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Context: Myxosporean endoparasites (phylum cnidarian) are critical pathogens that affect both wild and cultured freshwater and marine water fishes globally causing huge economic losses and high mortalities., Study Objective: The present study investigated myxosporean infections in Nile tilapia and African catfish collected from the natural resources., Methods: A total of four hundred Nile tilapia with an average weight (60 ± 5 g) and two hundred African catfish with an average weight (185 ± 30 g) were collected seasonally from Qalyubia Governorate, Egypt for parasitological and molecular diagnosis of isolated myxozoan species., Results: Microscopic examination revealed Myxobolus heterosporous, Myxobolus brachysporus, Myxobolus tilapiae, and Myxobolus amieti in Nile tilapia and Henneguya suprabranchiae, and Myxobolus brachysporus in African catfish. Sequencing of 18S rDNA gene for isolated Myxozoan spp. from Nile tilapia revealed Myxobolus tilapiae deposited in GenBank under accession numbers (OR766325 and OR766326). In African catfish, the isolated Myxobolus brachysporus sequence was deposited under accession numbers (OR766327 and OR766328). Henneguya suprabranchiae was also identified in African catfish (accession. No. OR763724 and OR763433)., Conclusion: Overall, these results indicate a high prevalence of myxozoan infection in naturally inhabiting Nile tilapia and African catfish. Curiously, Henneguya suprabranchiae was detected in the digestive tract and kidneys of African catfish, which is considered a rare form., Implication: This study highlighted the importance of parasitic surveys in natural resources that impact fish production., Competing Interests: Declarations. Ethical approval: The study was conducted following the protocol involving the use of animals that was approved by the Benha University, Faculty of Veterinary Medicine Animal Care and Use Committee (BUFVTM 13-10-22). All fish-handling procedures and regulations followed the ARRIVE guidelines for Animal Care and Use. All applicable institutional and governmental rules relating to the ethical use of animals were followed. No written consent was obtained from the fishermen for used fish in the study. Consent for publication: Not applicable. Competing interests: The authors declare no competing interests., (© 2024. The Author(s).)

Proliferative gill disease (PGD), caused by the myxozoan Henneguya ictaluri , has been the most notorious parasitic gill disease in the US catfish aquaculture industry. In 2019, an unusual gill disease caused by massive burdens of another myxozoan, Henneguya exilis , was described in channel ( Ictalurus punctatus ) × blue ( Ictalurus furcatus ) hybrid catfish. Targeted metagenomic sequencing and in situ hybridization (ISH) were used to differentiate these conditions by comparing myxozoan communities involved in lesion development and disease pathogenesis between massive H. exilis infections and PGD cases. Thirty ethanol-fixed gill holobranchs from 7 cases of massive H. exilis infection in hybrid catfish were subjected to targeted amplicon sequencing of the 18S rRNA gene and compared to a targeted metagenomic data set previously generated from clinical PGD case submissions. Furthermore, serial sections of 14 formalin-fixed gill holobranchs (2 per case) were analyzed by RNAscope duplex chromogenic ISH assays targeting 8 different myxozoan species. Targeted metagenomic and ISH data were concordant, indicating myxozoan community compositions significantly differ between PGD and massive branchial henneguyosis. Although PGD cases often consist of mixed species infections, massive branchial henneguyosis consisted of nearly pure H. exilis infections. Still, H. ictaluri was identified by ISH in association with infrequent PGD lesions, suggesting coinfections occur, and some cases of massive branchial henneguyosis may contain concurrent PGD lesions contributing to morbidity. These findings establish a case definition for a putative emerging, myxozoan-induced gill disease of farm-raised catfish with a proposed condition name of massive branchial henneguyosis of catfish (MBHC)., Competing Interests: Declaration of Conflicting InterestsThe author(s) declared no potential conflicts of interest with respect to the research, authorship, and/or publication of this article.

Multivalvulidan myxosporeans (Multivalvulida) of the genera Unicapsula Davis, 1924 and Kudoa Meglitsch, 1947 are mostly causative agents of latent and imperceptible infection in marine fishes. However, they are sometimes incriminated in causing post-mortem myoliquefaction or unsightly cyst formation in commercial fish. Despite the great commercial impacts of multivalvulidan infection, the biodiversity, host range and epidemiology of multivalvulidan species remain to be explored further, including infection of alternative annelid hosts. Therefore, this study aimed to identify multivalvulidan species and their host and/or distribution records in commercial fishes in China. Multivalvulidan infection was detected in ten commercial fish species of seven families from the South and East China Seas (Northwest Pacific Ocean) and the Eastern Central Atlantic Ocean (an imported Dagetichthys lusitanicus [de Brito Capello]). Based on morphological and molecular-genetic analyses of their small and large subunit of ribosomal RNA genes, five new host and/or geographical distribution records for five fish species are presented, and three new species in five fish species are described, namely Kudoa neoscomberomori sp. n. in Scomberomorus commerson (Lacépède); Kudoa pilosa sp. n. in Helicolenus hilgendorfi (Döderlein) (type host) and Sebastiscus tertius (Barsukov et Chen); and Kudoa tumidisporica sp. n. in Photopectoralis bindus (Valenciennes) (type host) and Nuchequula nuchalis (Temminck et Schlegel). This study provides new data on multivalvulidan diversity in the ocean ecosystem.

The Amazon is the largest river basin in the world and it is home to the greatest diversity of freshwater fish in the world. Mesonauta festivus is a cichlid popularly known as flag cichlid, widely distributed throughout South America. The diversity of parasites in fish from the Amazon region is still underestimated, due to the high fishes diversity. The Myxozoa class has a universal distribution, with some specimens being pathogenic to some fish. The aim of this work was to describe a new species of Hoferellus in M. festivus. The fish were collected in the lake region, municipality of Tartarugalzinho, in the state of Amapá, Brazil. The new species was found parasitizing the urinary bladder of M. festivus. Spores were 11.5 ±1.1 (10.4-12.6) µm long and 10.9 ±1 (9.9-11.9) µm wide, and polar capsules were equally sized, measuring 4.9 ±0.5 (4.4-5.4) µm long and 3.4 ±0.9 (2.5-4.3) µm wide, with a pyriform shape, convergent with the apical region of the spore. The polar filament was wound with 5 to 6 turns. Morphological, morphometric, molecular and phylogenetic analysis proved that it is a new species of Hoferellus in the Amazon region.

Myxozoa is a class of the Phylum Cnidaria made up of endoparasites from aquatic habitats. The genus Ceratomyxa preferentially infects marine fish, with the gallbladder being the main site parasitized. This study aimed to describe a new species of Ceratomyxa found in this organ in Boulengerella cuvieri using morphological, morphometric characterization and phylogenetic analysis of 18S rDNA gene sequences. Specimens of B. cuvieri were collected, anesthetized, desensitized and biometric measurements were performed. The organs were analyzed under a stereomicroscope and fragments of internal organs were extracted for light microscopy analysis, preserved in 80% ethanol for 18S rDNA gene analysis and fixed in Davidson solution for histological processing. Free spores of Ceratomyxa were observed in the gallbladder, in plasmodia with wave-like movements, with the following dimensions: spore width (24.5 ± 0.4) µm, spore length (5.2 ± 0.3) µm, polar capsule width (1.8 ± 0.2) µm, polar capsule length (2.1 ± 0.3) µm, number of polar tubule turns (4-5) and 100% prevalence. Phylogenetic analysis confirmed that Ceratomyxa matosi n. sp. is a new species, grouped with other freshwater Ceratomyxa species from the Amazon, representing the second description of species of this genus in the state of Amapá.

The coastal waters of Vietnam are home to a wide diversity of fishes, but the parasite diversity of these potential hosts is much less well characterized. To begin addressing this knowledge gap, we carried out surveys of myxozoan parasites in fishes collected from Nha Trang Bay in Vietnam's East Sea in 2018-2019. Mugilid fishes were collected in March-April 2018, January-February 2019, and November-December 2019, and examined for myxozoans. Myxospores consistent with those of the genus Ellipsomyxa were found in the gall bladder of four mullet species, and we thoroughly characterized those from Planiliza melinoptera. Myxospores were elliptoid and devoid of striation, with a distinct sinuous suture line. Polar capsules were pyriform and oriented toward the poles of the spore. Morphological features were compared to nominal species and this species from Vietnam was distinct. Phylogenetic analysis based on partial small subunit rDNA sequence revealed that broadly, Ellipsomyxa species split into three phylogenetic lineages, and although in some branches there are groupings by host family, habitat or locality, there are no clear phylogenetic patterns. The new species we encountered in P. melinoptera had a close sister relationship with Ellipsomyxa adlardi, with both species part of a larger subclade within the Ellipsomyxa lineage. Despite this phylogenetic similarity, these species were morphologically distinct, and partial large subunit DNA sequences were only 93% similar to each other. A combination of the morphological characteristics and molecular data suggest that this is an undescribed species and we propose the name Ellipsomyxa gordeyi n. sp., Competing Interests: Declaration of competing interest The authors declare they have no conflict of interest or competing interests. ORCID. Yurakhno, VM: 0000–0002–0571-6716. Ha,VT:0000–0001-7740-954×. Whipps, CM: 0000–0001–6139-0426., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Myxozoans are microscopical parasites widely distributed in fish, with over 2,600 described species, but their actual diversity is still underestimated. Among salmonids, more than 70 myxozoan species have been identified. This study focuses on species of Chloromyxum Mingazzini, 1890 that infect salmonid kidneys, particularly C. majori Yasutake et Wood, 1957 and C. schurovi Shulman et Ieshko, 2003. Despite their similar spore morphology, they exhibit distinct host preferences, tissue affinities and geographical distributions. Chloromyxum schurovi predominantly infects the renal tubules of Salmo salar Linnaues and S. trutta Linnaeus in Europe, while C. majori targets the glomeruli of Oncorhynchus mykiss (Walbaum) and O. tshawytscha (Walbaum) in North America. The sequence data for C. majori and C. schurovi have been either missing or questionable. In our study, we examined the kidneys of two salmonid species for chloromyxid infections, using both morphological and molecular data to characterise Chloromyxum species in salmonids. The sequence of C. schurovi obtained in our study did not match the previously published parasite data. Instead, it clustered as an independent lineage sister to the Paramyxidium Freeman et Kristmundsson, 2018 clade gathering the species from various fish organs, including the urinary tract. Our findings clarified the taxonomic origin of the previous C. schurovi sequence as Myxidium giardi Cépède, 1906, highlighting the risks associated with the presence of myxozoan blood stages in the bloodstream of their fish host and the challenges of non-specific PCR amplification. We redescribe C. schurovi, thus contributing to a better understanding of the diversity and phylogeny of kidney-infecting species of Chloromyxum.

European eel, Anguilla anguilla (Linnaeus) (Elopomorpha: Anguilliformes), is a critically endangered fish of ecological and economic importance, hosting numerous parasites, including myxozoans (Cnidaria). Since its initial discovery in the kidney of European eel, Myxidium giardi Cépède, 1906 has been reported with numerous spore sizes and shapes from various tissues of multiple anguillid species. Morphological variability, wide host and tissue spectrum, and lack of sequence data raised doubts about the conspecificity of reported isolates. Subsequent studies provided 18S rDNA sequences of several isolates from anguillids and other elopiform fish, and demonstrated a split of parasite data into two distinct phylogenetic lineages, one comprising the M. giardi sequence, and the other all species infecting elopiform fishes classified under the recently established genus Paramyxidium Freeman et Kristmundsson, 2018. Myxidium giardi was, however, transferred to this genus as Paramyxidium giardi n. comb. and designated as the type species of the genus. In line with this change, the sequence originally identified as M. giardi was considered to have been incorrectly associated with this species. To shed light on the status of M. giardi originally described by Cépède (1906), we conducted microscopic and molecular examinations of various organs of 24 individuals of European eel, originating from diverse Czech habitats. Through morphometric and molecular analyses, we demonstrated that spore and polar capsule morphology, morphometry and tissue tropism of our European eel kidney parasite isolates matched the features of the original M. giardi description. Our isolates clustered in the lineage encompassing the first published M. giardi sequence. Thus, the originally described M. giardi indeed represents an existing species within the genus Myxidium Bütschli, 1882, which we formally resurrect and redescribe. Due to the morphological and molecular differences between M. giardi and P. giardi of Freeman et Kristmundsson (2018), we additionally rename the latter species as Paramyxidium freemani nom. nov.

Myxospores discovered floating free in the bile of marine fishes from the south-central coast of Vietnam were identified using morphological and molecular methods, leading to the description of 2 new species. Ceratomyxa chauvanminhi n. sp. was detected in 16% (8/50) of cultured barramundi Lates calcarifer (Bloch) specimens, and Ceratomyxa sekoi n. sp. was found in 20% (5/25) of wild largehead hairtail Trichiurus lepturus Linnaeus specimens. The spores of C. chauvanminhi n. sp. are very shallowly ovoid, slightly crescent shaped, and 11.5 ± 0.5 (10.7-12.4) μm thick, 5.8 ± 0.2 (5.4-6.1) μm long, and 5.5 ± 0.2 (5.2-5.7) μm wide. Their posterior angles are slightly concave at 158.7° ± 4.2° (151.3°-164.8°), and they possess 2 equal spherical polar capsules 2.5 ± 0.2 (2.1-2.9) μm in diameter. The spores of C. sekoi n. sp. are 5.6 ± 0.2 (5.0-6.1) μm long, 75.5 ± 4.8 (68.9-90.0) μm thick, and 5.5 ± 0.1 (5.4-5.6) μm wide, with 2 equal, slightly anterior spherical polar capsules 2.1 ± 0.2 (1.7-2.4) μm in diameter. Although C. sekoi n. sp. spores resemble those of species of MyxodavisiaZhao, Zhou, Kent, and Whipps, 2008, characterized by long tapering valves, genetic analyses distinctly place this new species within the Ceratomyxa Thélohan, 1892 lineage. This study contributes to the understanding myxosporean diversity in Vietnamese waters and highlights the difficulty associated with distinguishing between the genera Ceratomyxa and Myxodavisia., (© American Society of Parasitologists 2024.)

A synopsis of Ortholinea Shulman, 1962 (Cnidaria: Myxosporea: Ortholineidae) is presented and identifies 26 nominal species presently allocated within this genus. Species morphological and morphometric features, tissue tropism, type-host, and type-locality are provided from original descriptions. Data from subsequent redescriptions and reports is also given. Accession numbers to sequences deposited in GenBank are indicated when available, and the myxospores were redrawn based on original descriptions. The information gathered shows that Ortholinea infect a wide taxonomic variety of freshwater and marine fish. Nonetheless, the broad host specificity reported for several species is not fully supported by morphological descriptions and requires molecular corroboration. The members of this genus are coelozoic and mainly parasitize the urinary system, with few species occurring in the gallbladder. Ortholinea visakhapatnamensis is the only exception, being histozoic in the visceral peritoneum. Molecular data of the small subunit ribosomal RNA gene (SSU rDNA) is available for about one third of Ortholinea species, with genetic interspecific variation ranging between 1.65% and 29.1%. Phylogenetic analyses reveal Ortholinea to be polyphyletic, with available SSU rDNA sequences clustering within the subclades of the highly heterogenous freshwater urinary clade of the oligochaete-infecting lineage. The life cycles of two Ortholinea species have been clarified based on molecular inferences and identify triactinomyxon actinospores as counterparts, and marine oligochaetes of the family Naididae as permissive hosts to this genus., (© 2024. The Author(s).)

Background: Myxozoa is a class of cnidarian parasites that encompasses over 2,400 species. Phylogenetic relationships among myxozoans remain highly debated, owing to both a lack of informative morphological characters and a shortage of molecular markers. Mitochondrial (mt) genomes are a common marker in phylogeny and biogeography. However, only five complete myxozoan mt genomes have been sequenced: four belonging to two closely related genera, Enteromyxum and Kudoa, and one from the genus Myxobolus. Interestingly, while cytochrome oxidase genes could be identified in Enteromyxum and Kudoa, no such genes were found in Myxobolus squamalis, and another member of the Myxobolidae (Henneguya salminicola) was found to have lost its entire mt genome. To evaluate the utility of mt genomes to reconstruct myxozoan relationships and to understand if the loss of cytochrome oxidase genes is a characteristic of myxobolids, we sequenced the mt genome of five myxozoans (Myxobolus wulii, M. honghuensis, M. shantungensis, Thelohanellus kitauei and, Sphaeromyxa zaharoni) using Illumina and Oxford Nanopore platforms., Results: Unlike Enteromyxum, which possesses a partitioned mt genome, the five mt genomes were encoded on single circular chromosomes. An mt plasmid was found in M. wulii, as described previously in Kudoa iwatai. In all new myxozoan genomes, five protein-coding genes (cob, cox1, cox2, nad1, and nad5) and two rRNAs (rnl and rns) were recognized, but no tRNA. We found that Myxobolus and Thelohanellus species shared unidentified reading frames, supporting the view that these mt open reading frames are functional. Our phylogenetic reconstructions based on the five conserved mt genes agree with previously published trees based on the 18S rRNA gene., Conclusions: Our results suggest that the loss of cytochrome oxidase genes is not a characteristic of all myxobolids, the ancestral myxozoan mt genome was likely encoded on a single circular chromosome, and mt plasmids exist in a few lineages. Our findings indicate that myxozoan mt sequences are poor markers for reconstructing myxozoan phylogenetic relationships because of their fast-evolutionary rates and the abundance of repeated elements, which complicates assembly., (© 2024. The Author(s).)

Members of the genus Ortholinea are among the worldwide distributed myxozoan parasites that mainly infect marine fish. In this study, a new myxosporean species, Ortholinea hamsiensis n. sp., was isolated from the urinary bladder of European anchovy Engraulis engrasicolus collected from the Sinop coasts of the Black Sea. The prevalence and density values of infection were 1.4% and 1–5 individuals in the field of view (1 + ), respectively. Mature myxospores are subspherical with slight tapering down to the less pronounced tip in the frontal view and subspherical in the sutural view. Myxospores measured 9.1 ± 0.25 (8.8–9.9) μ m in length, 9.2 ± 0.11 (8.9–9.4) μ m in thickness, and 8.4 ± 0.33 (8.2-9.1) μ m in width. Two polar capsules equal in size measured 3.1 ± 0.11 (3.0–3.3) μ m in length and 2.7 ± 0.11 (2.6–2.9) μ m in width. The polar tubule had 3–4 coils. Along with morphological peculiarities, the results of the 18S rDNA also revealed it to be a new species for science compared to the other species of the genus. In this study, another myxosporean species O. gobiusi was also detected in round goby Neogobius melanostomus with a prevalence of infection value of 4.8% and a density of 1–5 individuals in the field of view (1 + ). The present study also provided the first data of 18S rDNA of O. gobiusi from N. melanostomus and type species of the genus O. divergens from Gobius niger and the phylogenetic relationships of these species with other Ortholinea species have been revealed.

Purpose: The present study provides the complete morphological and molecular description of two new species of myxosporeans, Ceratomyxa zancli n. sp. and Ceratomyxa cornuti n. sp. infecting the gallbladder of Zanclus cornutus from the Lakshadweep Islands, Arabian Sea., Methods: Zanclus cornutus were screened for the presence of myxosporeans, and the recovered myxospores were morphologically characterized using Nomarski Differential Interference Contrast (DIC) optics. The sequences of SSU rDNA were employed for molecular and phylogenetic studies., Results: Both the parasites exhibited a prevalence of 21% each. C. zancli n. sp. is characterized by broadly cresentic myxospores with convex anterior and slightly concave to straight posterior margins and rounded ends. Spore valves two, unequal, measured 9.6 ± 0.7 μm × 25.2 ± 1.3 μm. Polar capsules two, unequal, spherical, measured 4 ± 0.6 μm × 3.5 ± 0.6 μm. Polar filament exceptionally long and arranged irregularly. Myxospores of C. cornuti n. sp. are elongated with convex anterior and slightly concave to straight posterior margins. Spore valves two, unequal, measured 7.00 ± 0.4 μm × 26.56 ± 1.8 μm. Polar capsules spherical, unequal, measured 3.52 ± 0.2 × 3.36 ± 0.35. Molecular analysis of C. zancli n. sp. (ON818297) and C. cornuti n. sp. (ON818298) resulted in 1469 and 1491 bp long SSU rDNA sequences, respectively. Molecularly C. zancli n. sp. is close to C. diplodae and C. barnesi with 91.39% similarity, while C. cornuti n. sp. appears closer to C. robertsthomsoni with 97.46% similarity. In phylogenetic analyses, C. zancli n. sp. branched separately within the Ceratomyxa clade while C. cornuti n. sp. clustered with C. robertsthomsoni and C. thalassomae., Conclusion: Based on the differences in morphological, morphometric, molecular, and phylogenetic characteristics, as well as differences in the host and geographic location, the above two species of myxosporeans are considered novel. The study forms the first report of a species of Ceratomyxa from Z. cornutus., (© 2024. The Author(s), under exclusive licence to Springer Nature Switzerland AG.)

The substantial development of microscopic techniques and histological examination methods during the past five decades allowed for many new insights into the histology and microanatomy of Rhizostomeae. The present review focuses on new findings about histologically important structures: nerves, senses, muscles, gonads, zooxanthellae and nematocysts. Different ontogenetic stages of rhizostome species were included in the literature research, supplemented with the authors' unpublished data and figures. The overview of the research results reveals that the application of chemo- and immunohistochemical techniques have provided deeper insights into neuronal and sensory structures and their interconnections. Modern microscopic methods led to new findings on the histological gonadal organization and details of the processes of gametogenesis, fertilization, cleavage, gastrulation, and brooding. Advanced optical methods also allowed for a better understanding of Rhizostomeae-zooxanthellae associations and the morphology and function of nematocysts. Improvements in molecular biology allowed for more precise identification of zooxanthellae associated with rhizostome species. Although there has been significant progress in all of the research subjects covered here, we identify several knowledge gaps and conclude with some recommendations for future research., (Copyright © 2024. Published by Elsevier Ltd.)

Myxidium rhodei Léger, 1905 (Cnidaria: Myxozoa) is a kidney-infecting myxosporean that was originally described from the European bitterling Rhodeus amarus. Subsequently, it has been documented based on spore morphology in more than 40 other cypriniform species, with the roach Rutilus rutilus being the most commonly reported host. This study introduces the first comprehensive data assessment of M. rhodei, conducted through morphological, ecological and molecular methods. The morphological and phylogenetic analyses of SSU rDNA sequences of Myxidium isolates obtained from European bitterling and roach did not support parasite conspecificity from these fish. In fact, the roach-infecting isolates represent three distinct parasite species. The first two, M. rutili n. sp. and M. rutilusi n. sp., are closely related cryptic species clustering with other myxosporeans in the freshwater urinary clade, sharing the same tissue tropism. The third one, M. batuevae n. sp., previously assigned to M. cf. rhodei, clustered in the hepatic biliary clade sister to bitterling-infecting M. rhodei. Our examination of diverse cypriniform fishes, coupled with molecular and morphological analyses, allowed us to untangle the cryptic species nature of M. rhodei and discover the existence of novel species. This underscores the largely undiscovered range of myxozoan diversity and highlights the need to incorporate sequence data in diagnosing novel species., (© D. Baiko et al., published by EDP Sciences, 2024.)

Myxosporean parasites Kudoa spp. have been reported in several marine fish species worldwide. However, little is known about the contamination of these parasites in raw fish in Southeast Asia, where the consumption demand of uncooked fish is increasing. In 2019, the occurrence of several cases of raw yellowfin tuna (Thunnus albacares) obtained from retail shops with the presence of unknown white, nodular cysts within the musculature have raised public health concerns for the consumption of raw marine fish in Vietnam. Microscopic examination revealed numerous myxospores with the quadratic shape of the Kudoidae. Morphologically, stained spores detected in this study are suspected to Kudoa thunni. To confirm the suspected Kudoa species, further examination of the 18S small-subunit (SSU) was conducted and the results of nucleotide sequence analysis obtained from nodular cysts revealed 99.18-100% identity to that of Kudoa thunni sequences available in GenBank. Detection of K. thunni infection in tuna in Southeast Asia highlights the need for appropriate surveillance and control measures to ensure high quality standards and safety on raw fish production and consumption., (Copyright © 2022 Elsevier B.V. All rights reserved.)

The present study describes Hoferellus jutubensis n. sp., a myxozoan parasite found in the urinary bladder of the driftwood catfish Ageneiosus inermis, captured on Jutuba Island in the state of Pará, northern Brazil. A total of 30 A. inermis specimens were examined, of which 26 (86.7%) had myxospores and polysporic plasmodia of varying shapes and sizes dispersed in the lumen of the urinary bladder, either floating freely or attached to the epithelium. In the apical view, the myxospores of Hoferellus jutubensis n. sp. are rounded, 6.1 ± 0.2 (5.7-6.3) μm long and 5.5 ± 0.3 (5.2-6.0) μm wide, with two sub-spherical polar capsules, equal in size and shape, 2.5 ± 0.2 (2.3-2.7) μm long and 1.7 ± 0.2 (1.4-2.2) μm wide. The phylogenetic analysis of a partial sequence of the SSU rDNA gene, indicated that the new species is the sister taxon of Hoferellus azevedoi, with these two species forming a Brazilian lineage of Hoferellus. The comparison of the morphological and molecular data with those of the existing members of the genus confirmed the species status of Hoferellus jutubensis n. sp., which adds one further Hoferellus taxon to the known myxosporean diversity of the Amazon basin., (Copyright © 2021 Elsevier B.V. All rights reserved.)

Kudoa ocellatus n. sp. was found in the musculature of Astronotus ocelattus (Agassiz, 1831) from the Arari River on Marajó Island in Pará, Brazil. The new species forms pseudocysts in the epaxial and hypaxial musculature composed of various spores that are pseudoquadrate in the apical view. In the lateral view, the spores were triangular or pyramidal. In the lateral view, the spores were 46 ± 0.11 μm (4.5-4.8) in length and 6.6 ± 0.3 μm (6.2-7.2) in width, with four pyriform polar capsules of equal size that measured 2.0 ± 0.16 μm (1.8-2.2) in length and 1.5 ± 0.18 μm (1.3-1.8) in width. Based on the partial (1418 bps) sequence of the SSU rDNA gene, Kudoa ocellatus n. sp. was distinct from all the other Kudoa species deposited in GenBank. The phylogenetic Bayesian Inference and P distance placed the new species together with the other Kudoa species that parasitize freshwater Amazonian fish. The morphological evidence, together with the SSU rDNA gene sequence, supported the description of Kudoa ocellatus n. sp., a distinct new species of the genus, which parasitizes a freshwater Amazonian cichlid., (Copyright © 2021 Elsevier B.V. All rights reserved.)

We described two novel myxozoan parasite species Ceratomyxa argentina n. sp. and Ceratomyxa raneyae n. sp. from the gall bladder of Raneya brasiliensis (Kaup) from the Patagonian coast of Argentina. Both species can be distinguished from other ceratomyxids by myxospore and polar capsule (nematocyst) morphology and morphometry, fish host and geographic locality. Phylogenetic reconstruction using ssrDNA gene sequences showed that the two new species are placed in a long-branching ceratomyxid clade which also include Ceratomyxa appendiculata Thélohan, 1892, Ceratomyxa anko Freeman, Yokoyama and Ogawa, 2008, Ceratomyxa pantherini Gunter, Burger and Adlard, 2010 and Pseudoalataspora kovalevae Kalavati, MacKenzie, Collins, Hemmingsen and Brickle, 2013. This study documents additional biodiversity of marine myxozoans in the South Atlantic, a region still largely unexplored for this group of parasitic cnidarians., (Copyright © 2021 Elsevier B.V. All rights reserved.)

Myxosporean species in the genus Cystodiscus are parasites of amphibians and have been reported from several continents. Typically used for the identification of myxozoans, the spores produced by these species are similar to one another, possessing 2 polar capsules and being ovoid. The number of transverse depressions on the spore can be useful for delineating species, but these can sometimes be difficult to distinguish. In North America, Cystodiscus serotinus and Cystodiscus melleni have been described, and for C. serotinus in particular, numerous reports and a wide range of hosts have been associated with this species. Given the challenges of identifying some of these species, we questioned whether all encounters of Cystodiscus species can be attributed to these 2 described species, or if there may be additional undescribed species or cryptic species. Over 7 yr, 383 amphibians representing 13 species of toads, frogs, and salamanders were collected from sites in Oklahoma and Arkansas. Cystodiscus infections were found in 56 individuals (14.6%). Tissues from these infected individuals were preserved in alcohol for genetic analysis. The small subunit (SSU) and large subunit (LSU) ribosomal RNA genes were partially sequenced and analyzed phylogenetically. Nine distinct SSU sequence types and 7 distinct LSU sequence types were identified. Phylogenetically, sequence types were attributable to C. serotinus, C. melleni, Cystodiscus axonis, and an undescribed species. For the previously described species, there were multiple SSU sequence types: 4 for C. serotinus and 2 for both C. melleni and C. axonis. Phylogenetic patterns were similar for the LSU sequence analysis using a shorter sequence than the SSU, and we propose that the LSU is useful for initial barcoding of Cystodiscus species in any future surveys. In our qualitative assessment of sequence types compared to geography and host species, SSU types C1 and C2 (C. axonis) were only found in Union County, Arkansas, and McCurtain County, Oklahoma, respectively. Also, salamanders were only infected with SSU types B or D (C. melleni), and type B was only found in salamanders. Our finding of C. axonis in North America is notable because this species was described in Australia and is associated with host pathology. Our work reveals that there are cryptic species of Cystodiscus in the United States, one of which may be a pathogen, highlighting the importance of genetic analysis for future surveys of these species., (© American Society of Parasitologists 2021.)

Myxosporeans are microscopic cnidarians associated with severe diseases in aquaculture and wild fish populations. This group of parasitic cnidarians thus warrants close attention concerning its potential impact on susceptible fish stocks. At present, little is known about this group of parasites infecting anguillid eels. From myxospore specimens collected from a freshwater eel (Anguilla marmorata) in the Solomon Islands, we describe a new species belonging to the genus Myxobolus based on an integrative taxonomic analysis of morphological, biological traits and molecular data. Furthermore, we determined the phylogenetic position and relationships of this species among other platysporine myxosporeans. Molecular phylogenetic assessment of small subunit ribosomal DNA showed that the species clusters together with Myxobolus portucalensis and Echinactinomyxon type 5 Özer, Wootten and Shinn, 2002, in a well-supported subclade. This is the first report of a myxosporean parasite infecting fish from the Solomon Islands., (Copyright © 2021 Elsevier B.V. All rights reserved.)

The present study describes a new coelozoic, eukaryotic microparasite of the genus Ellipsomyxa Køie, 2003 (Ceratomyxidae: Myxozoa) found parasitizing the gallbladder of Satanoperca jurupari Heckel, 1840 collected in the Curiaú River Environmental Protection Area in Macapá, Amapá state, Brazil. The fish were collected using mesh cast net. The gallbladders were examined, preserved in 80% alcohol for molecular analysis (SSU rDNA gene), and fixed in Davidson for histological slide preparation. The new parasite had a prevalence of 81% in the gallbladder, asymmetric plasmodia, irregular free spores in the bladder fluid, with no cyst formation. The spores are elliptical, with characteristics of the genus Ellipsomyxa, and they had a mean length of 10.11 (8.56-10.5) μm, mean width of 7.81 (5.96-9.56) μm, and thick walls. The polar capsules are sub-spherical in shape, slightly asymmetrical, with a mean length of 3.12 (2.31-3.99) μm and mean width of 2.5 (2.22-2.95) μm, containing polar filament with five or six coils perpendicular to the longitudinal axis of the capsule. The Bayesian Inference assigned the new species to a subclade formed by a lineage of Ellipsomyxa species from the Amazon region. Ellipsomyxa tucujuensis n. sp. is the sixth species of this genus described in fish from the Amazon region, and the first for the state of Amapá., (Copyright © 2021. Published by Elsevier B.V.)

Myxozoans are microscopic cnidarians that mainly parasitize fishes. The present study aimed to describe a new myxozoan parasite from the gills of Boulengerella cuvieri (Spix and Agassiz, 1829) by morphological and molecular analysis. The fish was collected in 2019 at the Pindaíba River, municipality of Cocalinho, Mato Grosso State, Brazil. Whitish and circular plasmodia were found in the primary gill filaments, occupying an intralamellar position, with an average of 0.5 mm in diameter. Henneguya Thélohan, 1892 myxospores found inside the plasmodia were elongated and ellipsoidal, consisting of two long and elliptical shell valves with two long, tapering caudal appendages. Morphometric measurements revealed a total spore length of 36.1 ± 2.0 μm; spore body length of 12.8 ± 0.5 μm; spore width of 4.9 ± 0.3 μm; tail length of 23.3 ± 1.6 μm; capsule length of 7.2 ± 0.4 μm; capsule width of 1.5 ± 0.2 μm; and 10 coils in the polar filament. Phylogenetic analysis showed that the isolates from this study were grouped into the main-clade of freshwater fishes, within a group of species parasitizing fishes from Brazil. Intergenotypic difference ranged from 23%-25.9% compared with other Brazilian myxozoan isolates. Using molecular and morphological characterization, this parasite was identified as a new species of the genus Henneguya., (Copyright © 2021 Elsevier B.V. All rights reserved.)

Turkey has more than 200 endemic freshwater fish species, one of which is the Ankara nase, Chondrostoma angorense Elvira, 1987 (Cypriniformes: Leuciscidae), a food fish in northern Turkey. Like most endemic fish species in Turkey, its myxosporean parasite fauna (Cnidaria: Myxosporea) are not yet described. We surveyed twenty C. angorense from Lâdik Lake in northern Turkey, and identified two myxosporean parasites from gills of these fish: Myxobolus arrabonensis Cech, Borzák, Molnár, Székely, 2015, and a co-infection of a novel species, Myxobolus polati sp. nov. We characterized both infections based on myxospore morphology, morphometry, tissue tropism, small subunit ribosomal DNA sequence and phylogenetic analysis. Plasmodia of both species were observed in gills, but had distinct tropism: M. arrabonensis is an intrafilamental vascular type, and M. polati sp. nov. is an intralamellar vascular type. We identified M. arrabonensis on the basis of myxospore characters and 100% similarity to the type DNA sequence from the closely-related host C. nasus. The small subunit ribosomal DNA sequence of M. polati sp. nov. (1946 base pairs; GenBank Accession number MH392318) had a maximum similarity of 98% with any Myxobolus sp. from other Eurasian cypriniforms. Phylogenetic analysis revealed that M. polati sp. nov. is most closely related to gill-infecting Myxobolus diversicapsularis from Rutilus rutilus (L.). The present study is the first record of myxosporean species infecting C. angorense comprising a novel species, M. polati sp. nov. and a known species M. arrabonensis., (Copyright © 2021 Elsevier B.V. All rights reserved.)

Characterising myxozoan taxa parasitising fish hosts in catfish aquaculture ponds is crucial to understanding myxozoan community dynamics in these diverse and complex ecological systems. This work investigated the myxozoan fauna of the western mosquitofish, Gambusia affinis, a common, incidental species found in catfish aquaculture ponds in the southeastern United States. 598 fish were sampled in May of 2018 and 2019 from the pond facility of the Thad Cochran National Warmwater Aquaculture Center in Stoneville, Mississippi, USA. Fish were examined microscopically using wet mount preparations of fresh tissue and histology for myxozoans. 18S rRNA gene sequences were amplified from myxospores obtained at necropsy. Updated morphologic, histologic, and 18S rRNA gene sequence features are provided for Henneguya gambusi, Myxobolus pharyngeus, and Myxidium phyllium. Two potentially novel myxozoans were observed during this survey, an undocumented Myxobolus sp. associated with chondrolysis of bones throughout the body and a putative Myxobilatus sp. observed histologically in the renal tubules, ureters, and urinary bladder. However, inadequate samples were obtained for proper species descriptions. Lastly, the life cycle of M. pharyngeus, which is thought to utilize the oligochaete worm Dero digitata as their definitive host, was putatively confirmed by 18S rRNA sequence matching to actinospore stages from oligochaetes in catfish ponds in Mississippi. This work provides novel and expanded morphologic, histologic, molecular and biologic data of five myxozoan parasites of G. affinis, expanding our knowledge of myxozoan diversity in catfish aquaculture ponds., (© 2021. The Author(s), under exclusive licence to Springer Nature B.V.)

Coelozoic parasites of the family Myxidiidae were observed in fish of the order Characiformes captured in the middle Tocantins River, Maranhão, Brazil, within the transition between the Cerrado savanna biome and the eastern extreme of the Brazilian Amazon Forest. The analysis of the morphological characteristics of the parasites, complemented with a Bayesian phylogenetic analysis, supported the description of new specie Myxidium imperatrizensis n. sp., found parasitising the gallbladder of specimens of the fish Triportheus angulatus. The study is also only the second report of the occurrence of Myxidium parasites in fish of the family Triportheidae in Brazilian rivers. The results of the present study expand the known distribution of the genera Myxidium the basin of the Tocantins River, the largest hydrographic basin located entirely within Brazil, which encompasses parts of both the Cerrado and Amazon biomes., (Copyright © 2021 Elsevier B.V. All rights reserved.)

An examination of 18 fishes caught in the South China Sea detected two Unicapsula spp. in the myofibers of the trunk muscles of carangid fishes: Unicapsula aequilobata n. sp. in the Japanese scad, Decapterus maruadsi, and Unicapsula seriolae in the yellowstripe scad, Selaroides leptolepis. They formed thin filamentous pseudocysts of 0.9-2.0 (mean 1.4) mm by 0.03-0.06 (0.04) mm (n = 5) and 0.9-3.4 (2.1) mm by 0.02-0.05 (0.04) mm (n = 12), respectively. Myxospores of U. aequilobata n. sp. are composed of three equal shell valves and measured 6.7-8.5 (7.3) μm in length and 7.1-8.8 (7.6) μm in width, and contained a prominent polar capsule (PC) 3.2-3.8 (3.6) μm in diameter (n = 18) and two rudimentary PCs. A nucleotide sequence (5127 bp) of the ribosomal RNA gene (rDNA) array was obtained for the genetic characterization of this new species. Based on morphological and phylogenetic criteria, we erect U. aequilobata n. sp. as the sixteenth species in the genus Unicapsula. Nucleotide sequences of the 18S and 28S rDNA obtained from U. seriolae from the yellowstripe scad were almost identical (99.6-100% or 99.0-99.6%, respectively) to those from fish found in the seawaters around Australia and Japan. Consequently, this is a new host and geographical distribution records for U. seriolae. In addition, we illustrated the predicted secondary structure of the available 5.8S rDNA sequences of multivalvulid species, including those obtained from U. aequilobata n. sp., to assess the significance of interspecific nucleotide variations in this short rDNA unit.

A new species of Ceratomyxa infecting the gallbladder of the marine ornamental fish Acanthurus xanthopterus collected from the Vizhinjam coast of Kerala is described. The parasite exhibited a prevalence of 100%. Mature spores recovered from the gallbladder were slightly crescentic with rounded lateral extremities and possessed convex anterior and slightly concave to straight posterior margins. Spore valves two, equal, joined by a straight and prominent suture. Myxospores measured 5.5 ± 0.6 μm in length and 15.9 ± 2.3 μm in thickness. Polar capsules two, equal, spherical, positioned anteriorly on either sides of the suture, 2.3 ± 0.2 μm long and 2.2 ± 0.2 μm wide. Polar filament with four to five coils, 21.2 ± 0.6 μm when extruded. Posterior angle 173.6 ± 5.2°. Early sporogonic stages and monosporic, disporic, and multisporic plasmodial stages were spherical to irregular in shape, with or without filopodia. Histopathologic analysis revealed that spores and developing stages were attached to the gallbladder wall as well as found free in the lumen. Morphologic and morphometric comparison of the present parasite with known species of Ceratomyxa indicated significant differences. In molecular and phylogenetic analyses, the present myxosporean revealed high divergence with related forms and occupied an independent position within the Ceratomyxa clade with high nodal support. Considering the morphological, morphometric, molecular, and phylogenetic dissimilarities with the previously described species of Ceratomyxa and the differences in host and geographic locations, the present species of myxosporean is treated as new and is named Ceratomyxa xanthopteri n. sp.

Recent anecdotal reports from seafood processors in eastern Australia have described an increased occurrence of post-mortem myoliquefaction ('jellymeat') in broadbill swordfish Xiphias gladius, and macroscopic cysts throughout the musculature of yellowfin tuna Thunnus albacares. A genus of parasitic cnidarians, Kudoa (Myxosporea, Multivalvulida), species of which are known to occur in economically important wild-caught fish species globally, can cause similar quality-deterioration issues. However, Kudoa sp. epizootiology within commercially harvested, high-value fish caught within Australia is poorly understood, despite the parasite's economic importance. To determine the causative agent responsible for the observed quality deterioration in swordfish and yellowfin tuna, muscle-tissue samples from seafood processors in Mooloolaba, Australia, collected from October 2019-February 2020, were examined for parasitic infection. Kudoid myxospores were identified from both hosts and were subquadrate in shape, with four equal-sized polar capsules. The SSU rDNA sequences from both fish shared > 99% identity to Kudoa species. Kudoa musculoliquefaciens was isolated from 87.1% of swordfish sampled, suggesting that it is a widespread parasite in swordfish from the southwest Pacific Ocean. This study provides the first molecular and morphological characterisation of Kudoa thunni in yellowfin tuna and K. musculoliquefaciens in swordfish harvested from the waters of eastern Australia, expanding the geographical distribution of K. thunni and K. musculoliquefaciens to include the Coral and Tasman Seas. We demonstrate that not all infected swordfish progress to jellymeat, show the usefulness of molecular tools for reliably identifying infection by Kudoa spp., and add to the overall knowledge of kudoid epizootiology in wild-caught fish.

Chloromyxum squali Gleeson and Adlard, 2012 originally described from Squalus acanthias Linnaeus, 1758 is reported for the first time from the gallbladder of Squalus blainville (Risso, 1827) caught from the eastern coast of Tunisia. In the present study, this chloromyxid was described based on morphological and ultrastructural features combined with molecular analysis of 18S rDNA. Young plasmodia were found attached to the gallbladder, while mature plasmodia and myxospores were observed floating free in the bile. Mature plasmodia were polysporic, subspherical in shape, measured 97.8 ± 0.5 µm long and 63.4 ± 0.4 µm wide. Mature myxospores were ovoid with a pointed anterior end, measuring 10.2 ± 0.5 µm long and 8.3 ± 0.5 µm wide. Two asymmetrical shell valves adhered together along an S-shaped suture line. Each valve has 5-7 elevated surface ridges parallel to suture line. A bundle of long caudal filaments extended from the basal end of shell valves. Four pyriform polar capsules equal in size, measuring 3.1 ± 0.4 µm long and 2.5 ± 0.3 µm wide, were situated at the same level in the anterior pole of the myxospore, each with a polar filament coiled in 7-8 turns. Pairwise comparisons among the SSU rDNA sequences revealed significant similarity between Chloromyxum squali infecting S. acanthias with the sequence obtained in this study. Phylogenetic analysis revealed that C. squali clustered in the clade of Chloromyxum species infecting the gallbladder of marine Chondrichthyes. Chloromyxum squali showed a seasonal variation of prevalence with significantly higher prevalence noted in summer and in autumn and absence of infection in winter.

Myxozoans are highly diverse and globally distributed cnidarian endoparasites in freshwater and marine habitats. They have adopted a heteroxenous life cycle, including invertebrate and fish hosts, and have been associated with diseases in aquaculture and wild fish stocks. Despite their importance, genomic resources of myxozoans have proven difficult to obtain due to their miniaturized and derived genome character and close associations with fish tissues. The first 'omic' datasets have now become the main resource for a better understanding of host-parasite interactions, virulence, and diversity, but also the evolutionary history of myxozoans. In this review, we discuss recent genomic advances in the field and outline outstanding questions to be answered with continuous and improved efforts of generating myxozoan genomic data., Competing Interests: Declaration of Interests The authors have no interests to declare., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Henneguya Thélohan, 1892 is the second most species rich genus of myxozoans, with reports from freshwater and marine fish worldwide. In the Great Lakes region of North America, muskellunge Esox masquinongy is an important game fish species that serves as an apex predator in the ecosystems of many inland lakes. The myxozoan fauna of esocid fish, especially muskellunge, remains largely understudied. During fish health assessments, muskellunge were examined for parasitic infections and myxozoan pseudocysts were observed on gill clip wet mounts. When ruptured under pressure, the intralamellar pseudocysts released thousands of myxospores consistent with those of the genus Henneguya. The myxospores were 67.3-96.6 (79.1 ± 5.9) µm in total length. The spore body was 18.6-22.6 (20.9 ± 1.0) µm × 5.4-6.9 (6.3 ± 0.4) µm in valvular view and 3.5-4.0 (3.8 ± 0.3) µm wide in sutural view. The two pyriform polar capsules positioned at the anterior of the spore body were 6.4-7.7 (7.0 ± 0.4) µm × 1.8-2.1 (2.0 ± 0.1) µm and each contained a tightly coiled polar filament with 9-10 turns. Two tapering caudal processes extended from the posterior of the spore body and were 47.3-75.6 (58.3 ± 5.8) µm in length. Histologically, large intralamellar polysporic plasmodia were surrounded by plump pillar cells and a distinct layer of plasma. Mild inflammation was present peripherally, with small numbers of necrotic germinative cells and intraplasmodial phagocytes internally. Ribosomal 18S rRNA gene sequence data were obtained from three gill pseudocysts. The three ~2000-bp sequences were identical, but shared no significant similarity with any publicly available sequence data. Phylogenetic analyses demonstrated sequence data from this Henneguya fell within a well-supported clade of Henneguya spp. reported from northern pike Esox lucius in Europe. Based on the distinct morphological, histological and molecular data, this species is designated as Henneguya michiganensis n. sp. from muskellunge in Michigan, USA.

This study undertook the first investigation of malacosporean infections in Neotropical fish. We used polymerase chain reaction detection with a primer set generally targeting known malacosporeans to assay for infection in the kidney of 146 fish in 21 species belonging to 12 families collected from two areas in the Amazon Basin. Infections were found in 13 fish variously belonging to seven species in six families and included the first identification of a malacosporean infection in cartilaginous fish (a freshwater stingray). Based on ssrDNA, all infections represented a single Buddenbrockia species (Buddenbrockia sp. E) that demonstrates an exceptionally broad range of fish species infected, and countered our expectations of high Neotropical malacosporean diversity. Infections were characterized at varying and often high prevalences in fish species but sample sizes were small. Ascertaining whether highly divergent malacosporeans have not been detected by current primers, and more comprehensive sampling may reveal whether malacosporeans are truly as species poor in the Amazon Basin as present data suggest. Our results prompt speculations about evolutionary scenarios including introduction via marine incursions and patterns of host use over time.

A novel myxozoan parasite is identified and described from mudskipper, Boleophthalmus dussumieri, collected from a brackishwater ecosystem in Maharashtra, India. Ellipsomyxa boleophthalmi sp. nov. was found in the gallbladder of 58 of 60 fish examined (96.7%). The parasite formed disporous plasmodia that varied in size and shape, and the thin-walled, ellipsoidal and elongated myxospores measured 9.0-10.7 × 6.0-7.8 μm. The two, spherical polar capsules measured 2.7 μm in diameter and enclosed 3-4 coils of polar tubules. Histological observations of infected gallbladder revealed the attachment of disporous plasmodial stages of the parasite to the gallbladder wall with fine pseudopodia. Under the scanning electron microscope (SEM), the myxospores showed a distinct central sutural line and two distinct depressions on the opposite sides at the openings of polar capsules. SEM also revealed the engulfment of microvilli of gallbladder wall by pseudopodia of the plasmodial stages. Analysis of the partial fragment of the SSU rDNA region (1386 bp) showed less than 98% sequence similarity with the other reported Ellipsomyxa spp. In the phylogenetic tree, the present species formed as a distinct subclade within the major clade of Ellipsomyxa spp. The unique morphological and morphometric features of the myxospore, together with the molecular analysis, allowed us to conclude that the present myxozoan is a new species and is named Ellipsomyxa boleophthalmi sp. nov., after the generic name of the host. This is the first report on the occurrence of the genus Ellipsomyxa in B. dussumieri.

During a survey of myxosporean parasites of marine fish in the coastal region of Vietnam, a species of the genus Henneguya (Myxosporea: Bivalvulida) was found in the gill of yellowfin seabream Acanthopagrus latus (Perciformes: Sparidae). White and oval cysts, measuring 145-220 μm in diameter, were detected in the gill lamellae of 4 of 15 fish examined (26.7%). Mature myxospores were elongate, with smooth valves, two similar polar capsules, and having the following dimensions: spore body length 9.9 ± 0.5 (8.9-12.5) μm, body width 6.7 ± 0.3 (6.1-7.6) μm, thickness 5.1 ± 0.2 (4.8-5.4) μm, caudal appendage length 10.0 ± 1 (8.3-11.6) μm, and total myxospore length 19.3 ± 1.4 (16.5-21.5) μm. The polar capsules were ovoid, measuring 3.2 ± 0.2 (2.8-3.9) μm long and 1.9 ± 0.2 (1.5-2.3) μm wide. Each polar capsule has a polar filament with 4-5 coils. Histological analysis revealed plasmodia in the connective tissues of the gill lamellae, but inflammation and deformation of the gills were not observed. In the phylogenetic tree reconstructed from the small subunit ribosomal DNA (SSU rDNA), sequences of the Henneguya specimens found in this study form a distinct branch. Morphological characteristics and molecular data identified a new species, namely Henneguya lata n. sp.

Commercial marine fishes caught locally in East Java, Indonesia, were examined for multivalvulid myxosporeans (Cnidaria: Myxozoa: Myxosporea). Plasmodia of Unicapsula pyramidata were detected in the trunk muscle of two fork-tailed threadfin breams (Nemipterus furcosus). Genetic comparisons of this sample to those collected in the Australian Coral Sea and South China Sea showed few nucleotide substitutions in the small subunit and large subunit ribosomal RNA gene (rDNA) with the species isolated in the Australian Coral Sea and South China Sea. Pseudocysts of two new Kudoa spp. with four shell valves and polar capsules were found in the trunk muscle of two shrimp scads Alepes djedaba and two flathead grey mullets Mugil cephalus. Kudoa javaensis n. sp. myxospores isolated from the shrimp scad were 5.1-7.2 (mean 6.2) μm thick, 6.2-7.9 (7.3) μm wide, and 4.6-6.3 (5.4) μm long, with polar capsules 1.9-2.5 (2.2) μm long and 1.1-1.4 (1.3) μm wide (n = 15). Kudoa surabayaensis n. sp. myxospores isolated from the flathead grey mullet were 5.8-6.7 (6.3) μm thick, 6.4-7.6 (6.9) μm wide, and 4.6-5.0 (4.7) μm long, with polar capsules 1.8-2.4 (2.1) μm long and 0.9-1.3 (1.1) μm wide (n = 25). These two Kudoa spp. showed critical differences in spore shapes (semiquadrate with unequal shell valves vs. equal shell valves), and absence vs. presence of uplifted shell valve termini. Nucleotide sequencing of rDNA supported the morphological differentiation of these two species. Furthermore, these two isolates were morphologically and phylogenetically distinct from any recorded Kudoa spp.

Myxozoan parasites of the genus Kudoa Meglitsch, 1947 are associated with post-mortem tissue degradation that causes great financial losses to commercial fisheries. Kudoa thyrsites (Gilchrist, 1924) is a species with a very wide host range including commercial tunas, mackerels, salmonids and flatfishes. A sample of 190 fishes of 18 species from the Madeira Archipelago and 30 Atlantic chub mackerel, Scomber colias Gmelin, and 30 blue whiting, Micromesistius poutassou (Risso), from the Portuguese mainland coast were examined for the presence of species of Kudoa. The prevalence of Kudoa spp. was 80% in M. poutassou and 60% in S. colias. No spore was detected in S. colias from Madeira, which was confirmed by specific PCR screening of the muscle from all individuals of S. colias. SSU rDNA analysis revealed that M. poutassou and S. colias from the Portuguese mainland coast were infected with K. thyrsites, an economically important myxozoan parasite. Both sequences were identical with sequences of the eastern Atlantic K. thyrsites genotype, including that from the type host of this parasite. This is the first report of K. thyrsites from M. poutassou and S. colias. The fact that spores of species of Kudoa were not detected in fishes screened in the Madeira Archipelago may be explained by various ecological factors, such as the absence of a continental shelf, a short insular shelf, and oceanic waters with low productivity, all resulting in reduced abundance of benthic organisms. Consequently, it is possible that as yet unknown annelid definitive hosts of Kudoa spp. are absent or very rare near Madeiran coasts.

A synopsis of the species of Zschokkella Auerbach, 1910 (Myxozoa: Bivalvulida: Myxidiidae) is presented, including 94 nominal species from piscine hosts and 3 additional nominal species from amphibian or reptilian hosts. The most relevant morphological and morphometric features of both myxospores and trophozoite stages (when available) are presented for each species in tabulated format.

Few studies have examined community structure among myxozoan species in fish. Herein myxozoan communities are described from 2 cyprinid species, the spottail shiner (Notropis hudsonius) and the common shiner (Luxilus cornutus), from mesotrophic and eutrophic localities in rivers in southwestern Quebec, Canada. Four myxozoan species were found, and total prevalence ranged from 60 to 86.7% in spottail shiners at 4 localities along the Richelieu River. Component species richness ranged from 2 to 4 and mean infracommunity richness from 0.87 to 1.47. Prevalence, component species richness, and infracommunity species richness in the spottail shiner were comparable to those from other localities in the Great Lakes and St. Lawrence River that were exposed to municipal effluents. Nine myxozoan species were found in common shiners from the Bras d'Henri micro-watershed. Component species richness varied from 6 to 8 at 4 localities, with total prevalence being 100% at all localities except 1, where it was 80%. Mean infracommunity richness ranged from 1.73 to 2.27. Prevalence, component species richness, and infracommunity species richness in the common shiner from the Bras d'Henri micro-watershed were among the highest observed for myxozoan communities from any host species to date. It is concluded that moderate levels of eutrophication are sufficient to generate species-rich communities of myxozoan parasites in fishes., (© American Society of Parasitologists 2021.)

A new coelozoic myxosporean species, Zschokkella epinepheli n. sp., collected from the gallbladder of the white grouper Epinephelus aeneus (Perciformes: Serranidae) from the bay of Bizerte, Tunisia, is described based on morphological and molecular characteristics. Myxospores and plasmodia were observed floating free in the bile. Mature plasmodia were polysporic and subspherical in shape, measuring 85.0-94.0 μm long and 70.0-82.0 μm wide. Mature myxospores were ovoid in valvular view, measuring 10.0 ± 1.7 (8.0-11.0) μm in length and 7.0 ± 0.3 (6.6-7.5) μm in width. Polar capsules were pyriform and equal in size, measuring 3.0 ± 0.2 (2.8-3.6) μm in length and 2.3 ± 0.3 (1.8-2.7) μm in width. Myxospore valves had 12-14 longitudinal striations. Based on the small subunit rDNA, the new species Z. epinepheli n. sp. differs from all other Zschokkella species for which there is a DNA sequence deposited in GenBank. Phylogenetic analysis revealed that Z. epinepheli n. sp. clustered in the marine subclade of Zschokkella species within the biliary tract IV clade. This is the first report of a Zschokkella species from the gallbladder of an epinephelin fishes.

The present study describes Henneguya tietensis n. sp. parasitizing Prochilodus lineatus (Valenciennes, 1837) from Brazil, through morphological and molecular analysis. Myxospores were found in the gill tissue of P. lineatus with a prevalence of 10%. The myxospores consisted of two elongate, elliptical shell valves each with a long, tapering caudal appendage. The morphology was consistent with Henneguya and the myxospore measured as follows (mean ± SD): total length 55.5 ± 2.1 μm, body length 16.2 ± 1.1 μm, body width 5.5 ± 0.1 μm, caudal appendages length 39.0 ± 2.0 μm. The polar capsules were 7.3 ± 0.2 μm long, 1.7 ± 0.2 μm wide and contained a polar filament coiled 11 to 13 turns. Histological analysis showed the plasmodia developing in the middle region of each gill filament and caused a focally extensive distension by forming a space-occupying mass within the gill filament. The ultrastructural analysis allowed the observation of the presence of mature myxospores throughout the plasmodium. Phylogenetic analysis with the SSU rDNA gene showed Henneguya tietensis n. sp. as a sister species of the subclade formed by Henneguya piaractus and Henneguya brachypomus that infect fishes of the genus Piaractus. The genetically closest species was H. piaractus, which showed a similarity of 82.4%. Using molecular and morphological characterization, the myxozoan parasite described herein represents a new species of the genus Henneguya.

The present study describes Henneguya lacustris n. sp. parasitizing Astyanax lacustris (Lütken 1875), from the Tietê River, State of São Paulo, Brazil, through morphological and molecular analysis. Myxospores of the parasites were found in the gills of A. lacustris with 13% prevalence of infection. The myxospores were oval and presented as measures (mean ± standard deviation): total length 18.3 ± 2.2 μm, body length 10.4 ± 1.6 μm, body width 4.9 ± 0.9 μm, tail length 7.2 ± 2.5 μm. The polar capsule was 4.8 ± 0.3 μm long and 1.5 ± 0.2 μm wide. The polar tubules, present inside the polar capsules, had 6 to 7 turns. The ultrastructural analysis allowed the observation of the asynchronous development of the myxospores. Phylogenetic analysis showed H. lacustris n. sp. as a sister species of H. chydadea Barassa and Cordeiro 2003 in a subclade formed by species that parasitize the gills of Characiform fish from Brazil. Using molecular and morphological characterization, this parasite was identified as a new species of the genus Henneguya Thélohan 1892.

Myxozoans of the family Myxobolidae are common parasites in fish. The diversity and ecology of the species of the genus Unicauda are poorly known, which hampers the understanding of the distribution and prevalence of this group of parasites. In the present study, cysts containing parasites whose morphology was consistent with the genus Unicauda were found in the circumorbital region of the ocular conjunctiva of the freshwater fish Moenkhausia grandisquamis Müller & Troschel, 1845 (Characiformes: Characidae) and Triportheus angulatus Spix & Agassiz, 1829 (Characiformes: Triportheidae). The spores have an oval body and long caudal appendage, with a mean total length of 65.2 ± 5.9 μm and width of 5.2 ± 0.7 μm, with two oval and symmetrical polar capsules of 4.9 ± 0.5 μm in length and 1.4 ± 0.2 μm in width, containing polar filaments with five or six coils. An integrated comparative analysis of the morphological characteristics of this parasite and partial sequences of the SSU rDNA gene supported the identification of a new species of histozoic parasite of the genus Unicauda found in fish from the Tocantins River basin, in the eastern Brazilian Amazon region. The new species was denominated by Unicauda tavaresii n. sp.

This paper reviews current knowledge of the structure, genesis, cytochemistry and putative functions of the haplosporosomes of haplosporidians (Urosporidium, Haplosporidium, Bonamia, Minchinia) and paramyxids (Paramyxa, Paramyxoides, Marteilia, Marteilioides, Paramarteilia), and the sporoplasmosomes of myxozoans (Myxozoa - Malacosporea, Myxosporea). In all 3 groups, these bodies occur in plasmodial trophic stages, disappear at the onset of sporogony, and reappear in the spore. Some haplosporidian haplosporosomes lack the internal membrane regarded as characteristic of these bodies and that phylum. Haplosporidian haplosporogenesis is through the Golgi (spherulosome in the spore), either to form haplosporosomes at the trans-Golgi network, or for the Golgi to produce formative bodies from which membranous vesicles bud, thus acquiring the external membrane. The former method also forms sporoplasmosomes in malacosporeans, while the latter is the common method of haplosporogenesis in paramyxids. Sporoplasmogenesis in myxosporeans is largely unknown. The haplosporosomes of Haplosporidium nelsoni and sporoplasmosomes of malacosporeans are similar in arraying themselves beneath the plasmodial plasma membrane with their internal membranes pointing to the exterior, possibly to secrete their contents to lyse host cells or repel haemocytes. It is concluded that these bodies are probably multifunctional within and between groups, their internal membranes separating different functional compartments, and their origin may be from common ancestors in the Neoproterozoic.

Two myxosporean species of the genus Sphaeromyxa were isolated from the gallbladders of marine fish in the South China Sea. Sphaeromyxa scorpaena n. sp. was collected from Scorpaenodes albaiensis Evermann and Seale, 1907. The mature myxospores were arcuate-shaped with tapered to pointed ends, and a length of 14.1 ± 0.7 (13.8-15.1) μm and a width of 5.2 ± 0.3 (4.9-5.8) μm. The polar capsules (PCs) were pyriform with a length of 3.2 ± 0.2 (3.1-3.5) μm and a width of 1.6 ± 0.1 (1.4-1.8) μm, and containing ribbon-like polar filaments irregularly folded 1.5-2.5 turns. Molecular characteristics and phylogenetic analysis based on 18S rDNA as well as morphological comparison confirmed that S. scorpaena n. sp. was a previously undescribed species. Sphaeromyxa theraponi, isolated from Terapon jarbua Forsskål, 1775, was reported for the first time from the South China Sea. The mature myxospores were slightly arched, tapering to bluntly rounded ends, with a length of 17.3 ± 0.9 (15.5-19.4) μm and a width of 4.8 ± 0.3 (4.1-5.3) μm. A sporoplasm was situated in the space between PCs in the myxospore. The PCs were pyriform, which contained ribbon-like polar filaments irregularly folded by 2-3 turns, with a length of 7.0 ± 0.5 (5.8-8.1) μm and a width of 2.6 ± 0.2 (2.2-3.0) μm. Our morphological and phylogenetic analyses suggest that the pointed ends of S. scorpaena n. sp. might be a secondarily acquired characteristic rather than an ancestral trait.

Four new actinospore types belonging to the sphaeractinomyxon collective group (Cnidaria, Myxosporea) are described from the coelomic cavity of a marine Baltidrilus sp. (Oligochaeta, Naididae) inhabiting a northern Portuguese estuary. Host identification supports the usage of marine oligochaetes, namely of the family Naididae Ehrenberg, 1828, as definitive hosts for myxosporeans inhabiting estuarine/marine environments. The absence of mixed infections in the host specimens analysed is suggested to reflect the influence of host-, parasite- and environmental-related factors regulating myxosporean-annelid interactions. Molecular analyses matched the SSU rDNA sequences of three of the four new types with those of mugiliform-infecting Myxobolus spp., namely Myxobolus mugiliensis and a Myxobolus sp. from flathead grey mullet Mugil cephalus, and Myxobolus labrosus from thicklip grey mullet Chelon labrosus. These results directly link, for the first time, the sphaeractinomyxon collective group to a myxospore counterpart, further confirming their previously hypothesized specific involvement in the life cycle of myxobolids that infect mullets. Acknowledging this life cycle relationship, the functionality of the sphaeractinomyxon morphotype is suggested to have been decisive for the evolutionary hyperdiversification of the genus Myxobolus in mullets. Unlike other actinospore morphotypes, sphaeractinomyxon lack valvular processes, which implies a limited capability for buoyancy. Considering the benthic-feeding nature of mullets, this feature is most likely crucial in promoting successful transmission to the vertebrate host.

The specimens of Trachinus draco collected from the Bay of Bizerte were found to be infected with a new Ceratomyxa species described as Ceratomyxa draconis n. sp. The sequence of small subunit ribosomal RNA gene obtained in this study differs from other Ceratomyxa sequences available in GenBank. Mature spores of this species were elongated and crescent-shaped in sutural view, measuring 7.4 ± 0.77 (6.4-8.0) μm in thickness and 30.8 ± 1.65 (28.8-32.8) μm in width. The polar capsules were spherical, equal in size, and measuring 3.3 ± 0.2 (3.6-4.0) μm in diameter. The Ceratomyxa draconis n. sp. showed a clearly seasonal variation of prevalence with highest prevalence noted during summer months.

Members of the myxosporean genus Kudoa are defined as having a myxospore with four or more shell valves (SVs) and a corresponding number of polar capsules (PCs). Here, we employed integrated taxonomic approaches to four kudoid isolates from Acentrogobius chlorostigmatoides and Konosirus punctatus from the South China Sea, off Guangdong, Pentanemus quinquarius from the Southeast Atlantic Ocean, off West African coast, and Atheresthes stomias from the Bering Sea, off Alaska, and concluded that all these kudoids were novel species, named Kudoa acentrogobia n. sp., Kudoa guangdongensis n. sp., Kudoa iidae n. sp., and Kudoa aburakarei n. sp., respectively. Kudoa guangdongensis n. sp., forming pseudocysts in the trunk muscle myofibers of K. punctatus, had large-sized tripod myxospores with three wing-like SV extensions and three PCs (one prominent PC and two rudimentary PCs). Phylogenetic analyses based on the 18S and 28S ribosomal RNA gene (rDNA) demonstrated its affinity to the genus Kudoa, not to Unicapsula characterized by a myxospore with one prominent PC and two rudimentary PCs, suggesting the atypical nature of this new species in the context of myxospore morphology. Three other kudoid isolates had four SVs and PCs in a semi-quadrated, ray-like, or cruciform myxospore, respectively, forming pseudocysts in the trunk muscle myofibers. Kudoa iidae n. sp. forming pseudocysts in the muscles of P. quinquarius from Southeast Atlantic Ocean had unique myxospores with ray-like form, showing close morphological resemblance to Kudoa rayformis, which were recorded from the muscle of Scomberomorus sierra from the Pacific Ocean off Panama. These two species had a phylogenetic relationship of morphological convergence, evolving separately in different sea areas. It is fairly difficult to differentiate K. acentrogobia n. sp. and K. aburakarei n. sp., prevalent in their host fishes, from kudoid species with similar myxospore morphology (e.g., Kudoa nova and Kudoa thyrsites, respectively), but distinct in phylogeny from known Kudoa spp. Combined taxonomic identification of multivalvulid myxosporeans based on both morphological criteria of isolated myxozoans and their molecular characterization could disclose their real biodiversity and phylogeny.

This study describes Henneguya sacacaensis n. sp. in specimens of the Osteichthyes Satanoperca jurupari (Heckel, 1840), collected in the Rio Curiaú Environmental Protection Area in the city of Macapá, state of Amapá Brazil. Using optical microscopy and molecular analysis, these cyst-shaped parasites were analyzed. The gills of 57.14% of the analyzed S. jurupari contained hundreds of spores. The cysts found on the gill lamellae were oval-shaped and whitish. The Henneguya spores had an average length of 46.5 (41.3-56.92) µm. The fusiform body of the Henneguya measured 16.5 (13.16-20.01) µm long and 5.1 (3.91-6.12) µm in width, the two polar capsules had a taper of 3.83 (3.4-4.32) µm and a width of 1.68 (1.4-1.99) µm, and the tail measured 30 (22.47-41.67) µm in length, containing a polar filament coiled seven to nine times. Morphogical and phylogenetic analysis allowed the preposition of a new species, Henneguya sacacaensis n. sp, that belongs to the family Myxobolidae and the genus Henneguya.

Thelohanellus magnacysta n. sp. (Bivalvulida: Myxobolidae) infects the skeletal muscle of blacktail shiner, Cyprinella venusta Girard, 1856 (Cypriniformes: Cyprinidae) in Bull Creek, Chattahoochee River Basin, eastern Georgia. Although numerous members of ThelohanellusKudo, 1933 have overlapping myxospore dimensions with the new species, it differs from all nominal congeners by polar filament coil number and polar capsule width as well as by lacking a mucous envelope, iodinophilic vacuole, and sutural markings. With the use of novel primers for Myxozoa, a phylogenetic analysis of the small subunit ribosomal DNA (SSU rDNA) suggests that the new species shares a recent common ancestor with a clade of cyprinid-infecting species of Myxobolus Bütschli, 1882 (Bivalvulida: Myxobolidae) and Thelohanellus. Consistent with other published research concerning the systematics of Thelohanellus, this result suggested that Thelohanellus and Myxobolus are polyphyletic and need revision. Histological sections of infected blacktail shiners confirmed that myxospores were only found within a plasmodium and only infected skeletal muscle and that plasmodia were encapsulated by a granuloma comprising varying degrees of acute granulomatous inflammation. The new species is the fourth of Thelohanellus reported from North America and the first reported from Cyprinella, as well as the first myxozoan described from the blacktail shiner., (© American Society of Parasitologists 2020.)