Myxobolus acanthogobii Hoshina, 1952 and Myxobolus selari n. sp. (Myxosporea: Myxobolidae) infecting brain of commercial fishes in Terengganu, Malaysia.

Myxosporean infection in marine water fishes has drawn less attention than in freshwater fishes, which resulted in a higher taxonomic variety in freshwater in Malaysia. This study aimed to address the gap by conducting a myxosporean survey on two commercially significant marine fish species, Nemipterus furcosus (Valenciennes) (Eupercaria incertae sedis: Nemipteridae) and Selar crumenophthalmus (Bloch) (Carangiformes: Carangidae), collected from the northeastern part of peninsular Malaysia. During the examination of the organs, two distinct Myxobolus Bütschli, 1882 species were discovered in the brain tissue of these fishes, despite the absence of any observable pathological signs. The two Myxobolus species were characterized through morphometry, morphology, and analysis of partial small subunit ribosomal RNA (18S rDNA) gene. As a result, Myxobolus acanthogobii Hoshina, 1952, which infects 2.3% of N. furcosus, is synonymous with a myxobolid species commonly found in Japanese waters, based on its morphological traits, tissue tropism, and molecular diagnostics. Furthermore, a novel species, Myxobolus selari n. sp., was described, infecting the brain of one (11%) individual S. crumenophthalmus. This unique species displayed distinctive features, placing it within a well-supported subclade primarily comprising brain-infecting myxobolids. Maximum likelihood analysis further revealed the close relationships among these brain-infecting myxobolids, underscoring the significance of tissue tropism and host taxonomy for myxobolids. This study represents the initial documentation of Myxobolus species within the southern South China Sea, shedding light on the potential diversity of marine myxosporean in this region. This article was registered in the Official Register of Zoological Nomenclature (ZooBank) as urn:lsid:zoobank.org:pub:7C400E35-7CB8-4DEE-92B7-F75FF3926441.

A new myxozoan parasite, Myxobolus allami sp. n. (Myxozoa: Myxobolidae) from the intestinal wall of Sparidentex hasta (Valenciennes) in Arabian Gulf.

Myxobolus allami sp. n. is described from the intestinal wall of the silvery black porgy, Sparidentex hasta (Valenciennes), off Saudi Arabian coast of Arabian Gulf. Two of 20 examined fish were found to be infected with irregular-shaped plasmodia 3-8 mm long × 2-3 mm wide. Mature myxospores are subspherical to elliptical in the valvular view and oval in the sutural view, and are 11-13 (12) µm long, 7-8 (7.5) µm wide and 10-12 (10.8) µm thick. Spores have relatively thin valves and mostly (~ 72%) end with short caudal appendages of ~3 µm long. The spores also have two polar capsules, which are oval to elliptical and measure 5-7 (5.7) µm in length and 2-3 (2.7) µm in width. Polar filaments are coiled, with three turns. Transmission electron microscopy revealed that caudal appendages originated from the sutural edge at the posterior pole of the myxospore with density similar to that of its valves. The SSU rRNAgene sequence of the present species does not match any available sequences in GenBank. Phylogenetically, this species is sister to Myxobolus khaliji Zhang, Al-Qurausihy et Abdel-Baki, 2014 within a well-supported clade of Myxobolus-Henneguya with species infecting marine fishes. The combination of molecular data and morphological differences between this and other species of Myxobolus Bütschli, 1882 lead us to propose that the present form be established as a new species, M. allami. The present study also provides more evidence for the idea that caudal appendages cannot be reliably used to distinguish the species of the genera Myxobolus and Henneguya Thélohan, 1892.

Synopsis of the species of Myxobolus (Cnidaria, Myxozoa, Myxosporea) described between 2014 and 2020.

A synopsis of the species of Myxobolus Bütschli, 1882 (Cnidaria, Myxosporea, Myxobolidae) described from 2014 up till now is presented. It includes 122 nominal species described all over the world. For each of the species, the most relevant morphological and morphometric data, as well as data are provided related to the location in the host, type host and type locality. The GenBank accession numbers are provided whenever possible, and the spores were redrawn based on the original descriptions. The bibliography includes all the papers containing the species descriptions.

