhner, 1900 (Trematoda: Gymnophallidae) polyphyletic? A new hypothesis based on phylogenetic position of Gymnophallus deliciosus (Olsson, 1893).

Gymnophallus deliciosus is a type species of the genus Gymnophallus. We collected this trematode species from gallbladder of Larus argentatus caught in the White Sea (Kandalaksha Bay, Chupa Inlet) and obtained the sequences of its nuclear 18S and 28S rDNA genes. Our recently obtained phylogenetic data support a sister group relationship of this species with G. choledochus. However, other species of the genus Gymnophallus-G. australis (KM246854) and G. minutus (KM268111)-do not branch with the group G. choledochus + G. deliciosus or with each other. Our study revealed that the genus Gynmnophallus is probably a polyphyletic taxon, and the genus affiliation of its representatives should be re-examined in future.

Phylogenetic position of Atriophallophorus minutus (Trematoda: Microphallidae), the type-species of the genus Atriophallophorus Deblock & Rosé, 1964, based on partial 28S rDNA gene sequence.

We found metacercariae of a microphallid trematode Atriophallophorus minutus in freshwater snails Bithynia tentaculata. In this study, we provide a morphological description of whole-mount specimens and semi-thin sections of experimentally grown adults of this species. Our morphological examination supports the idea that the adults of Atriophallophorus spp. have a ventral sucker with a sinistral interruption of the outer edge. In the 28S rDNA gene-based phylogenetic analyses, our specimens of A. minutus were grouped into Atriophallophorus spp. clade, as a sister taxon to Atriophallophorus winterbourni + Atriophallophorus sp. Analysis of the pairwise genetic distances between coI mtDNA gene sequences revealed a low divergence between the two specimens of A. minutus (1.1%) and a greater divergence (up to 16.6%) between them and A. winterbourni. Since other Atriophallophorus spp. are known to have a strict specificity to the polyvalent intermediate host, we suggest that A. minutus reported from different snail species may represent a complex of cryptic species.

Oribatid mites (Acariformes, Sarcoptiformes: Oribatida) in the gills of Salmo spp. (Actinopterygii: Salmonidae) parr - Short communication.

Live oribatid mites of the family Malaconothridae were found on Salmo spp. parr caught in the rivers of Northwest Russia. The mites were localised in the gill filaments and enclosed in connective tissue capsules. The encapsulation was accompanied by hyperplasia and displacement of the respiratory epithelium. One mite specimen was an adult female, while all the other specimens were protonymphs. The adult female and one protonymph specimen were identified as Tyrphonothrus sp. Other protonymphs could be identified only at the family level. The obtained partial 18S rDNA sequence of one protonymph was 100% identical to that of Tyrphonothrus maior (NCBI accession No. KY922215). This is the first report of living malaconothrid mites encapsulated in fish gills, and the phenomenon may assume parasitic behaviour. However, the nature of the relationship between the mites and the fish requires further investigations.

Genetic diversity of Loma acerinae (Microsporidia: Glugeida) from different fish hosts and localities - Short communication.

Loma acerinae is a xenoma-forming fish microsporidium described from common ruffe Gymnocephalus cernua (Perciformes: Percidae) and also found in Ponto-Caspian gobies (Gobiiformes: Gobiidae). This casts doubt on the strict host specificity of this parasite. The largest subunit RNA polymerase II (rpb1) was used as a genetic marker of the parasite isolated from six host species of Perciformes (G. cernua from the Baltic Sea), Atheriniformes (Atherina boyeri from the Azov Sea) and Gobiiformes (Neogobius spp. and Zosterisessor ophiocephalus from the Black Sea and Ponticola kessleri from the Caspian Sea basin). Two major rpb1 haplogroups were found with 98.5% identity between the groups. Notably, Haplogroup I was associated with Neogobius spp. samples (n = 6) only, whereas Haplogroup II included the samples from other host species (n = 7). These findings confirm the broad distribution and host range of L. acerinae, but also indicate that certain patterns of host-driven intraspecific polymorphism may exist. Furthermore, the study revealed low similarity between the ribosomal RNA gene sequences of L. acerinae and the type species, Loma morhua (as well as other species of the genus). This suggests loose genetic association within the genus, and may raise the need for the taxonomic revision of L. acerinae.

First Description of the Metacercaria of Nematostrigea serpens serpens (Nitzsch, 1819) (Trematoda, Strigeidae) and Phylogenetic Affiliation of Nematostrigea vietnamiensis Zhokhov & Mishina, 2011.

