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During surveys on myxosporean parasites of Lake Balaton and River Danube fishes, two Thelohanellus spp. were found on tench (Tinca tinca) and on common nase (Chondrostoma nasus). They were identified as Thelohanellus pyriformis and Thelohanellus cf. fuhrmanni, respectively. Myxospores of T. pyriformis from tench were collected from arteria branchialis afferens of gill filaments. The mature myxospores of this species were pyriform in shape and 19 ± 0.6l (18-19.5) long, 8.2 ± 0.54 (7.5-9) wide, 7.3 ± 0.25 (7-7.5) thick containing polar tubules with 9-10 turns. The plasmodia of Thelohanellus cf. fuhrmanni were collected from under the skin of snout of the common nase. The myxospores were pyriform, 16.3 ± 0.39l (15.5-16.5) long, 6.5 ± 0.55 (6.3-7) wide, 6.3 ± 0.53 (5.8-7) thick containing polar tubules with 6 turns. Small subunit ribosomal DNA sequences of both Thelohanellus species differed from other known myxozoans. The myxospores morphology, histopathology and ssrDNA sequences supported a diagnosis of T. pyriformis from tench and T. cf. fuhrmanni from common nase.

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About forty-two species of Myxobolus have been previously described to parasitize characiform fishes in South America. Here, we described a new myxozoan species, Myxobolus parodontidis n. sp., in the gills of Apareiodon piracicabae (Characiformes, Parodontidae) from the streams of the Middle Paranapanema River basin, Brazil. The proposed new species is supported by a combination of morphological, ultrastructural, and molecular characterization (small subunit ribosomal DNA). Thirteen specimens of A. piracicabae were analyzed and 30.8% was infected by Myxobolus parodontidis n. sp. The myxospores was classified as intralamellar asymmetric type. A few aberrant myxospores with three polar capsules were observed: the spore length and width were the same of normal myxospores, but the polar capsules had smaller sizes. Ultrastructural analysis showed that the plasmodial membrane of Myxobolus parodontidis n. sp. was in direct contact with the host tissue and a connective capsule surrounding the plasmodium was not observed. The phylogenetic analysis showed that the main influence in the clustering of species of myxobolids seems to be related to the phylogenetic relationships established among fish hosts, mainly at the level of family and order. This is the first record of a myxozoan species parasitizing parodontid fish, contributing to the knowledge of the biodiversity of myxozoans from Brazil.

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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.

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The American pumpkinseed Lepomis gibbosus, was introduced to Europe more than one hundred years ago. Currently it is a common fish in European freshwaters but relatively few specific parasites infect this fish in this new habitat. In Europe only a single species, Myxobolus dechtiari seems to represent the American myxosporean fauna of centrarchid fishes. M. dechtiari was found in both Portugal and Hungary. This species forms plasmodia with elongated shape inside the cartilaginous rays of gill filaments. In the advanced stage of infection, after disruption of plasmodia, small groups of myxospores remain enclosed in the cartilaginous gill rays causing distortions in the filaments. Myxospores were ellipsoidal in frontal view and lemon-shape in sutural, length 12.5 ± 0.46 (12-13.4) µm, width 10 ± 0.37 (9.6-10.4) µm, and thickness 7.4 ± 0.37 (7-8) µm; the polar capsules were pyriform, equal in size, length 5.6 ± 0.21 (5.3-6) µm, width 3.2 ± 0.16 (3-3.6) µm; Seven to eight polar tube coils were arranged perpendicularly to the capsule length. There was a small, round, 0.4 ± 0.1 (0.3-05) (N = 50) intercapsular appendix in the spores. The small subunit ribosomal DNA (ssrDNA) of M. dechtiari differed from other myxozoans sequenced to date. Phylogenetic analysis of the ssrDNA gene sequence placed this species in a clade including actinospores and Myxobolus species: Raabeia type1, Triactinomyxon sp., and Myxobolus osburni infecting the same host fish. The focus of our study was to prove that the pumpkinseed, a fish originated from North-America introduced one of its myxosporean parasite to Europe. Emphasis was put on to demonstrate the unique feature of this parasite causing infection in the cartilaginous gill rays.

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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.

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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.

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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.

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This study increases the known biodiversity of cnidarian parasites in neotropical bryconid fishes. Two novel Myxobolus species are described based on morphology, ultrastructure and small subunit ribosomal DNA (ssrDNA) sequencing: Myxobolus vetuschicanus n. sp. infecting fins of Salminus franciscanus and Myxobolus mineirus n. sp. infecting the mesentery of Brycon orthotaenia from the São Francisco River basin, Minas Gerais State, Brazil. Ultrastructural analysis of the two species revealed an asynchronous sporogenesis process, with germinative cells and young developmental stages of myxospores in the periphery of the plasmodia. In M. vetuschicanus n. sp., the plasmodia were surrounded by a layer of fibroblasts and in M. mineirus n. sp., the plasmodial membrane had direct contact with the host tissue. The phylogenetic analysis based on the ssrDNA of Henneguya/Myxobolus species showed that the two novel Myxobolus species grouped in subclades together with other parasite species of bryconid fishes.

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

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Plerocerci of the monotypic Paranybelinia otobothrioides were found parasitizing the subtropical neritic krill Nyctiphanes simplex in the Gulf of California, Mexico. The plerocerci were recovered from two microhabitats of the intermediate host, typically embedded inside the digestive gland (hepatopancreas) or rarely in the hemocoel. The morphology of the simple, single-layered blastocyst surrounding the entire scolex is unique within the Trypanorhyncha by having four large funnel-like pori or openings possibly with feeding and/or excretory function. One of the openings is located anteriorly and three at the posterior end. Scolex surface ultrastructure shows hamulate and lineate spinitriches covering the bothrial surface, capilliform filitriches at the anterior scolex end and on the scolex peduncle, and short papilliform filitriches on the long appendix. This pattern resembles that of species of the Tentaculariidae; but differs in that the hamulate spinitriches, which appear lineate at the bothrial margins, densely cover the entire distal bothrial surface. Tegumental grooves are present on the posterior bothrial margin, lacking spinitriches. Paranybelinia otobothrioides and Pseudonybelinia odontacantha share the following unique combination of characters: two bothria with free lateral and posterior bothrial margins, homeoacanthous homeomorphous armature, tegumental grooves, the distribution of the hamulate spinitriches, and the absence of prebulbar organs. Both genera infect euphausiids as intermediate hosts. Sequence data of the partial ssrDNA gene place Pa. otobothrioides sister to the family Tentaculariidae, and the Kimura two-parameters (K2P) distance between Pa. otobothrioides and species of the family Tentaculariidae ranged from 0.027 to 0.039 (44-62 nucleotide differences). These data suggest both species be recognized in a family, the Paranybeliniidae, distinct from, albeit as sister taxon to, the Tentaculariidae. High prevalence of infection (<14%) and ontogenetic changes of Pa. otobothrioides support N. simplex as a required intermediate host and suggest a zooplanktophagous elasmobranch as final host in the Gulf of California.

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Twelve Myxobolus species have been previously described to parasitize Bryconidae fish in South America. