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Salmonids are affected by the economically significant whirling disease (WD) caused by the myxozoan parasite Myxobolus cerebralis . In the past, it was endemic to Eurasia, but it has now spread to different regions of North America, Europe, New Zealand, and South Africa. Among salmonids, rainbow trout is considered the most highly susceptible host. Upon entering to the host's body, the parasite invades the spine and cranium, resulting in whirling behaviour, a blackened tail, and destruction of cartilage. The disease is characterized by the infiltration of numerous inflammatory cells, primarily lymphocytes and macrophages, with the onset of fibrous tissue infiltration. Several efforts have been undertaken to investigate the role of various immune modulatory molecules and immune regulatory genes using advanced molecular methods including flow cytometry and transcriptional techniques. Investigation of the molecular and cellular responses, the role of STAT3 in Th17 cell differentiation, and the inhibitory actions of suppressors of cytokine signaling (SOCS) on interferons and interleukins, as well as the role of natural resistance-associated macrophage proteins (Nramp) in WD have significantly contributed to our understanding of the immune regulation mechanism in salmonids against M. cerebralis . This review thoroughly highlights previous research and discusses potential future directions for understanding the molecular immune response of salmonids and the possible development of prophylactic approaches against WD.

Myxobolus Bütschli, 1882 is the most speciose myxozoan genus, although some species have only been described according to the morphological characteristics of spores. In the present study, a new Myxobolus species infecting the gill lamellae of goldfish from Chongqing, China, was described using a comprehensive analysis of morphological, molecular, and histological data. Mature spores were flat-pear in valvular view with tapering anterior and rounded posterior ends, measuring 11.0 ± 0.4 (10.4-11.6) μm in length and 10.3 ± 0.3 (9.6-11.0) μm in width. Two equal-sized elongate pyriform polar capsules were 5.6 ± 0.6 (4.5-6.4) μm long and 3.5 ± 0.5 (2.4-4.1) μm wide. Polar tubules were coiled with 8 or 9 turns. The small-subunit ribosomal DNA gene sequence length of the present species was 1951 nt, and the highest similarity was 97.99% with M. pyramidis. Comparative analysis of the morphological and molecular data revealed that the present species was distinct from other known myxosporeans. Plasmodia were located at the interlamellar troughs nearing the top of the primary gills. Infection by the present species destroyed the original structure of gill lamellae and caused an inflammatory response, eventually leading to fish dyspnea. The morphological, molecular, and pathological data from the present study can be used for aquaculture since they provide guidance for easy detection and future control of this myxosporidiosis., Competing Interests: Declaration of Competing Interest The authors declare no conflicts of interest., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Ceratomyxa Thélohan, 1892 is one of the largest genera of the sub-phylum Myxozoa, and has a worldwide geographical distribution. Ceratomyxa species mainly infect the gallbladder of marine fish and rarely infect histozoically or coelozoically freshwater fish. In the present study, yellow catfish, Trachysurus fulvidraco (Siluriformes, Actinopteri) collected from the low reach of Yellow River was first found to be infected with an unknown Ceratomyxa species in their gallbladder which was identified to be new to science and nominated as Ceratomyxa huangheensis n. sp. by an integrative taxonomic approach for myxosporeans. Spores are typical of the genus Ceratomyxa, and matures spores are arcuate, with posterior angle of 139.2 ± 11.6 (137.0-156.0)° and rounded ends, and measures 4.7 ± 0.6 (3.3-5.5) μm in length, and 10.7 ± 1.3 (8.5-13.3) μm in thickness. Two spore valves are slightly unequal and present elongated ovoid in the lateral view. Two equal spherical polar capsules, measuring 2.2 ± 0.4 (1.4-2.8) μm × 2.0 ± 0.4 (1.0-2.5) μm in size locates adjacent to the straight suture line. The obtained partial small subunit ribosomal DNA (SSU rDNA) sequence was unique among all myxozoans in GenBank, and the highest similarity is 85.3% with Ceratomyxa sparusaurati, a marine fish-infecting congener. Phylogenetic analysis further indicated that this novel species did not cluster with other freshwater fish-infecting congeners from South America to form an independent clade, but was phylogenetically positioned within the marine fish-infecting clade. The possible presence of different radiation trajectories between Ceratomyxa huangheensis n. sp. and Amazonian freshwater Ceratomyxa lineage was discussed. This is the first report of Ceratomyxa species in the Yellow River basin and the second freshwater Ceraotomyxa species in China, even in Asia., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Objective: Myxosporidiosis of bagrid fishes has been a focus of aquaculture research in recent years. The purpose of this study is to characterize a novel myxobolid, named Myxobolus xiushanensis n. sp., infecting Yellowhead Catfish Tachysurus fulvidraco in China., Methods: We used molecular biology, morphology, phylogeny, and histopathology in the present study., Result: Mature myxospores were circular to ellipsoidal in valve view, measuring 12.2 ± 0.4 μm (mean ± SD; range = 11.2-13.2 μm) in length and 10.6 ± 0.4 μm (9.5-11.1 μm) in width. Two oval polar capsules were equal in width (3.4 ± 0.2 μm; 3.0-3.8 μm) but slightly unequal in length: 5.6 ± 0.3 μm (5.3-6.1 μm) and 4.7 ± 0.2 μm (4.4-5.5 μm). The polar capsule was packed with five to seven spirals of polar tubules. Histopathological investigation demonstrated that the plasmodium under the cuticular layer of the gill arch only induced a local inflammatory response and did not cause serious damage to the gill arch's internal structure. The two small subunit (SSU) ribosomal DNA sequences of M. xiushanensis n. sp. showed 100% similarity and uniqueness, and the highest similarity with other myxosporean sequences in GenBank was 90.27% (query coverage = 94%). The secondary structures of the SSU ribosomal RNA revealed that the present species was distinctly different from related species in regions V4 and V7. Phylogenetic analysis showed that M. xiushanensis n. sp. clustered independently within a branch., Conclusion: These results enrich our understanding of the biodiversity of myxobolids infecting bagrid fishes and provide fundamental data for the diagnosis of myxosporidiosis., (© 2023 American Fisheries Society.)

During an investigation of Myxobolus diversity in the Chinese longsnout catfish Tachysurus dumerili (Bleeker), a new species infecting the intracranial epidermis of the host was discovered. Upon opening the cranial cavity, several round whitish plasmodia measuring 0.55-0.80 mm in diameter were observed. Fresh spores (n= 50) were pyriform in the frontal view and fusiform in the sutural view, with a length of 15.4±0.6 (13.9-16.5) μm, width of 9.1±0.4 (8.3-9.8) μm, and thickness of 7.0±0.4 (6.3-7.9) μm. The spores had smooth shell surfaces and transparent membrane sheaths in the posterior. No folds, intercapsular appendix, and caudal appendages were observed. Two equal polar capsules were pyriform and measured 7.5±0.5 (6.7-8.7) μm in length and 3.2±0.3 (2.5-3.6) μm in width. The polar filaments were coiled with five to six turns and perpendicular to the polar capsule length. A BLAST search indicated M. dumerilii sp. n. was closely related to five Myxobolus species (with sequences similarities ranging from 90.54% to 96.52%) found in different organs of yellow catfish Tachysurus fulvidraco (Richardson), rather than the T. dumerili-infecting species M. branchiola Dong and Zhao, 2014 (with 90.5% sequence similarity). Phylogenetic analysis showed that M. dumerilii sp. n. didn't form sister clade with brain-infecting Myxobolus spp, but clustered with M. jianlinensis Gao et Zhao, 2020 and M. voremkhai Akhmerov, 1960 within the Siluriformes-clade with highly supported values, indicating that the host specificity may play a stronger signal than site infections during the evolution of Myxobolus species. Based on the morphological, ecological, and molecular differences observed between the newly discovered species and other available Myxobolus species, M. dumerilii sp. n., is proposed and described in this study., (© 2023. The Author(s), under exclusive licence to Springer Nature B.V.)