BACKGROUND: The genus Nematostrigea comprises trematodes parasitising in fish-eating birds of Europe, Africa, and North America. Their life cycle is unknown. PURPOSE: To provide the first description of metacercariae of Nematostrigea serpens serpens, a nominative subspecies of the type species of Nematostrigea, and to record metacercaria of Nematostrigea vietnamiensis, with molecular data. METHODS: Encysted metacercariae of N. serpens serpens and N. vietnamiensis were collected from fish in Russia and Vietnam, respectively, and were processed, identified, and documented using standard morphological techniques. The 28S rRNA gene of metacercariae of both species and the cox1 gene of metacercariae an earlier studied adult of N. serpens serpens were sequenced and used for phylogenetic analysis. RESULTS: Metacercaria of N. serpens serpens have a spatulate body with a foliate forebody and a short hindbody, two long lateral rectilinear pseudosuckers, and the holdfast organ with bifurcated anterior and entire posterior external lobes. The analysis of the cox1and 28S rRNA gene sequences unequivocally showed the conspecificity of metacercariae and adult stage of N. serpens serpens. Based on the 28S rRNA gene sequence-based phylogenetic analysis, the genus Nematostrigea is a sister taxon to the group of strigeid genera Cotylurus + Ichthyocotylurus. Morphological and phylogenetic data demonstrated that N. vietnamiensis does not belong to the genus Nematostrigea and is possibly be affiliated with the crassiphialine trematodes. CONCLUSION: This is the first record of metacercariae of N. serpens serpens. N. vietnamiensis, renamed Prodiplostomulum vietnamiense comb. nov., must be moved to the crassiphialine 'Prodiplostomulum'-type metacercariae group.

Phylogeny of dracunculoid nematodes (Chromadorea: Rhabditida: Spirurina: Dracunculoidea) from some Eurasian freshwater fishes.

Sets of small ribosomal DNA (SSU rDNA) and large ribosomal DNA (LSU rDNA) sequences were obtained for Philometroides moraveci Vismanis Yunchis, 1994, Philometra kotlani (Molnár, 1969), Philometra rischta Skrjabin, 1923, Philometra cf. obturans (Prenant, 1886) (Philometridae), Sinoichthyonema amuri (Garkavi, 1972), Agrachanus scardinii (Molnár, 1966), Kalmanmolnaria intestinalis (Dogiel Bychowsky, 1934) and Skrjabillanus tincae Shigin Shigina, 1958 (Skrjabillanidae). Phylogenetic analysis of SSU rDNA data shows that dracunculoid nematodes are divided into two well-supported clades designated as Clade I and Clade II, respectively. Clade I includes the type species of the genus Philonema Kuitunen-Ekbaum, 1933, some species from the family Daniconematidae Moravec Køie, 1987 and two subfamilies of skrjabillanids, Skrjabillaninae Shigin Shigina, 1958 and Esocineminae Moravec, 2006. Clade II unites species from the families Dracunculidae Stiles, 1907, Micropleuridae Baylis Daubney, 1926 and Philometridae Baylis Daubney, 1926. Within the Philometridae, there are several well-supported groups of species, one of which unites freshwater Philometra spp. from the Palearctic cyprinids, identified as P. kotlani, P rischta, P. ovata (Zeder, 1803) and P. cyprinirutili (Creplin, 1825). However, the phylogenetic relationships of most philometrids are unresolved. An analysis of partial SSU and LSU rDNA sequences indicates that there is no direct phylogenetic relationship between Agrachanus Tikhomirova, 1971 (type species Skrjabillanus scardinii Molnár, 1966) and Skrjabillanus Shigin Shigina, 1958 (type species Sk. tincae), which means that the genus Agrachanus can be resurrected. Our study confirms that Philonematinae Ivashkin, Sobolev Khromova, 1971 should be elevated to the family rank. We formally establish the family Philonematidae Ivashkin, Sobolev Khromova, 1971 stat. nov. We also suggest combining the superfamilies Dracunculoidea Stiles, 1907 and Camallanoidea Railliet Henry, 1915 into the infraorder Camallanomorpha Roberts, Janovy Nadler, 2013.

Phylogenetic position of deep-sea opecoelid digenean Tellervotrema beringi (Mamaev, 1965) (Trematoda: Opecoelidae) based on novel genetic data.

Adult opecoelid digeneans consistent with Tellervotrema beringi (Mamaev, 1965) were found in the intestine of the deep-water fish Antimora microlepis Bean collected off the northern Kuril Islands (Russia). Earlier, T. beringi was recorded only in macrourid fish. This is the first record of a species of Tellervotrema Gibson & Bray, 1982 from fishes of the family Moridae (Gadiformes). A partial 28S rRNA gene sequence was obtained for this species and was compared with the available sequences on GenBank from other opecoelids. The phylogenetic analysis did not support the current taxonomic hypothesis that the genus Tellervotrema belongs to the subfamily Podocotylinae Dollfus, 1959. Our results demonstrated that species of this genus formed a clade with those of the genera Mesobathylebouria Martin, Huston, Cutmore & Cribb, 2019 and Abyssopedunculus Martin, Huston, Cutmore & Cribb, 2019. The morphological synapomorphies of the Tellervotrema + (Mesobathylebouria + Abyssopedunculus) clade are currently unclear, and this group does not have an adequate subfamilial classification.

Amended diagnosis, validity and relationships of the genus Acrolichanus Ward, 1917 (Digenea: Allocreadiidae) based on the 28S rRNA gene, and observations on its lineage diversity.

Adult specimens of the allocreadiid trematode Acrolichanus auriculatus (Wedl, 1858) were collected from Acipenser schrenckii Brandt from the River Amur and Amur Estuary, Acipenser ruthenus L. from the Rivers Irtysh and Oka (Asian and European parts of Russia, respectively), and Acipenser fulvescens Rafinesque from Lake Winnebago, Wisconsin, USA, and used for phylogenetic analyses based on 28S rDNA sequence data. The results supported the monophyly of Acrolichanus populations from both continents and a clear separation of A. auriculatus from other allocreadiid taxa, thus supporting the validity of the genus Acrolichanus Ward, 1917. This study formally reestablishes the genus Acrolichanus. Species of this genus differ from other allocreadiids by the following combination of morphological features: triangular mouth opening with rounded posterior corners, simple ventral, dorsolateral and dorsomedian muscular lobes on the oral sucker, stout cirrus-sac, pars prostatica with dorsal pocket, and massive cirrus with conspicuous triradiate lumen when invaginated. Specimens of the trematode from North America and the Rivers Irtysh and Oka were more closely related to each other than to the specimens from River Amur and Amur Estuary. This reflects the evolutionary divergence among sturgeons into Pacific and Atlantic clades. The phylogenetic analyses also confirmed the paraphyly of the genus Crepidostomum Braun, 1900.

Helminths of Fedorov's lumpsucker Eumicrotremus fedorovi Mandrytsa, 1991 (Actinopterygii: Cyclopteridae) in the Simushir Island area (Pacific Ocean).

Fedorov's lumpsucker, Eumicrotremus fedorovi Mandrytsa, 1991, is native to the Kuril Islands area. Two sets of 69 and 30 adult specimens of E. fedorovi caught on the continental shelf and slope of Simushir Island in the Pacific Ocean were dissected and studied for macroparasites. The studied specimens were heavily infected with plerocercoids of Nybelinia surmenicola and Ascarophis pacifica. For the first time, 15 parasite taxa were recorded for E. fedorovi. None of the parasites found are solely specific to the E. fedorovi or to the representatives of the family Cyclopteridae. The infection of E. fedorovi could be dramatically different in different parts of the Simushir Island area in terms of parasite species composition and intensity of infection.

Asaccotrema vietnamiense n. gen.; n. sp. (Trematoda: Monorchioidea), a new aberrant representative of lissorchiid trematodes from the sidestripe rasbora, Rasbora paviana Tirant (Actinopterygii: Cyprinidae), Vietnam.

In this paper we erect Asaccotrema n. gen. of Lissorchiidae Magath, 1917 (Trematoda, Monorchioidea) and provide a description of Asaccotrema vietnamiense n. sp., that infects freshwater fish in Vietnam. The morphology of Asaccotrema n. gen. is significantly different from that of other lissorchiid genera due to the absence of the cirrus-sac, and position of the genital pore sublaterally, near the middle level of the esophagus. The male terminal genitalia are characterized by the presence of the seminal vesicle, which is surrounded by prostate gland-cells, a tubular pars prostatica and a long unarmed ejaculatory duct. The terminal portion of the cluster of prostate gland-cells, which surrounds pars prostatica and the distal end of seminal vesicle, is covered with a thin-walled open-ended membrane, and the other constituents of the male terminal genitalia are naked. Asaccotrema vietnamiense n. sp. specimens are found as progenetic metacercariae on the liver of the sidestripe rasbora, Rasbora paviana Tirant (Actinopterygii: Cyprinidae). A partial sequence was obtained for the 28S rRNA gene (rDNA) of A. vietnamiense n. sp. The 28S rDNA-based phylogenetic analyses support the integration of A. vietnamiense n. sp. into the Lissorchiidae clade, which also includes Lissorchis kritskyi Barnhart Powell, 1979 and Asymphylodora perccotti Besprozvanykh, Ermolenko Atopkin, 2012. This is the second record of a lissorchiid species in fish in Vietnam.

Phylogenetic analysis of the superfamily Hemiuroidea (Platyhelminthes, Neodermata: Trematoda) based on partial 28S rDNA sequences.

In the present paper, the phylogenetic relationships between genera, subfamilies and families of the Hemiuroidea are explored. Twelve new sequences of 28 rDNA and data taken from GenBank (NSBI) on 43 species affiliated to 34 genera were included in the analysis. Most of the hemiuroidean trematodes form two highly supported clades (A and B), which are sister groups to each other. Hemipera manteri joined with Gonocerca spp. with moderate statistical support. This clade is basal relative to the clades A and B. Сlade A is polytomic and contains representatives of the families Accacoeliidae, Syncoeliidae, Didymozoidae, Hirudinellidae and Sclerodistomidae, and derogenid subfamilies Derogeninae and Halipeginae. At the same time, the Syncoeliidae, Hirudinellidae and Accacoeliidae form a well-supported monophyletic group. The phylogenetic relationship between Derogeninae and Halipeginae is poorly resolved. Сlade B unites the isoparorchiid, bunocotylid, lecithasterid and hemiurid trematodes. Our data re-establishes the family Bunocotylidae, which consists of two subfamilies, Opisthadeninae and Bunocotylinae, and the Machidatrema chilostoma + Hysterolecithoides frontilatus group. The Bunocotylidae is the sister group to the Hemiuridae + Lecithasteridae group and the Isoparorchiidae is a basal relative to the representatives of these three hemiuroid families.

Phylogenetic position of the genus Gonocerca Manter, 1925 (Trematoda, Hemiuroidea), based on partial sequences of 28S rRNA gene and a reconsideration of taxonomic status of Gonocercinae Skrjabin et Guschanskaja, 1955.

Adult trematodes of the genus Gonocerca Manter, 1925, are parasites of marine fishes. Identification of the phylogenetic positions and a revision of the taxonomic status of the subfamily Gonocercinae Skrjabin et Guschanskaja, 1955 (Derogenidae) are the main purposes of this research article. Four Gonocerca species were used in the study, including the type-species G. phycidis Manter, 1925. Molecular phylogenetic analysis, based on partial sequences of 28S rRNA gene, revealed that Gonocerca spp. are phylogenetically distant from other hemiuroid trematodes, including Derogenes varicus (Müller, 1784), representative of the type-genus of the family Derogenidae. The taxonomic rank of Gonocercinae should be raised to the family level. The generic composition of the family Gonocercidae Skrjabin et Guschanskaja, 1955 stat. nov., requires further clarification as the molecular data do not support the inclusion of the genus Hemipera Nicoll, 1913, in this family.

Comparison of the developmental stages of some European allocreadiid trematode species and a clarification of their life-cycles based on ITS2 and 28S sequences.

Genetic data were used to examine the diversity in some allocreadiid trematodes. Nuclear ribosomal DNA (ITS2 and partial sequences of 5.8S and 28S) was sampled from sexual adult and 'larval' stages. From these and previous reference datasets phylogenetic trees were constructed. The results uncovered genetically distinct lineages within Bunodera luciopercae (Müller, 1776), suggesting that the two Palaearctic subspecies, B. l. luciopercae and B. l. acerinae Roitman & Sokolov, 1999, and Nearctic B. luciopercae from Perca flavescens may represent distinct species with a restricted host-specificity. Identical rDNA was revealed for the sexual adult of B. l. acerinae and 'larval' B. luciopercae described by Wisniewski (1958). An unexpected match between the rDNA sequences of adult B. l. luciopercae and 'larval' Allocreadium isoporum (sensu Wisniewski, 1958) was also detected. The adult A. isoporum (Looss, 1894) differs significantly from the 'larval' A. isoporum, the level of rDNA sequence divergence between them (8.6 % for 5.8S-ITS2-28S sequences and 6.26% for 28S) being consistent with the level expected for intergeneric variation. These results revealed the possible existence of a cryptic species complex within the nominal species B. luciopercae and a clear need for reconsideration of some of the accepted, but largely untested, tenets regarding allocreadiid life-cycles.