Morphological and molecular characterization of a novel Myxobolus species from the gastrointestinal tract of brown trout (Salmo trutta) in Spain.

The genus Myxobolus Bütschli, 1882 is the largest group within the class Myxosporea and includes 905 nominal species, 18 of which have been found to infect fish belonging to the family Salmonidae. In the present study, microscopic analysis enabled detection of myxospores in 43 of 613 (7.0%) gastrointestinal tracts from brown trout (Salmo trutta) captured in several rivers in the northwest of the Iberian Peninsula. Measurement of the whole myxospores, polar capsules and other morphological characteristics, together with identification of the site of infection, has led us to propose a novel salmonid-myxobolid species, Myxobolus compostellanus n. sp. Molecular analysis of the small subunit ribosomal RNA (SSU-rRNA) gene yielded the same consensus sequence of 2039 bp in 14 fish specimens. A BLAST search indicated 97.6% similarity to Myxobolus neurobius. Phylogenetic analysis revealed that M. compostellanus n. sp. is clustered with other salmonid-infecting myxobolids. The present findings contribute to the existing knowledge about the genus Myxobolus, providing both morphological and molecular data on a novel species of Myxobolus found to infect the gastrointestinal tract of salmonids, M. compostellanus n. sp. in the brown trout (S. trutta).

A new species of Myxobolus Bütschli, 1882 (Bivalvulida: Myxobolidae) infecting stratum spongiosum of the imperiled sicklefin redhorse, Moxostoma sp. (Cypriniformes: Catostomidae) from the Little Tennessee River, North Carolina, USA.

The sicklefin redhorse, Moxostoma sp. (Cypriniformes: Catostomidae), is an innominate imperiled catostomid endemic to the Hiwassee and Little Tennessee river basins, which has been restricted to a few tributaries of these systems by impoundments. During collections to propagate sicklefin redhorse for reintroduction, a myxozoan, described herein, was observed infecting sicklefin redhorse in the Little Tennessee River Basin, North Carolina. Myxobolus naylori Ksepka et Bullard sp. n. infects the stratum spongiosum covering the scales of sicklefin redhorse. Myxospores of the new species differ from all congeners by the combination of having a mucous envelope, intercapsular process, and sutural markings as well as lacking an iodinophilic vacuole in the sporoplasm. A phylogenetic analysis of the 18S rDNA gene recovered the new species in a polytomy with Myxobolus marumotoi Li et Sato, 2014 and a clade comprised of species of Myxobolus Bütschli, 1882; Thelohanellus Kudo, 1933, and Dicauda Hoffman et Walker, 1973. Histological sections of infected sicklefin redhorse skin revealed myxospores within a plasmodium in the stratum spongiosum dorsal to scales, encapsulated in collagen fibres, and associated with focal erosion of scales directly beneath the plasmodium; in some instances, the scale was perforated by the plasmodium. The specificity of the new species to sicklefin redhorse may make it a useful biological tag to differentiate sicklefin redhorse from morphologically similar species. The new species is the first parasite reported from sicklefin redhorse, a species of concern to the United States Fish and Wildlife Service. No species of Myxobolus has been reported from species of Moxostoma in the Southeast United States. As it was observed that Myxobolus minutus Rosser, Griffin, Quiniou, Alberson, Woodyard, Mischker, Greenway, Wise et Pote, 2016 is a primary junior homonym of Myxobolus minutus Nemeczek, 1911, we propose the replacement name Myxobolus diminutus (Rosser, Griffin, Quiniou, Alberson, Woodyard, Mischker, Greenway, Wise et Pote, 2016).

Necroulcerative dermatitis associated with Myxobolus dermatoulcerans n. sp. (Cnidaria: Myxobolidae) in red-bellied piranha, Pygocentrus nattereri Kner (Characiformes: Serrasalmidae), from Peru.

A group of red-bellied piranha, Pygocentrus nattereri Kner, recently imported from Peru exhibited multifocal, cutaneous ulcerations with exposure of the underlying musculature. Skin scrapes yielded moderate numbers of myxospores morphologically consistent with Myxobolus Bütschli, 1882. Myxospores from these fish were morphologically and molecularly distinct from other myxobolids infecting piranha. Myxospores are pyriform to capsular with a rounded posterior and slightly rounded to tapering anterior aspect in valvular view. Myxospore bodies are 14.3-17.8 (mean 16.1) µm long and 7.6-10.3 (mean 8.9) µm wide. Polar capsules are symmetrical, slender, elongate, and measure 7.4-10.2 (mean 9.2) µm long and 2.1-3.7 (mean 3.0) µm wide. Sequence generated for the 18S rRNA gene had no direct matches to any sequence available on GenBank but demonstrated less than 89% nucleotide similarity to various published and unpublished Myxobolus spp. from Piaractus brachypomus (Cuvier) and Colossoma macropomum (Cuvier). This paper provides the morphological and molecular characterisation of Myxobolus dermatoulcerans n. sp. from red-bellied piranha and describes associated pathological lesions.

Morphology and molecular data of two novel cnidarian myxosporean (Myxobolidae) infecting Piaractus brachypomus from the Amazon basin.

The objective of this study was reports, through morphological and small subunit ribosomal DNA (SSU rDNA) sequencing, two novel myxobolid myxosporeans infecting Piaractus brachypomus, an economicaly important Amazonian fish popularly known as "pirapitinga". Of a total of 25 specimens of P. brachypomus examined 68% had the gill filament parasitized by Henneguya tapariensis n. sp. and 16% had infection of Myxobolus arapiuns n. sp. in the pyloric cecum. The morphological analysis revealed H. tapariensis n. sp. myxospores with an ellipsoid shape and caudal process larger than the length of the body. The polar capsules of same size were elongated and occupied less than half the body. Sequencing of the SSU rDNA generated a partial sequence of 1946 bp. In M. arapiuns n. sp. the myxospores had oval-shaped body and polar capsules of the same size, occupying less than half the body. Sequencing of the SSU rDNA generated a partial sequence of 1950 bp. Phylogenetic analysis revealed a cluster according to the order/family of the host, where H. tapariensis n. sp. was grouped in a subclade with Henneguya brachypomus and Henneguya piaractus and M. arapiuns grouped in a subclade with Myxobolus colossomatis, Myxobolus matosi and Myxobolus pirapitingae.

Taxonomy, phylogeny and host-parasite interaction of two novel Myxobolus species infecting Brycon orthotaenia from the São Francisco River, Brazil.

Two new Myxobolus species were described infecting Brycon orthotaenia from the São Francisco River, in the state of Minas Gerais, Brazil. From a total of 39 B. orthotaenia collected, two specimens (5.1%) exhibited infection of the ovary and 12 specimens (30.8%) displayed infection of the liver. The plasmodia of both Myxobolus species were white and spherical measuring around 1 mm in length. The plasmodium found in the ovary showed mature myxospores, which were oval shaped from the frontal view and measured 9.2-11.0 (9.8 ± 0.4) µm in length, 5.9-6.9 (6.5 ± 0.3) µm in width and 4.6-5 (4.9 ± 0.1) µm in diameter. The two polar capsules were the same size and measured 3.9-6.2 (4.7 ± 0.5) µm in length and 1.8-2.4 (2.1 ± 0.2) µm in width. The polar tubules had 9 coils. The plasmodium found in the liver showed mature myxospores which were ellipsoidal in shape from the frontal view and measured 10.0-11.4 (10.7 ± 0.5) µm in length, 7.3-8.6 (8.1 ± 0.4) µm in width and 5.3-7.0 (6.8 ± 0.4) µm in diameter. The two polar capsules were the same size and measured 4.2-5.4 (4.9 ± 0.3) µm in length and 1.9-2.9 (2.7 ± 0.3) µm in width. The polar tubules had 8 coils. Ultrastructural analysis revealed an asynchronous sporogenesis process, with young developmental myxospore stages more often found in the periphery of the plasmodium and mature myxospores in the centre of the plasmodium. The plasmodial wall was formed by a single membrane which was not surrounded by a layer of host tissue. A thick layer of fibrous material was found in the peripheral ectoplasm close to the plasmodial wall of the plasmodium found in the ovary. Phylogenetic analysis based on the small-subunit ribosomal DNA - ssrDNA sequences and using the closest myxozoan sequences to each one of the species studied here based on previous GenBank data and Henneguya/Myxobolus/Thelohanellus species parasitizing fish from South American, revealed that the new species are grouped in a subclade together with other Myxobolus species parasitizing bryconid hosts.

Morphological and molecular characterization of myxobolids (Cnidaria, Myxozoa) infecting cypriniforms (Actinopterygii, Teleostei) endemic to the Iberian Peninsula.

The Iberian Peninsula provides a unique freshwater ecosystem for native and endemic cypriniforms to thrive. Despite cypriniforms being hosts to multiple myxobolids worldwide, little research has been performed in this geographic location. In this study, the examination of three Iberian endemic cypriniforms showed that myxosporean richness in the Iberian Peninsula is underestimated, with three new and one known myxobolid species being reported based on morphological and molecular data (SSU). Myxobolus arcasii n. sp. is described from the kidney and gonads of the "bermejuela" Achondrostoma arcasii, M. duriensis n. sp. from the gills of the Northern straight-mouth nase Pseudochondrostoma duriense, and Thelohanellus paludicus n. sp. from the intestine of the Southern Iberian spined-loach Cobitis paludica. Myxobolus pseudodispar Gorbunova, 1936 is further reported from several organs of P. duriense, and from the spleen of A. arcasii. The occurrence of M. pseudodispar in endemic Iberian species reveals that host-shift followed its co-introduction with central European leuciscids into this geographic location. Several other myxobolids originally described from barbels in central Europe have also been reported from the Iberian endemic cypriniform Luciobarbus bocagei. Nonetheless, except for M. musculi, the identification of these myxobolids in L. bocagei is here shown to be dubious and require molecular confirmation. Phylogenetic analyses reveal M. arcasii n. sp. and M. duriensis n. sp. clustering within different lineages of leuciscid-infecting species, showing that myxobolids entered Leuciscidae as hosts multiple times during their evolution. Constituting the first myxobolid reported from the subfamily Cobitinae, Thelohanellus paludicus n. sp. stands alone in the tree topology.

Morphological and molecular characterization of a new species Myxobolus gutturocola n. sp. (Myxozoa: Myxobolidae) from the throat of Hypophthalmichthys molitrix in China.

Myxobolus gutturocola n. sp. was isolated from the throat of silver carp, Hypophthalmichthys molitrix, in Chongqing, China. Myxospore valves are unsymmetrical and smooth. Mature spores are ellipsoidal in frontal view, measuring 12.5 ± 0.2 µm (n = 25) in length, 8.4 ± 0.2 µm (n = 25) in width and 7.1 ± 0.2 µm (n = 25) in thickness. Each spore has two pyriform and unequal sizes polar capsules, the large one with 5.7 ± 0.2 µm in length × 3.6 ± 0.2 µm in width and the small one with 4.6 ± 0.2 µm in length × 2.6 ± 0.1 µm in width. Polar filaments are coiled seven or eight turns in the large polar capsule and four or five turns in the small polar capsule. The coils are arranged almost perpendicularly to the longitudinal axis of the polar capsule. Morphological analysis revealed that M. gutturocola n. sp. is distinct from related species of Myxobolus Bütschli, 1882. Molecular analysis has demonstrated that its SSU rDNA sequences do not match with any available sequences in GenBank. Phylogenetic analysis of the SSU rDNA sequences indicated this species clustered in a clade composed exclusively of parasites infecting the fishes of the Leucisini lineage and most closely related to Myxobolus pavlovskii isolated from the gill filaments of silver carp in Hungary.

Morphological and molecular characterization of Myxobolus dibombensis sp. n. (Myxozoa: Myxobolidae), a parasite of the African carp Labeobarbus batesii (Teleostei: Cyprinidae) from Dibombe River, Cameroon.

Myxobolus dibombensis sp. n. (Cnidaria: Myxosporea: Bivalvulida) is described from the fins of the African carp, Labeobarbus batesii, based on morphological and molecular data. Prevalence of infection was 51.9% (67/129). Ovoid to spherical cyst-like plasmodia were found in the intrasegmental region and among the fin rays. No pathological changes were found in the fish host tissue surrounding the cyst-like plasmodia. Mature myxospores were ovoid in frontal view and lenticular in lateral view, with slightly truncated anterior and rounded posterior ends. Myxospores measured 16.8 (15.8-18.0) µm long and 11.4 (10.0-13.0) µm wide. There was a triangular intercapsular appendix measuring 3.8 (2.6-4.5) µm long. Polar capsules were ovoid and slightly unequal in size, occupying approximately one-third of the myxospore length. The larger polar capsule measured 7 (6-8) µm long and 3.6 (3-4) µm wide, while the smaller one measured 5.8 (4.8-7.0) µm long and 3 (2-4) µm wide. The larger polar capsule contained nine to 11 filament coils, whereas the smaller one contained seven to nine coils. SSU rDNA gene sequence of M. dibombensis sp. n. did not match any sequences available in the GenBank. The similarity with available Myxobolus spp. sequences ranged from 65 to 81%. The novel species clustered with M. algonquinensis, which infects the cyprinid Luxilus cornutus from Canada.

Phylogeny and comprehensive revision of mugiliform-infecting myxobolids (Myxozoa, Myxobolidae), with the morphological and molecular redescription of the cryptic species Myxobolus exiguus.

Mullets inhabit a wide range of habitats from tropical to temperate regions and play a critical role in their ecosystems. This commercially important fish group constitutes a significant source of food in several geographic regions, and the production of some species for consumption is an increasing trend. About 64 myxosporean species have been reported in mullets, some of which are cryptic, as is the case of Myxobolus exiguus, and M. muelleri. This paper provides, for the first time, a detailed and critical revision of the data available for myxobolids reported in mullets, determining the species that have bona fide mugiliform fish hosts, in accordance with the original species descriptions, the available molecular data and the currently accepted taxonomic and phylogenetic criteria. Phylogenetic analyses using Bayesian inference and maximum-likelihood methodologies suggest that the evolutionary history of myxobolids with bona fide mugiliform fish hosts reflects that of its vertebrate hosts, while reinforcing known evolutionary factors and old systematic issues of the clade of myxobolids. A comprehensive morphological, ultrastructural and molecular redescription is also provided for the cryptic species M. exiguus, from infections in the visceral peritoneum of the thinlip-grey mullet Chelon ramada in the River Minho, Portugal.

Morphological plasticity in Myxobolus Bütschli, 1882: a taxonomic dilemma case and renaming of a parasite species of the common carp.

BACKGROUND: Myxozoans are a group of cnidarian parasites, the present taxonomy of which favors a more comprehensive characterization strategy combining spore morphology, biological traits (host/organ specificity, tissue tropism), and DNA data over the classical morphology-based taxonomy. However, a systematist might again run into a taxonomic dilemma if more than two of the following exceptional cases were encountered at the same time: extensive intraspecific polymorphism, interspecific morphological similarity, identical interspecific biological traits and blurred small-subunit (SSU) rDNA-based species boundaries. In the present study, spores of a species of Myxobolus Bütschli, 1882 with two morphotypes (wide type and narrow type) were collected from the gills of common carp Cyprinus carpio Linnaeus. Confusingly, the wide type was found to be identical to Myxobolus paratoyamai Kato, Kasai, Tomochi, Li & Sato, 2017 in spore morphology and SSU rDNA sequence, which confidently suggested their conspecificity; while the narrow type, was highly similar to Myxobolus toyamai Kudo, 1917 based on spore morphology and SSU rDNA sequence and thus could not be easily classified. This discordance between wide type and narrow type has caused a taxonomic dilemma. To address this problem, a hypothesis about the conspecificity of the narrow type and M. toyamai was addressed. RESULTS: It was found that if the narrow type is conspecific with M. toyamai, it would be paradoxical for the SSU rDNA sequence of the narrow type to be more similar to M. paratoyamai (99.3%), Myxobolus acinosus Nie & Li, 1973 (98.6%) and Myxobolus longisporus Nie & Li, 1992 (98.7%) than to M. toyamai (97.6%). According to the results of the above what-if analysis, the narrow type and M. toyamai were considered to be different species. All in all, the present dual-morphotype species is estimated to be conspecific with M. paratoyamai Kato, Kasai, Tomochi, Li & Sato, 2017. Considering that this species name was preoccupied by Myxobolus paratoyamai Nie & Li, 1992, the replacement name Myxobolus pseudoacinosus nom. nov. is proposed. CONCLUSIONS: This work addresses the taxonomic dilemma in polymorphic myxozoans and demonstrates that M. pseudoacinosus is a distinct species with two morphotypes. The present study may serve as a baseline for future studies that encounter similar classification complexities.

Morphological, histological and molecular characterization of three Myxobolus species (Cnidaria: Myxosporea) from silver carp Hypophthalmichthys molitrix Valenciennes and bighead carp Hypophthalmichthys nobilis Richardson in China.

Three Myxobolus species were obtained from silver carp Hypophthalmichthys molitrix Valenciennes and bighead carp Hypophthalmichthys nobilis Richardson in China. In the present study, we supplemented their taxonomic characteristics by the morphological, histological and molecular methods. Myxobolus kiuchowensis Chen in Chen et Ma, 1998 formed small ellipsoidal plasmodia in the intestinal wall of bighead carp. Its spores appeared asymmetrical obovate in frontal view and fusiform in lateral view. Tiny mamillary protrusion in the anterior of some spores was observed. Two pyriform polar capsules were unequal. Histologically, M. kiuchowensis infected the tunica muscularis of host intestine. Myxobolus abitus Li et Nie, 1973 formed sausage-like plasmodia in the gills of silver carp. Its spores appeared oblate in frontal view and fusiform in lateral view. Two pyriform polar capsules were unequal and an obvious inter-capsule appendix was observed. Histological examination revealed that M. abitus developed in the interlamellar-epithelium of host gills. Myxobolus pavlovskii (Akhmerov, 1954) Landsberg et Lom, 1991 formed sausage-like plasmodia both in the gills of silver carp and bighead carp. Spores of M. pavlovskii were proximate oval in frontal view and fusiform in lateral view. Two pyriform polar capsules were unequal. The BLAST search indicated the SSU rDNA sequences of M. kiuchowensis and M. abitus were not identical to any sequence, however, the SSU rDNA sequences of M. pavlovskii were identical to that of M. pavlovskii recorded previously. Phylogenetic analysis showed that the present three species robustly clustered together in Cyprinid group and Asia group.

Morphological, histological and molecular characterization of Myxobolus kingchowensis and Thelohanellus cf. sinensis infecting gibel carp Carassius auratus gibelio (Bloch, 1782).

A Myxobolus species and a Thelohanellus species infecting Carassius auratus gibelio (Bloch, 1782) were redescribed by their morphological, histological and molecular characterization. In the present study, the Myxobolus species infecting the muscle was identified as Myxobolus kingchowensis Chen et Ma, 1998 by the morphological and molecular data. Histologically, mature spores of M. kingchowensis were observed in the intercellular and connective tissue of muscle, though the plasmodia were not found. In addition, scattered spores also occurred in the intercellular of haematopoietic cells, intraepithelial of the renal tubules and interior of the melano-macrophage centres. Phylogenetic analysis showed that M. kingchowensis clustered in the clade of muscle-infecting Myxobolus species, further supporting muscle as the infection site of M. kingchowensis. The present Thelohanellus species infecting the gills was identified conspecific as Thelohanellus sinensis reported in Sun (2006) (mark it as T. sinensis-Sun)based on spore morphology, biological traits (host specificity and organ specificity), and molecular data. However, compared with the original description of T. sinensis Chen et Hsieh, 1960, the present Thelohanellus species and T. sinensis-Sun both infecting the gills of gibel carp are distinguishable from the original description in the host and infection site, which made the validity of T. sinensis-Sun dubious. Due to the absence of molecular data in the original description of T. sinensis, we suggest marking the present species and T. sinensis-Sun as T. cf. sinensis to avoid the confusion until T. sinensis is obtained from the type host and type infection site.

Myxobolus okamurae sp. nov. (Myxosporea: Myxozoa) causing severe gill myxoboliosis in the cyprinid Labeo bata in a cold water wetland, Punjab (India).

During the present study on myxozoan parasites infecting gills of cyprinid carps inhabiting Ranjit Sagar Wetland, a new parasite, Myxobolus okamurae sp. nov. infecting gills of Labeo bata has been described based on morphology, histopathology and partial 18S rDNA sequencing. For M. okamurae sp. nov., hundred fish specimens were examined, out of which thirty-three had large cylindrical to round, white plasmodia in gills, each plasmodium measured 0.9-3.0 mm in diameter. The myxospore body was pyriform in shape, measuring 12.25 × 4.93 µm, with a small intercapsular process at the anterior end. The polar capsules were equal and pyriform in shape, measuring 6.06 × 1.45 µm having polar filaments forming coils up to 13-14 in number. The intensity of infection was recorded to be heavy as indicated by gill plasmodium index (GPI = 3). Sequence analysis showed that M. okamurae sp. nov. is 91% similar with M. catlae infecting gills of Catla catla from India followed by M. intimus infecting gills of Leuciscus idus from Hungary. The phylogenetic tree based on the final edited alignment (403 bp) with Maximum-Likelihood showed the high bootstrap value of 75 and formed two major clades involving M. okamurae sp. nov. with M. pendula M. catlae and M. dispar in one clade with a low bootstrap value of 23 and the rest of the species in a separate clade. The plasmodium was located in the gill lamella and typed as "intralamellar vascular type, LV3".

Myxobolus pseudowulii sp. n. (Myxozoa: Myxosporea), a new skin parasite of yellow catfish Tachysurus fulvidraco (Richardson) and redescription of Myxobolus voremkhai (Akhmerov, 1960).

Two species of Myxobolus Bütschli, 1882 were found in yellow catfish Tachysurus fulvidraco (Richardson). A species of Myxobolus infecting the gills was morphologically identified as Myxobolus voremkhai (Akhmerov, 1960) and it was characterised here with additional morphological and molecular data. The other species of Myxobolus infecting the host's skin did not conform to any known myxosporean species. It is characterised by the presence of round, black or milky white plasmodia with black spots. Myxospores are pyriform in frontal view and lemon-shaped in lateral view, measuring 12.9-16.2 µm (14.6 ± 0.7 µm) in length, 8.1-10.8 µm (9.4 ± 0.5 µm) in width, and 6.1-8.1 µm (7.0 ± 0.4 µm) in thickness. Two ampullaceous polar capsules are slightly unequal in size, larger polar capsule 7.2-9.5 µm (7.9 ± 0.4 µm) long by 3.0-3.9 µm (3.5 ± 0.2 µm) wide, smaller capsule 6.9-8.0 µm (7.4 ± 0.3 µm) long by 2.9-3.9 µm (3.4 ± 0.2 µm) wide. Polar filaments are coiled with seven to nine turns. Histologically, the plasmodia develop in the stratum spongiosum of skin dermis, resulting in epithelial cell shedding and immunological cell infiltration. Given the morphological and molecular differences between this species and other species of Myxobolus, we proposed the name of Myxobolus pseudowulii sp. n. for this parasite from the skin of yellow catfish. Interestingly, some spores of the new species possess Henneguya-like caudal appendages. Phylogenetically, M. pseudowulii sp. n. and M. voremkhai infecting yellow catfish group together in one clade with other parasites of Siluriformes, indicating that parasites clustering according to the fish host order may be an important factor affecting the evolution of species within the Myxobolus clade.

Morphological and molecular description of Myxobolus batalhensis n. sp. (Myxozoa, Myxosporea), a liver and ovary parasite of Salminus hilarii in Brazil.

Plasmodia containing myxospores belonging to the genus Myxobolus Bütschli, 1882 were found in the ovaries and liver of Salminus hilarii. Despite its economic value, this fish host has no previous reports of myxozoan infections. Herein, we describe Myxobolus batalhensis n. sp. using morphological and ultrastructural data, as well as histological and SSU rDNA molecular data. The mature myxospores were elongated, measuring in average 15.2 ± 0.8 µm in length, 8.4 ± 0.4 µm in width, and 5.1 ± 0.2 µm in thickness. Polar capsules were elongated and measured 5.3 ± 0.3 µm in length and 2.8 ± 0.3 µm in width. Polar filaments had 6-9 coils. Histopathological analysis showed coagulation necrosis associated with cell lysis as a response of the host cell to the parasite in the ovaries. No inflammatory reaction was observed in the liver, although the presence of the plasmodia caused changes in tissue structure. The phylogenetic analysis of South American myxobolid species showed M. batalhensis n. sp. as sister species of Myxobolus aureus. This is the first report of a myxozoan species parasitizing S. hilarii and the first myxozoan species described in the Batalha river.

Myxobolus sp. and Henneguya sp. (Cnidaria: Myxobolidae) natural co-infection in the kidney of Piaractus mesopotamicus (Characiformes: Serrasalmidae).

This study evaluated the myxozoan infection and histopathology of the kidney of freshwater fish Piaractus mesopotamicus from intensive fish farming in Brazil. A total of 55 fish were examined for myxozoan infection. Infected organs were processed by usual histology and stained with hematoxylin-eosin (H&E) and Ziehl-Neelsen (ZN). From the total of 55 fish analyzed, 47 (85.45%) presented myxospores, being 9.09% (5/55) only with Myxobolus sp., 5.45% (3/55) only with Henneguya sp., and 70.91% (39/55) presenting both parasites. The presence of myxospores was associated with histological alterations in both stromal and renal parenchyma. Myxospores were found mostly in the peritubular interstitial tissue and in low intensity in the glomerulus which caused nuclear hypertrophy and loss of Bowman space. An increase in the glomerular tuft and a reduction in the lumen of the collector tubules were also observed, besides the high number of melanomacrophage cells in the glomerulus. This study reports for the first time detection of myxozoan mixed infection in one organ of pacu and discuss the possible transportation of myxospores in the circulating blood.

Myxobolus taibaiensis sp. n. (Myxozoa: Myxosporea) infecting the intestinal wall of common carp Cyprinus carpio Linnaeus in China.

Myxobolus taibaiensis sp. n. was found in the inner intestinal wall of common carp, Cyprinus carpio Linnaeus, during the investigation of fish parasite fauna in Lake Taibai, located in the middle reach of the Yangtze River, China. The whitish ellipsoidal plasmodia, up to 2.9 mm long and 1.7 mm wide, developed in the circular muscle layer of the intestinal wall and produced significant compression into adjacent tissues, but no significant inflammatory responses were observed against this infection. Mature spores are oval in frontal view and lemon-like in lateral and apical view, averaging 10.2-11.2 µm (10.8 ± 0.2 µm) in length, 9.1-9.9 µm (9.6 ± 0.2 µm) in width and 6.1-6.6 µm (6.3 ± 0.1 µm) in thickness. Polar capsules are pyriform, equal in size, slightly converging anteriorly, measuring 4.4-5.4 µm (5.0 ± 0.2 µm) in length by 3.2-3.6 µm (3.4 ± 0.1 µm) in width. Polar filaments coiled with four to five turns and arranged perpendicular to the polar capsule length, measuring up to 106 µm. Myxobolus taibaiensis sp. n. is morphologically similar to Myxobolus rotundatus Achmerov, 1956 which also infects the inner wall of the intestine of common carp. However, the small subunit ribosomal DNA sequence identity was only 94%, generally beyond the intraspecies variation in the genus. Phylogenetically, this new species is sister to M. rotundatus and then clusters with M. shantungensis Hu, 1965 to form an independent common carp-infecting cluster within the Henneguya-Myxobolus clade.