A new species of Chloromyxum Mingazzini, 1890, C. dasyatidis n. sp., is described from the gallbladder of the elasmobranch fish Dasyatis pastinaca (Linnaeus) from the Mediterranean coast off Tunisia. Mature myxospores were subspherical measuring 13.0 ± 0.3 (12.5-13.9) µm in length, 11.3 ± 0.5 (10.2-12.2) µm in width, and 11.8 ± 0.5 (11.4-12.7) µm in thickness. Four pyriform polar capsules 4.4 ± 0.3 (4.0-4.9) µm long and 3.5 ± 0.3 (3.0-3.8) µm wide. Valves were ornamented by 5 to 7 elevated surface ridges and displayed short caudal filaments. Chloromyxum dasyatidis n. sp. had an overall prevalence of infection of 38.5%, with significant seasonal variation, being more prevalent during summer (60.0%) and autumn (70.8%). Phylogenetically, this species grouped within the Chloromyxum sensu stricto lineage, forming a small clade together with the genetically more similar species C. acuminatum and C. myolibati, both from stingray hosts., (© 2023. The Author(s), under exclusive licence to Springer Nature B.V.)

This paper provides an updated checklist of species-level identified myxosporeans from marine and freshwater fishes in Vietnam. The list includes 51 nominal species (38 marine and 13 freshwater) belonging to 9 genera: Myxobolus Bütschli, 1882 (26 species); Kudoa Meglitsch, 1947 (6 species); Henneguya Thélohan, 1892 (6 species); Thelohanellus Kudo, 1933 (5 species); Unicapsula Davis, 1924 (2 species); Ceratomyxa Thélohan, 1892 (2 species), Zschokkella Auerbach, 1909 (2 species); Auerbachia Meglitsch, 1960 (1 species), and Meglitschia Kovaleva, 1988 (1 species). For each parasite species, information on myxospore morphology, line drawings, fish hosts, infection sites, and collection locality in Vietnam are reported. Where available, we also provide GenBank accession numbers for nucleotide sequence data. In addition, taxonomic status of several species was discussed and Myxobolus eszterbaueri nom. nov. is proposed as a junior homonym for Myxobolus hakyi Baska, Voronin, Eszterbauer, Müller, Marton & Molnár 2009, which is preoccupied., (© 2023. The Author(s), under exclusive licence to Springer Nature B.V.)

The southern striped shiner, Luxilus chrysocephalus isolepis (Hubbs & Brown) is a relatively large minnow belonging to the true minnow family Leuciscidae Bonaparte. Between May 2020 and January 2022, 55 L. c. isolepis were collected from watersheds in Montgomery (n = 6), Polk (n = 17) and Sevier (n = 32) counties, Arkansas, USA, and their gills, gallbladders, urinary bladders, fins, integument, other major organs, and musculature were examined for myxozoans. Gills of 11 (34%) individual southern striped shiners from Sevier County were infected with a new myxozoan, Myxobolus carlhubbsi n. sp. A qualitative and quantitative morphological description was based on formalin-fixed preserved myxospores, and molecular data consisted of a 1,970 base pair sequence of the partial small subunit rRNA gene from ethanol-preserved specimens. Histologically, plasmodia filled and expanded interlamellar troughs. Hyperplastic epithelial and goblet cells filled interlamellar troughs adjacent to plasmodia, but inflammatory response was limited to scattered lymphocytes. Phylogenetic analysis revealed that M. carlhubbsi n. sp. is a member of a clade of species with pyriform myxospores parasitizing North American Pogonichthyinae Girard and North American and Eurasian Leuciscinae Bonaparte. This is the first report of a myxozoan from L. c. isolepis. This article was registered in the Official Register of Zoological Nomenclature (ZooBank) as urn:lsid:zoobank.org:pub:D10D71C2-2C75-4A1C-80ED-B98FF36CB509., (© 2023. The Author(s), under exclusive licence to Springer Nature B.V.)

The genus Henneguya Thélohan, 1892 (Cnidaria: Myxosporea: Myxobolidae) encompasses a large number of species that mostly infect freshwater fish belonging to 71 families of Actinopterygii. A synopsis of Henneguya species described between 2012 and 2022 is herein presented. It includes 57 species described during the last decade, and one species missing from the previous synopses, adding to a total of 254 species that have been formally described within this genus. Biological characters and myxospore morphometry are presented for each species record., (© 2023. The Author(s), under exclusive licence to Springer Nature B.V.)

We describe Ceratomyxa saurida Zhao et al . 2015 and Ceratomyxa mai sp. nov. (Myxozoa: Ceratomyxidae) from the East China Sea. C. saurida was found in the gallbladders of 3/13 specimens of its type host, Saurida elongata Temminck and Schlegel 1846 (Aulopiformes). Myxospore characters were consistent with the original description to which we have added small subunit (SSU) rRNA gene data. C. mai sp. nov. was found in gallbladders of 3/13 specimens of S . elongata and 5/13 specimens of Neobythites sivicola Jordan and Snyder 1901 (Ophidiiformes). Mature myxospores of C. mai sp. nov. were crescentic in sutural view, with a deeply concave posterior angle 142.2±8.2° (125.8‒158.2°) and an arched anterior side. Shell valves were smooth and equal, 20.9±1.9 (17.3‒24.7) µm thick and 9.2±0.5 (8.1‒9.9) µm long, and joined at a straight, thin sutural plane passing between two nematocysts (polar capsules). The nematocysts were equal-sized, pyriform, 2.6±0.2 (2.4‒2.9) µm long and 2.7±0.2 (2.4‒3.3) µm wide, with their tapered ends pointed toward each other, located in the anterior third of the spore. Sequences of the SSU rRNA gene and internal transcribed spacer 1 showed that the isolates of C. mai sp. nov. obtained from S. elongata and N. sivicola were identical. The SSU rRNA gene sequence of C. mai sp. nov. was distinct from all known myxosporeans and clustered with C. saurida , and then with Ceratomyxa filamentosi Kalatzis, Kokkari and Katharios 2013, both of which also infect Aulopiformes fishes.

A myxozoan survey was performed on specimens of thicklip grey mullet Chelon labrosus (Risso) captured from the Douro River estuary, northern Portugal. Eleven new species, all belonging to the genus Myxobolus Bütschli, 1882 (M. abdominalis n. sp., M. aestuarium n. sp., M. caudalis n. sp., M. chelonari n. sp., M. cucurbitiformis n. sp., M. douroensis n. sp., M. intestinicola n. sp., M. invictus n. sp., M. labicola n. sp., M. peritonaei n. sp., and M. pinnula n. sp.) are described based on microscopic and molecular data, confirming the known high radiation of these myxozoans in mullets. Additionally, Myxobolus pupkoi Gupta et al., 2022 is reported for the first time from C. labrosus, bringing forth a novel case of morphological plasticity between geographic isolates. We consider that molecular-based comparisons are imperative for the description of mugiliform-infecting Myxobolus, with distance estimation further matching two of the novel Myxobolus spp. with sphaeractinomyxon types previously reported from another Portuguese estuary. This finding supports sphaeractinomyxon as specific life cycle counterparts of Myxobolus that infect mullets. Phylogenetic analyses of 18S rDNA retrieved a monophyletic clade of mugiliform-infecting myxobolids comprising well-supported lineages of species parasitizing mullets from the genera Chelon, Mugil, Crenimugil, and Planiliza. The existence of more than one Chelon- and Planiliza-infecting lineage reveals that myxobolids parasitized members of these genera multiple times during their evolution. Lastly, the elevated number of unmatched sphaeractinomyxon sequences included in the Chelon-infecting lineages clearly shows that Myxobolus diversity hosted by this genus remains underrated., (© J. Guimarães et al., published by EDP Sciences, 2023.)

A synopsis of 43 nominal species from five genera of tailed Myxobolidae infecting Indian freshwater and marine fishes is presented. The main characteristic of this group is the presence of at least one tail-like caudal process. For each species, relevant morphological and morphometric data are provided, such as the host(s), site(s) of infection within the host and sampling state. A key for the identification of 13 genera of tailed Myxobolidae is also included.

Tetracapsuloides bryosalmonae is a malacosporean endoparasite that causes proliferative kidney disease (PKD) in wild and farmed salmonids in Europe and North America. The life cycle of T. bryosalmonae completes between invertebrate bryozoan and vertebrate fish hosts. Inside the fish, virulence factors of T. bryosalmonae are induced during infection or interactions with host cells. T. bryosalmonae genes expressed in vivo are likely to be important in fish pathogenesis. Herein, we identify in vivo induced antigens of T. bryosalmonae during infection in brown trout ( Salmo trutta ) using in vivo induced antigen technology (IVIAT). Brown trout were exposed to the spores of T. bryosalmonae and were sampled at different time points. The pooled sera were first pre-adsorbed with antigens to remove false positive results. Subsequently, adsorbed sera were used to screen a T. bryosalmonae cDNA phage expression library. Immunoscreening analysis revealed 136 immunogenic T. bryosalmonae proteins induced in brown trout during parasite development. They are involved in signal transduction, transport, metabolism, ion-protein binding, protein folding, and also include hypothetical proteins, of so far unknown functions. The identified in vivo induced antigens will be useful in the understanding of T. bryosalmonae pathogenesis during infection in susceptible hosts. Some of the antigens found may have significant implications for the discovery of candidate molecules for the development of potential therapies and preventive measures against T. bryosalmonae in salmonids., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Kumar, Sudhagar, Shivam, Nilsen, Bartholomew and El-Matbouli.)

The European anchovy represents the main fisheries for countries in the Mediterranean and Black Sea basins. The skeletal muscle of 13 of 48 (27.1%) Engraulis encrasicolus (L.) specimens from North East Atlantic waters (FAO 27.8.c) was found infected with interfibrillar elongated plasmodia (130-980 µm in length) containing mature myxospores belonging to the genus Kudoa Meglitsch, 1947. No flesh softening was found associated with infection. Fresh myxospores were 10.8 ± 0.7 (9.1-12.3) µm in width 1, 11.3 ± 0.9 (9.5-13.4) µm in width 2, 6.7 ± 0.4 (5.8-7.4) µm in thickness, and 6.9 ± 0.5 (5.8-7.5) µm in length. They were almost stellate in apical view having three pointed-edged shell valves bearing three small polar capsules equal in size 5.0 ± 0.3 (4.4-5.4) μm long and 2.4 ± 0.2 (2.0-3.0) μm wide, and one rounded- to rarely bluntly pointed-edged shell valve bearing a large and particularly wide polar capsule 6.8 ± 0.4 (5.9-7.6) μm long and 4.1 ± 0.2 (3.6-4.4) μm wide. Morphological and morphometrical comparisons between these myxospores and those of Kudoa thyrsites (Gilchrist, 1923) from the clupeid Sardina pilchardus (Walbaum) (North East Atlantic waters, FAO 27.9.a), with which exhibited a similarity of 98.9% and 96.2% using SSU and LSU rDNA sequences, respectively, support the creation of Kudoa encrasicoli n. sp. Morphometrical analysis of the polar capsules of flattened myxospores is suggested as a useful approach to differentiate phylogenetically related kudoids with stellate or almost stellate myxospores bearing four polar capsules., (© 2022. The Author(s).)

Purpose: The present study describes a new species of myxosporean, Sphaeromyxa cornuti n. sp. infecting the gallbladder of the Moorish idol, Zanclus cornutus (Linnaeus 1758) collected from Lakshadweep waters of the Arabian Sea., Methods: Fish were collected using traps and cages. The morphology of mature spores recovered from the gallbladder of Z. cornutus was studied under Nomarski Differential Interference Contrast (DIC) optics. The molecular and phylogenetic analyses were based on SSU rDNA., Results: Sphaeromyxa cornuti n. sp. is characterized by arcuate myxospores with tapering extremities and round ends in valvular, and slightly sigmoid in sutural views (19.2-24.7 µm × 4.1-5.7 µm). The two polar capsules are unequally elongate-ovoid in shape and positioned at opposite ends of the spore (6.2-9.7 µm × 1.7-2.6 µm). Each encloses an irregularly folded, ribbon-like polar tubule, which is oriented parallel to polar capsule axis. In molecular and phylogenetic analyses, the present myxosporean revealed significant differences with related forms and clustered together with S. hellandi within the 'incurvata' group of the Sphaeromyxa clade with high nodal support., Conclusions: Morphological, morphometric, molecular and phylogenetic differences between our material and previously described species of Sphaeromyxa, along with host and geographic variations indicate that the present myxosporean is unique and the name Sphaeromyxa cornuti n. sp. is proposed. This forms the first report of a myxosporean parasite-infecting Z. cornutus., (© 2022. The Author(s) under exclusive licence to Witold Stefański Institute of Parasitology, Polish Academy of Sciences.)

In the Columbia River Basin (CRB), USA, anthropogenic factors ranging from dam construction to land use changes have modified riverine flow and temperature regimes and degraded salmon habitat. These factors are directly implicated in native salmon and steelhead (Oncorhynchus species) population declines and also indirectly cause mortality by altering outcomes of ecological interactions. For example, attenuated flows and warmer water temperatures drive increased parasite densities and in turn, overwhelm salmonid resistance thresholds, resulting in high disease and mortality. Outcomes of interactions between the freshwater myxozoan parasite, Ceratonova shasta, and its salmonid hosts (e.g., coho O. kisutch and Chinook salmon O. tshawytscha) are well-described, but less is known about effects on chum salmon O. keta, which have a comparatively brief freshwater residency. The goal of this study was to describe the distribution of C. shasta relative to chum salmon habitat in the CRB and assess its potential to cause mortality in juvenile chum salmon (listed as threatened in the CRB under the U.S. Endangered Species Act). We measured C. shasta densities in water samples collected from chum salmon habitat throughout the lower CRB during the period of juvenile chum salmon outmigration, 2018-2020. In 2019, we exposed caged chum salmon fry from two hatchery stocks at three C. shasta-positive sites to assess infection prevalence and survival. Results demonstrated: (1) C. shasta was detected in spawning streams from which chum salmon have been extirpated but was not detected in contemporary spawning habitat while juvenile chum salmon were present, (2) spatiotemporal overlap occurs between C. shasta and juvenile chum salmon in the Columbia River mainstem, and (3) low densities of C. shasta caused lethal infection in chum salmon fry from both hatchery stocks. Collectively, our results suggest C. shasta may limit recovery of chum salmon now and in the future., Competing Interests: The authors have declared that no competing interests exist.

We describe Myxobolus cataractae n. sp., found parasitizing Imparfinis mirini from Cachoeirinha stream, a tributary of Pardo River, Paraná River basin, municipality of Botucatu, São Paulo State, Brazil. We based the description on myxospore morphology, histology, and small subunit ribosomal DNA partial sequences. Whitish and elongated mass of myxospores were found in gill filaments, occupying an intrafilamental position. The myxospores were slightly ellipsoidal and the morphometric measurements revealed a myxospore body length of 7.8 ± 0.4 μm; myxospore width of 5.9 ± 0.4 μm; and thickness of 3.9 ± 0.3 μm. The polar capsules are equal in size and measured 3.5 ± 0.2 μm in length and 1.7 ± 0.2 μm in width. The polar tubules had 6-7 coils. Myxobolus cataractae n. sp. is classified as the intrafilamental type and chondreal subtype by the histological analysis of gills. Phylogenetic analysis showed a well-supported subclade formed by species that parasitize gills of Siluriformes, with M. imparfinis as a sister species of Myxobolus cataractae n. sp. Using molecular and morphological characterization, this cryptic species was identified as a new species of the genus Myxobolus., (Copyright © 2022 Elsevier B.V. All rights reserved.)

Ceratomyxa marginati n. sp. and C. aenei n. sp. are two new coelozoic myxosporean species infecting the gallbladder of the dusky grouper (Epinephelus marginatus) and the white grouper (Epinephelus aeneus), respectively. These two ceratomyxids were described using morphological characteristics and molecular analysis of the SSU rDNA. Ceratomyxa marginati n. sp. exhibits disporic plasmodia measuring 12-14 µm long and 11-12 µm wide and mature myxospores which are slightly crescent-shaped, measuring 6.0 ± 0.3 (5.6-6.8) µm in length and 12.9 ± 0.9 (11.5-14.0) μm in thickness. Plasmodia of C. aenei n. sp. were disporic and ellipsoidal and measured 28-32 μm in length and 19-22 μm in width. Mature myxospores of C. aenei n. sp. were elongated with unequal shell valves and measured 7.4 ± 0.6 (6.9-9.0) μm in length and 26.9 ± 2.4 (23.2-30.0) μm in thickness. Based on the SSU rDNA sequences, Ceratomyxa marginati n. sp. and C. aenei n. sp. are distinct from all other Ceratomyxa sequences available in GenBank. Phylogenetic analysis reveals that the two new species branched together within a clade with other Ceratomyxa species from different host families and different geographical localities with maximum support (100%)., (© 2022. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Purpose: The class Myxozoa consists of microscopic spores that typically present uniform morphological simplifications, containing one or two sporoplasms surrounded by valve cells. The present study analysed the morphological and phylogenetic characteristics of what proved to be a new species of Ceratomyxa found in the gallbladder of Mesonauta festivus Heckel, 1840 captured on the Piririm River in the municipality of Macapá, in Amapá state, Brazil., Methods: The fish were collected in gillnets, and were transported alive to Amapá State University in Macapá for the analysis of the organs. Crescent- or arch-shaped free spores were observed under light microscopy, containing two polar capsules and a suture line, which is typical of the morphology of the genus Ceratomyxa, contained within plasmodia that had snake-like movements. The gallbladder was conserved in 80% alcohol for the analysis of the 18S rDNA gene, and in Davidson solution for standardhistology., Results: Ceratomyxa spores (n = 20) were observed in 11 of the 50 samplesanalysed, and thus had a prevalence of 22%, and had a mean length of 4.2 ± 0.5 µm, mean thickness of 22.75 ± 0.3 µm, and two polar capsules, 1.86 ± 0.3 µm long and 1.63 ± 0.1 µm thick., Conclusions: The phylogenetic analysis indicated that the new species forms a cluster with other freshwater Amazonian Ceratomyxa species, described previously. The new species represents the first description of a taxon of the genus Ceratomyxain fish from the Brazilian state of Amapá., (© 2021. Witold Stefański Institute of Parasitology, Polish Academy of Sciences.)

The taxonomy of myxosporeans was traditionally dependent solely upon the spore morphological and morphometric data. Intensive reports of intraspecific morphological variation, however, are increasingly challenging the taxonomic approaches for myxosporeans. In the present work, the morphological pleomorphism of myxospores of Myxobolus drjagini (Akhmerov, 1954) was observed. More interestingly, all of these pleomorphic myxospores occurred in the same plasmodium of M. drjagini, which refutes the previous hypothesis that morphological variation of M. drjagini was derived from its responses to differences in nutrition and immunological responses associated with different host tissues. Bearing the intraspecific morphometric and morphotype variation in mind, the combination of morphological, ecological and molecular data should be applied to the species identification and delimitation for myxosporeans. This is the first reported myxobolid species with high pleomorphic myxospores which are present in the same plasmodium.

A new myxozoan species, Ceratomyxa binhthuanensis n. sp. (Myxosporea: Ceratomyxidae), was found in the gall bladder of blacktip grouper Epinephelus fasciatus (Perciformes: Serranidae) in the East Sea of Vietnam. Myxospores were observed floating free in the gall bladder of 3 out of 20 fish examined (15%). Mature myxospores were elongate and slightly crescent-shaped and measured 12.2 ± 1.3 (10.8-16.0) μm in thickness and 5.8 ± 0.6 (4.8-6.9) μm in length, with two smooth equal shell valves. The two polar capsules were spherical and equal in size, measuring 2.6 ± 0.3 (2.3-2.9) μm in diameter. The posterior angle was slightly concave, 153.7° ± 5.6° (148.9°-166.0°). Molecular analysis of SSU rDNA sequence showed that Ceratomyxa binhthuanensis n. sp. differs from other Ceratomyxa spp. available in GenBank. Phylogenetic analysis indicated that C. binhthuanensis n. sp. was closely related to three species, Ceratomyxa nolani, Ceratomyxa yokoyamai, and Ceratomyxa cutmorei, which also infect fish hosts of the genus Epinephelus., (© 2022. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Proteases and their inhibitors play critical roles in host-parasite interactions and in the outcomes of infections. Ceratonova shasta is a myxozoan pathogen that causes enteronecrosis in economically important salmonids from the Pacific Northwest of North America. This cnidarian parasite has host-specific genotypes with varying virulence, making it a powerful system to decipher virulence mechanisms in myxozoans. Using C. shasta genome and transcriptome, we identified four proteases of different catalytic types: cathepsin D (aspartic), cathepsin L and Z-like (cysteine) and aminopeptidase-N (metallo); and a stefin (cysteine protease inhibitor), which implied involvement in virulence and hence represent target molecules for the development of therapeutic strategies. We characterized, annotated and modelled their 3D protein structure using bioinformatics and computational tools. We quantified their expression in C. shasta genotype 0 (low virulence, no mortality) and IIR (high virulence and mortality) in rainbow trout Oncorhynchus mykiss , to demonstrate that there are major differences between the genotypes during infection and parasite development. High proliferation of genotype IIR was associated with high expression of the cathepsin D and the stefin, likely correlated with high nutrient demands and to regulate cell metabolism, with upregulation preceding massive proliferation and systemic dispersion. In contrast, upregulation of the cathepsin L and Z-like cysteine proteases may have roles in host immune evasion in genotype 0 infections, which are associated with low proliferation, low inflammation and non-destructive development. In contrast to the other proteases, C. shasta aminopeptidase-N appears to have a prominent role in nematocyst formation in both genotypes, but only during sporogenesis. Homology searches of C. shasta proteases against other myxozoan transcriptomes revealed a high abundance of cathepsin L and aminopeptidase homologs suggesting common gene requirements across species. Our study identified molecules of potential therapeutic significance for aquaculture and serves as a baseline for future research aimed at functional characterisation of these targets., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Alama-Bermejo, Bartošová-Sojková, Atkinson, Holzer and Bartholomew.)

Although species of the genus Ceratomyxa Thélohan, 1892 mostly parasitize marine fish around the world, a surprising diversity of the genus has recently been reported from Amazonian freshwater fish. In this study, we report a freshwater Ceratomyxa species parasitizing Hemiodus orthonops (Hemiodontidae) from the Paraná River (La Plata Basin) in a watershed flowing into the southern part of South America, which expands the geographic distribution of this fish parasite in the freshwater resources of the continent. We applied a combination of morphological, small subunit ribosomal DNA (SSU rDNA), and phylogenetic analyses, and vermiform-shaped plasmodia endowed with motility were found swimming in the bile of the fish. The characteristics of the plasmodia and myxospores of the Ceratomyxa species found in the Paraná River resembled those of Ceratomyxa fonsecai, a parasite of the congeneric host Hemiodus unimaculatus from the Tocantins River basin in northern Brazil. Due to the close morphological and morphometric resemblances and the impossibility of genetic comparison, the parasite found in H. orthonops from the Paraná River was designated as Ceratomyxa cf. fonsecai, and the definition of its taxonomic status was left for further study. Maximum likelihood and Bayesian analyses showed Ceratomyxa cf. fonsecai clustering within a well-supported clade, together with other Amazonian freshwater ceratomyxids. The present study suggests that shifts of the complex host/parasite between marine and freshwater environments were facilitated by marine incursions into South America in the Early Miocene., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Two new species of Myxobolus Bütschli, 1882 (Bivalvulida: Myxobolidae) are described from the gill of the black redhorse (Moxostoma duquesnei [Leueur][Cypriniformes: Catostomidae]) from the Little Tennessee River Basin, North Carolina, United States. Myxobolus branchiofilum n. sp. infects lumen of the lamellar arterioles and Myxobolus branchiopecten n. sp. infects the bone and cartilage at the tip of the gill rakers. They differ from all congeners by a combination of myxospore dimensions and the presence or absence of an iodinophilic vacuole in the sporoplasm, mucous envelope, intercapsular process, and sutural markings. A phylogenetic analysis of the small subunit ribosomal DNA recovered M. branchiopectin sister to Myxobolus sp. (AF378343) in a clade composed of 6 species of Myxobolus, which infect predominately cypriniform intermediate hosts. Myxobolus branchiofilum was recovered sister to Myxobolus ictiobus Rosser, Griffin, Quiniou, Alberson, Woodyard, Mischker, Greenway, Wise & Pote, 2016 in a clade composed of 8 species of Myxobolus, which predominately infect catostomid intermediate hosts. Histological sections of infected gill revealed intra-lamellar plasmodia of M. branchiofilum in the lumen of the lamellar arterioles and foci of M. branchiopecten developing in the bone and cartilage of the gill raker tip. These are the first myxozoans reported from the black redhorse. Given that these two new species are morphologically congeneric but recovered in distantly related clades, we discuss the persistent issue of myxobolid genera paraphyly/polyphyly., (© 2021. The Author(s), under exclusive licence to Springer Nature B.V.)

Infestation by parasitic copepods is a substantial problem in the cage culture of European sea bass (Dicentrarchus labrax). The Copepoda parasite, Lernanthropus kroyeri (Lernanthropidae), is one of the threats to the mariculture of sea bass. In the present study, we evaluated the data of abundance for L. kroyeri pre-adults and adult males (PAAM) and ovigerous adult females (AF o ) to estimate the internal infection pressure (IIP) in the same cage environment. The sea bass infested by L. kroyeri was collected from grow-out sea cages located in Gulluk Bay (Turkey) in September and October 2019. Mean fish weight and length of sea bass were 75.79 ± 1.66 g and 21.40 ± 0.56 cm, respectively. The fixed lag method was used to predict the abundance of L. kroyeri adult females. The overall prevalence of L. kroyeri was 60%. The mean abundance of PAAM and AF o varied from 0.8 ± 0.24 to 2.5 ± 0.67 and 2.9 ± 0.40 to 4.3 ± 0.55, respectively. The abundance of AF was strongly correlated with PAAM. The pattern of AF o and PAAM was interpreted as an indication of the continuous infestation of L. kroyeri on sea bass. Our results showed that the correlation of AF o abundance for five consecutive weeks was significant, representing the main determinative factor for the continuity of the parasitic load. In our approach, internal infestation pressure is the quantitative estimation of the potential infective copepodids, which are mainly characterized by AF abundance and the prevalence. We predicted that the internal infestation pressure could be high, even exceeding the 50.000 × 10 6 potential infective copepodids for one sea cage with the fish density of 20 sea bass/m 3 ., (© 2021 John Wiley & Sons Ltd.)

Members of the class Myxosporea Btschli, 1881 have a cosmopolitan distribution in a wide variety of fish species worldwide. In the present study, the black scorpionfish Scorpaena porcus collected from the Sinop coasts of the Black Sea was investigated for myxosporean parasites using both conventional and molecular methods in the period between September 2015 and August 2019. Using morphological and morphometric data, the myxosporean parasite Ceratomyxa scorpaeni Garbouj, Rangel, Castro, Hmissi, Santos, Bahri, 2016 was identified in the gall bladder of host fish. Molecular analysis of the 18S rDNA gene confirmed the identity of this parasite as C. scorpaeni. This is the first report of its occurrence in the Black Sea.

Investigations on myxozoan parasites of fish from Chongqing in China, revealed two Myxidium cuneiforme-like myxosporeans infecting the gallbladder of Cyprinus carpio carpio and Carassius auratus. We researched their myxospore morphology, and analyzed their genetic similarity and phylogenic relationships to other myxozoans based on small subunit ribosomal DNA (18S rDNA) sequences. Although both parasites recovered were morphologically similar, the myxosporean isolated from C. auratus was consistent in morphology to Myxidium cuneiforme, which was described from this host species. The parasite isolated from C. c. carpio had overlapping myxospore dimensions to M. cuneiforme, but on average, the polar capsules were not as long. More importantly, this parasite was genetically distinct from M. cuneiforme with 96.3% and 96.5% similarity in two sequences of 18S rDNA, and we propose the name Myxidium pseudocuneiforme n. sp. for this myxozoan from common carp. Its mature myxospores are ellipsoidal and asymmetric with pointed ends in valvular view, arc-shaped or fusiform in sutural view. The pyriform polar capsules are equal in size, and polar filament with 5-6 coils. This study highlights that molecular characteristics and host specificity are indispensable for myxozoan species identification when presented with the taxonomic dilemma of whether we are observing one species that exhibits slight morphological differences or multiple, but similar, species in different hosts., (© 2021 The International Society of Protistologists.)

The present study describes the morphological, histopathological and SSU rDNA data of a new myxosporean species infecting farmed Astyanax lacustris fish from the state of São Paulo, Brazil. Henneguya lambariensis sp. nov. was found infecting the gills, and the plasmodial development resulted in displacement, blood congestion, compression, deformation and lamellar fusion, stretching of the epithelia, hyperplasia of the epithelial cells, edema, and mild infiltration of the mast cells and lymphocytes. The SSU rDNA sequencing resulted in the sequencing of 1804 nucleotides that did not correspond to any myxosporean sequences deposited in GenBank. The closest phylogenetic affinity of the new species was to the South American Henneguya loreotoensis and Henneguya guanduensis, which also parasite the gills. The present study suggests this new myxosporean species has considerable pathogenic potential, and health monitoring should be implemented in A. lacustris fish farms to ensure production., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Enteromyxum leei is a causative agent of enteromyxosis, with a wide range of marine fish hosts. Recently, massive morbidity and mortality were caused by E. leei infection in cultured olive flounders in Korea. To reveal a relationship between E. leei abundance in culture water and the occurrence of parasite infection in host fish, we used a quantitative PCR assay targeting the 28S rDNA of E. leei in three fish farms (two where enteromyxosis had occurred and one where it did not) from April to November 2018. The gene of E. leei was detected at levels greater than 10 cells/L in the culture water where enteromyxosis occurred from July to September. Furthermore, 2 months after the detection in the water, the parasite gene (with more than 5,000 cells per 100 mg) was detected in fish intestine samples. However, in the fish farms where enteromyxosis had not occurred, the E. leei gene was detected at <10 cells in culture water (1 L) and fish intestine samples (100 mg). The quantification method used in this research provides a baseline of the infection timeline in olive flounder to develop effective management practices., (© 2021 John Wiley & Sons Ltd.)

The present study describes a new species of the genus Sphaerospora found in the urinary bladder of the flag cichlid, Mesonauta festivus collected in Corre Água district of the municipality of Macapá, Amapá State (Brazil). The study includes morphological and phylogenetic analyses of the new parasite, to determine the relationship of the new species with related myxosporean species. The new species has polysporous plasmodia, which vary in size and shape. The mature myxospores are subspherical shape in valvar view. In the sutural view, the myxospores are 5.3±0.2 (5.2-5.6) μm in length and 7.0±0.7 (6.3-7.7) μm in width, with two piriform polar capsules equal size, 2.5±0.2 (2.3-2.8) μm in length and 1.8±0.2 (1.6-2.0) μm in width. The phylogenetic analyses of a partial sequence of the 18S rRNA gene confirmed the status of the new species and determined the relationship of the new species and related myxosporean species.The sum of the evidence indicates that, Sphaerospora festivus n. sp. belongs to the family Sphaerosporidae, and is the first record of the genus Sphaerospora from Brazil.

Myxosporidiosis is an infectious disease caused by myxozoans of the Phylum Cnidaria, Class Myxosporea, and Order Bivalvulida, considered a common parasite in fresh and saltwater fishes that parasitize many organs, especially gills. In the present study, 49 specimens of fishes belonging to eight genera: Tetragonopterus, Leporinus, Myleus, Pirinampus, Rhapiodon, Pygocentrus, Ageneiosus, and Serrasalmus were collected and blood smears were made, fixed with absolute methanol, and stained with Giemsa 10% to survey hemoparasites. However, myxospores were found in the circulating blood of five (10.20%) fishes belonging to genus Tetragonopterus, Myleus, and Pygocentrus. Two morphological types of Myxobolus spp. were identified in all the five fish specimens analyzed. Usually, investigations on myxozoans in fish are carried out with the search for plasmodia or cysts in the fish organs and observation of the cavity of organs. Nevertheless, this study highlights the importance of also examining the blood of these animals, since these parasites can cause severe pathogenic diseases in fish. Thus, the blood analyses can proportionate preventive sanitary control for commercial fish avoiding economic loss.

Recent range expansions of whirling disease impelled us to understand the impacts of its causative agent, the myxozoan parasite Myxobolus cerebralis, on lesser-studied fish hosts. Mountain Whitefish Prosopium williamsoni overlap broadly with M. cerebralis across the western United States and Canada, and populations have experienced widespread declines since the 1990s. To evaluate effects of the parasite on Mountain Whitefish, we revisit formerly unpublished work of the Colorado Division of Wildlife (now Colorado Parks and Wildlife), comparing infection in age-matched Mountain Whitefish, Rainbow Trout Oncorhynchus mykiss, and Brown Trout Salmo trutta. To complement the original report, we reanalyze mortality data and include additional SEM imagery. Infection of M. cerebralis in juvenile Mountain Whitefish was characterized by a brief but heavy period of mortality in the first 2 weeks after exposure, with limited pathology. This clinical effect is unique among the known salmonid hosts of M. cerebralis., (© 2021 American Fisheries Society.)

The myxozoan parasite Myxobolus xiantaoensis is a fin pathogen of commercially important yellow catfish Tachysurus fulvidraco Richardson, 1846, in the freshwater ponds of China. In the present work, four geographical isolates of M. xiantaoensis were sampled from the fins of yellow catfish. It was found that the spores of four isolates exhibited few markable differences in morphometrics. The small subunit ribosomal DNA (SSU rDNA) sequences of four isolates were conspecific to the SSU rDNA sequence of M. xiantaoensis. No genetic level variation was observed, even in the characteristically more variable internal transcribed spacer (ITS) region. This absence of variability suggests high gene flow as a result of panmixia in the parasitic populations. ITS phylogeny placed four isolates of M. xiantaoensis in a clade together with myxozoans species infecting Siluriformes. The M. xiantaoensis infection inflicted severe hemorrhages on epidermis of ray-fins, which grew into inflammatory epithelial hyperplasia and lytic cartilage signs. The histochemical analysis of infected fins biopsies is characterized by damage of collagen components of cartilage, resulting in weakness, breaks, and missing fin rays. These tissue sections also had a remarkable inflammatory response around the fin cartilage, with the absence of mature spores and chondrocytes. These results indicate that the fin cartilage damage appeared before the development of tissue inflammation and the parasitic infestation of the fins. The present four geographical isolates of M. xiantaoensis were identified by a holistic approach of species characterization based on biological, morphological, and molecular evidence. These four isolates showed some morphological and genetic variations but within the intraspecific range., (Copyright © 2021. Published by Elsevier Ltd.)

The farming of Colossoma macropomum has intensified in recent years, leading to an increased need for research into the health of the fish. We therefore investigated the diversity of myxosporeans (Cnidaria: Myxozoa) infecting C. macropomum in a breeding system in the municipality of Rio Branco, in the state of Acre, Brazil. Twenty-four fish specimens were examined from June to August 2018. Our results revealed a high prevalence of infection, with 23 specimens (95.8%) exhibiting myxosporean plasmodia. Morphological analysis, based on light and electron microscopies, and molecular analysis (small subunit ribosomal DNA [SSU rDNA] sequencing) revealed the occurrence of three novel species of the genus Myxobolus. Plasmodia of Myxobolus guttae n. sp. were found in the fins of 75% of the specimens, and the myxospores were pear-shaped, measuring 12.3 ± 0.6 (10.3-13.5) μm in length, 8.1 ± 0.3 (7.1-8.6) μm in width, and 5.1 ± 0.6 (4.5-6.5) μm in thickness. The polar capsules were elongated and equal in size, measuring 6.8 ± 0.5 (5.8-7.6) μm in length and 2.5 ± 0.3 (1.8-3.1) μm in width, exhibiting polar tubules with 9-10 coils. The plasmodia of Myxobolus longus n. sp. were found in the gills of 45.8% of the C. macropomum specimens, and the myxospores were fusiform, measuring 16.4 ± 0.6 (14.7-17.3) μm in length, 7.1 ± 0.2 (6.8-7.7) μm in width and 5.5 ± 0.6 (4.4-6.6) μm in thickness. The polar capsules occupied more than half of the myxospore, exhibiting different sizes, the largest measuring 9.2 ± 0.5 (7.3-10.1) μm in length and 2.5 ± 0.3 (2.0-3.1) μm in width, while the smallest measured 8.5 ± 0.4 (7.1-9.1) μm in length and 2.4 ± 0.2 (1.9-3.0) μm in width. Both polar capsules contained polar tubules with 10-12 coils. For Myxobolus tambaquiensis n. sp., plasmodia were found in the opercular cavity of 41.7% of the fish specimens, and the myxospores had an oval shape, measuring 10.0 ± 0.4 (9.5-11.3) μm in length, 6.5 ± 0.2 (6.1-7.1) μm in width, and 4.6 ± 0.4 (3.9-5.6) μm in thickness. The polar capsules were elongated and equal in size, measuring 4.9 ± 0.2 (4.4-5.3) μm in length and 1.9 ± 0.2 (1.5-2.2) μm in width, closing with 8-9 coils of the polar tubule. The morphological and sequencing data of the SSU rDNA showed that the three species studied herein remain unknown to science, increasing the diversity of myxosporeans infecting C. macropomum, an iconic fish in South American freshwater fish farming. The SSU rDNA based phylogenetic analysis revealed that Myxobolus spp. parasites of C. macropomum did not have a monophyletic origin, identifying different times and pathways of the acquisition of parasites by this host species., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

A total of 30 specimens of the Amazonian electric knifefish, Brachyhypopomus beebei Schultz, 1944 (Gymnotiformes: Hypopomidae), were collected from the Peixe-Boi River in the state of Pará, Brazil (1°06'59" S; 47°18'26" W). Fragments of the brain tissue were extracted for analysis via optical microscopy, and 18 specimens (60%) presented microparasites of the genus Myxobolus, with unequal capsules. The spores were 18.6 µm (17.7-19.8 µm) long and 8.6 µm (8.4-9.0 µm) wide; the largest polar capsule was 13.0 µm (12.4-13.4 µm) long and 5.6 µm (5.3-6.0 µm) wide, and the smallest capsule was 5.0 µm (4.5-5.3 µm) long and 2.5 µm (2.3-2.6 µm) wide. Infected brain fragments were extracted for histological processing and staining with hematoxylin-eosin and Ziehl-Neelsen. Some fragments were conserved in ethanol for molecular genetics analysis. A partial sequence of the 18S DNA gene was obtained from the spores, which did not correspond to any other sequences deposited in GenBank, although it did form a clade with other Myxobolus parasites of the nervous system. The morphological data, together with molecular phylogeny, supported the designation of a new species Myxobolus freitasi n. sp.

We identified Myxobolus anatolicus Pekmezci, Yardimci, Yilmaz & Polat, 2014 and 4 novel Myxobolus species from the Anatolian khramulya Capoeta tinca (Cyprinidae) in northern Turkey based on morphology, histology, and phylogenetic analysis. M. karaeri sp. nov. plasmodia were observed in the skin doublets between fin rays, the surfaces of the operculum, the gill arch membrane, and in the skin of the fin base. M. samsunensis sp. nov. plasmodia were observed in epithelial tissue inside and on the surface and midline of the gill filaments. M. cakmaki sp. nov. presented as a typical vascular species, which develops in large plasmodia at the end of the gill filaments. The chondrophilic M. ekingeni sp. nov. was detected by histology inside the cartilaginous gill arch and the cartilaginous gill rays of the filaments. Phylogenetic analysis of small subunit ribosomal DNA sequences revealed that M. karaeri sp. nov. and M. samsunensis sp. nov. were clustered with Myxobolus species that infect gills, scales, and fins of cyprinids. M. cakmaki sp. nov. grouped with Myxobolus species that exclusively infect the gills of cyprinids. No molecular data were available for M. ekingeni sp. nov.

A new cnidarian myxosporean infecting the spleen of an economic and ecological important bryconid fish (Salminus franciscanus) is described based on integrative taxonomic approach including morphological, ultrastructural, biological traits, geography, molecular data and phylogenetic analysis. In a total of thirty specimens examined, nineteen (63.3%) were infected by an undescribed parasite species belonging to the genus Myxobolus. Plasmodial development was asynchronous, with young development in the periphery and mature myxospores in the central area and without projections and microvilli in the plasmodial wall. Mature myxospores were ovoid in shape and measured 7.9 ± 0.2 μm (7.6-8.1 μm) in length and 5.4 ± 0.1 μm (5.0-5.6 μm) in width. The two polar capsules were equal in size, occupying a little more than half of the myxospore body, measuring 4.0 ± 0.2 μm (3.9-4.1 μm) in length and 1.7 ± 0.1 μm (1.5-1.8 μm) in width. The polar tubules coiled in six turns, perpendicular to the long axis of polar capsule. Phylogenetic analysis placed the new species within a clade containing nine myxobolid species from South American characiforms fish and appears as a close species of Myxobolus pantanalis. Nevertheless, the sequences of the new species and M. pantanalis have a large genetic divergence of 13.5% in their SSU rDNA. In light of the differences observed from the integrative taxonomy, we confidently considered that this isolate is a new species of cnidarian myxosporean, M. douradae n. sp., increasing the knowledge of diversity of this enigmatic group of cnidarians., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Out of the total 2600 myxozoan species reported so far, Thelohanellus Kudo, 1933 is the second most prevalent genus comprising of about 150 species after Myxobolus Butschli, 1882 infecting both freshwater and marine fishes. In general, these are mostly histozoic (within the tissues) and sometimes coelozoic (in body cavities). The present paper deals with the study of a novel Thelohanelloid T. neocyprini infecting gills of Catla catla and species complex among the member species of Thelohanellus genus from the Indian subcontinent infecting gills, fins and muscles. The species forming species complex are T. rohitae, T. catlae, T. jiroveci, T. seni, T. bifurcata, T. dykovae, T. neocyprini, T. filli, T. muscularis and T. theinensis. The phylogenetic analysis was also done on the basis of 18S rDNA. The homogeneity was found to be between 90 and 99%. The factors responsible for the species complex could be phylogeography, host reluctant, organ and tissue specificity of the these myxozoan parasites. Study of more genetic markers facilitated with morphotaxonomy can be used to sort out the occurrence of species complex among the morphologically different species having similar genetic makeup and vice-versa., (Copyright © 2020. Published by Elsevier Ltd.)

A new myxozoan species belonging to the genus Henneguya was isolated from the serous membrane of the visceral cavity of the hognosed catfish Brochis multiradiatus from Peruvian Amazon. Whitish plasmodia, macroscopically visible, were found in four of the thirty examined fishes. Mature myxospores were ellipsoidal in shape in frontal view and had a total length of 44.5 ± 0.6 μm (43.9-45.1), spore body measured 18.7 ± 0.9 μm (16.8-19.6) in length, 7.1 ± 0.2 μm (6.6-7.4) in width and 5.5 ± 0.3 μm (4.9-5.6) in thickness. The two polar capsules were elongated and equal in size, measuring 9.1 ± 0.1 μm (8.8-9.4) in length and 1.7 ± 0.1 μm (1.6-1.8) in width, occupying half of the myxospore body. Polar tubules coiled in 10-11 turns perpendicular to the long axis of the polar capsule. The caudal appendage was not bifurcated and measured 25.8 ± 0.6 μm (24.7-26.5) in length. The sequencing of the 18S rDNA gene resulted in 1400 bp and this sequence did not match any of the myxozoans available in GenBank. Phylogenetic analysis placed the new species in a well-supported subclade of Henneguya spp. infecting callichthyid fishes, with Henneguya loretoensis being the closest species. This study is the first description of a myxozoan species, Henneguya multiradiatus n. sp. from a fish of the genus Brochis., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

In the present study, we provide supplementary data for Myxidium cf. rhodei Léger, 1905 based on morphological, histological and molecular characterization. M. cf. rhodei was observed in the kidneys of 918 out of 942 (97%) roach Rutilus rutilus (Linnaeus, 1758). Myxospores of M. cf. rhodei were fusiform with pointed ends, measuring 12.7 ± 0.1 SD (11.8-13.4) µm in length and 4.6 ± 0.1 (3.8-5.4) µm in width. Two similar pear-shaped polar capsules were positioned at either ends of the longitudinal axis of the myxospore: each of these capsules measured 4.0 ± 0.1 (3.1-4.7) µm in length and 2.8 ± 0.1 (2.0-4.0) µm in width. Polar filaments were coiled into 4 to 5 turns. Approximately 18-20 longitudinal straight ridges were observed on the myxospore surface. The suture line was straight and distinctive, running near the middle of the valves. Histologically, the plasmodia of the present species were found in the Bowman's capsules, and rarely in the interstitium of the host. Phylogenetic analysis revealed that M. cf. rhodei was sister to M. anatidum in the Myxidium clade including most Myxidium species from freshwater hosts.

Reproduction of Trachurus picturatus off the western Portuguese coast was studied over 1 yr. During histological analyses, the presence of Kudoa sp. was detected in advanced vitellogenic oocytes. Kudoa infections are known to cause economic loss through the induction of post-mortem liquefaction of fish muscles, but ovarian infection as reported in this study will seriously affect the reproductive potential of the species and thus impact fisheries productivity. Only females showed gonad infection which led to total degradation of advanced vitellogenic oocytes. No macroscopic, somatic or condition indices revealed differences between infected and uninfected females, rendering this infection event a concealed suppression of reproduction.

Malacosporeans are a group of endoparasitic cnidarians (Myxozoa) that use freshwater bryozoans and fish as final and intermediate hosts, respectively. The malacosporean Tetracapsuloides bryosalmonae causes proliferative kidney disease (PKD), an emerging disease in aquaculture and wild fish populations, including threatened salmonids in Europe and the USA. Mixed infections of malacosporeans are often encountered, and a monitoring tool for screening of multiple malacosporean species in either their fish or bryozoan hosts is therefore desirable. We describe an inexpensive method that combines PCR amplification of the partial 18S rRNA gene (~260 bp) and a single-step restriction fragment length polymorphism (RFLP) method for identification of 10 malacosporean lineages and species. We demonstrate and test this methodology on a set of DNA extracted from malacosporeans infecting fish kidney and tissues sampled from bryozoan colonies and compare the results with Sanger sequencing of the same parasite DNA isolates. The PCR-RFLP and Sanger sequencing methods agreed in 100% of cases. The PCR-RFLP method offers a number of opportunities, including screening large panels of host tissue samples to gain insights into infection patterns, characterizing mixed infections, and confirming highly pathogenic T. bryosalmonae infections. The method can also be further refined as new sequence data become available for malacosporeans.

Metacercariae of Diplostomidae are widely distributed in America and may cause diplostomiasis, an ocular disease in fishes. The aim of this study is to report the occurrence of metacercariae of Austrodiplostomum compactum in Plagioscion squamosissimus (non-native fish species) from Nova Avanhandava Reservoir, Tietê River, Brazil and an explanation for the high infection rates with this parasite in the Paraná River Basin is proposed. Eyes of 70 hosts were examined, the metacercariae were preserved and identified. The prevalence (P), mean intensity of infection (MII) ± standard deviation, mean abundance (MA) ± standard deviation, were calculated and a bibliographic review was performed. There was no difference in parasitism between male and female hosts. The values of P = 80%, MII = 21.55 ± 3.25 and MA = 17.24 ± 2.91 were high, as in most studies in areas where P. squamosissimus were introduced, while these values were low in areas of natural occurrence. This may be explained by the genetic susceptibility of the host to the parasite. The entire population of P. squamosissimus from the Upper Paraná has been founded by a few specimens, resulting in very low genetic variability. Consequently, the population may be highly susceptible to A. compactum.

Proliferative kidney disease (PKD) of salmonids, a chronic immunopathology caused by the myxozoan parasite Tetracapsuloides bryosalmonae, is exacerbated by increased water temperatures. PKD causes economic concerns to trout farmers and contributes to the decline of wild salmonid populations in North America and Europe. The parasite occurs as far north as Norway and Iceland in Europe and was confirmed from California to southern British Columbia in the American continent. In mid-September 2011 adult chum salmon (Oncorhynchus keta) were sampled from Kantishna River, a tributary to Yukon River in Alaska. Clinical PKD was diagnosed based on the macroscopic appearance of mottled kidneys that were uniformly swollen and by the detection of tumultuous histozoic extrasporogonic and coelozoic sporogonic stages of T. bryosalmonae in renal tissue by histopathology. Archived samples provided the molecular confirmation and local strain identification, representing the first confirmed case of PKD in wild adult chum salmon, also co-infected with Parvicapsula minibicornis that represents another novel myxozoan detection in Alaska. Our investigation was extended to another case from August/September 1997, with mortality following furunculosis and ectoparasite co-infections, in sockeye salmon (Oncorhynchus nerka) pre-smolts net-pen reared in English Bay Lakes, Alaska. Immunohistochemistry on archived histological preparations confirmed T. bryosalmonae sporogonic and extrasporogonic stages, indicating a severe to resolving PKD, with concomitant Chloromyxum spp. infection. Those cases provide the first documentation that this parasite is present in Alaska and causes PKD in wild and cultured salmonids in the region. The known geographic range of T. bryosalmonae can be extended to ~267 km south of the Arctic Circle, representing the northernmost detection in America. Given the vast size of Alaska and small resident population, it is likely that T. bryosalmonae remained undetected, but more recently became evident due to the clinical manifestation of PKD, possibly linked to increasing water temperatures reported at the sample locations., (Copyright © 2020 Australian Society for Parasitology. All rights reserved.)

Myxosporea (Cnidaria: Myxozoa) are common fish parasites with complex life cycles that involve annelid hosts. Two economically important salmonid-infecting myxosporeans from rivers of the northwestern United States, Ceratonova shasta (Noble, 1950) and Parvicapsula minibicornis Kent et al., 1997, have life cycles that require a freshwater annelid host, identified previously as Manayunkia speciosa Leidy, 1859. This species was described originally from Pennsylvania, with subsequent records from New Jersey, the Great Lakes and west coast river basins. Despite apparent widespread distributions of both suitable fish hosts and the nominal annelid host, both parasites are restricted to river basins in the northwestern US and have never been recorded from the Great Lakes or the eastern US. In this study, we sampled 94 infected and uninfected annelids from two northwestern US rivers to confirm the identity of the host. We found these new specimens had mitochondrial COI sequences with no more than 4.5% distance from each other, but with at least 11% divergence from M. speciosa sampled from near the type locality (New Jersey) and Lake Superior. We did not recover any M. speciosa from either west coast river. The annelid from the Klamath and Willamette rivers showed marked sexual dimorphism that has not been reported in any Manayunkia described to date, though it is apparent that this had been missed in M. speciosa. Accordingly, we describe a new taxon, Manayunkia occidentalis sp. nov., and show that it can host both C. shasta and P. minibicornis. We suspect that previous records of Manayunkia from Pacific Northwest watersheds are likely to be M. occidentalis sp. nov. and not M. speciosa. Sampling of Manayunkia from additional localities is underway to test if the novel Manayunkia species is the only freshwater fabriciid annelid present across the Pacific Northwest.

Parasite composition can be affected by physiological and ecological changes during host ontogeny. Intertidal fish do not travel long distances and live in the same area throughout their lifetimes, meaning that parasite communities can differ across geographic ranges. The objective of this study was to analyse the parasite communities of three fish species (Hypsoblennius sordidus, Helcogrammoides cunninghami and Scartichthys viridis) collected from the Chilean coast. The composition of parasite species was compared among host ontogenetic stages (larvae, juveniles and mature fish) and geographic areas. A total of 184 larval, 252 juveniles and 217 mature individuals were collected in the northern area (c. 24°S), and 186 larval, 192 juveniles and 112 mature individuals from the central area (c. 33°S). Ectoparasites were most prevalent in fish from the central area, whereas endoparasites were most prevalent in the northern area. The parasite species richness varied significantly between geographical areas for H. sordidus and H. cunninghami, but the parasite composition varied significantly between geographical areas for all fish species analysed. Therefore, the geographical area was the most important factor determining the parasite composition of intertidal fish species. The absence of endoparasites in fish larvae and the increased infestation in juvenile and mature fish may be explained by the shift in habitat from the water column to intertidal pools where prey abundance and availability are higher. On the other hand, hydrographic barriers affecting prey distributions may also offer an explanation as to the differences in parasite composition.

Kudoa ajurutellus n. sp. (Multivalvulida: Kudoidae) is described as a parasite of the Bressou catfish, Aspistor quadriscutis (Siluriformes) collected at Ajuruteua beach, northeastern Pará, Brazil. The new species is described based on the morphology of the spores and the sequence of the 18S rDNA. This parasite was found only in the skeletal muscular tissue of fish hosts. In the apical view, the spores were pseudo-quadrangular in shape, with rounded borders, and four symmetrical capsules. The spores are 6.63±0.53 μm in length and 8.16±0.75 μm in width. The polar capsules are piriform, 3.45±0.30 μm in length and 1.87±0.15 μm in width. The phylogenetic analysis based on the sequence of the 18S rDNA gene indicated that the new species is clearly distinct from all other Kudoa species, and that the degree of differentiation is constant with the existence of a new species of the genus Kudoa. This is the first marine Kudoa species recorded in northeastern of the State of Pará.

During an ongoing investigation into the myxosporean diversity of common carp in Chongqing, China, Myxobolus parakoi sp. nov. was found to infect the gill lamellae of Cyprinus carpio (Linnaeus, 1758). Both morphological and SSU rDNA data revealed that M. parakoi sp. nov. was distinct from other myxosporeans. Mature myxospores of M. parakoi sp. nov. were pyriform in the frontal view. The spores were 15.98 ± 0.78 (14.59-17.72) μm in length and 7.84 ± 0.78 (6.66-9.75) μm in width. The two polar capsules were pyriform and equal in size, exhibiting 8.72 ± 0.50 (7.76-9.92) μm in length and 3.03 ± 0.23 (2.63-3.56) μm in width. Polar filaments within the polar capsules were coiled with 11 or 12 turns. Phylogenetic analysis revealed that M. parakoi sp. nov. and Myxobolus koi (FJ710800) were clustered together, forming a sister subclade with Myxobolus tanakai. This study also implied that the morphology of the myxospores was generally correlated with their phylogenetic relationship, while such correlation was not strong or consistent.

Chloromyxum thymalli infects the gall bladder of fishes belonging to the genus Thymallus. Here, we provide morphological, histological, and molecular data to complete the original description. Rounded plasmodia measured from 25 to 30 µm in diameter and contain 8 developing spores at most. Spores are typical of the genus Chloromyxum. Mature spores are subspherical, (mean ± SD) 9.05 ± 0.08 µm (range: 7.75-10.03 µm) long, and 8.81 ± 0.09 µm (range: 7.19-10.01 µm) wide. Four pyriform polar capsules are equal in size, 3.4 µm long and 2.7 µm wide on average. Histopathological analysis showed that presporogonic stages were found in the lamina propria under the epithelial cells of the host's gall bladder and destroyed the integrity of the epithelium. The obtained SSU rDNA sequences of C. thymalli did not match any available sequences in GenBank. Phylogenetic analysis showed that C. thymalli forms a sister cluster with C. truttae, with a subclade of Chloromyxum spp. that infect the gall bladder of freshwater teleosts.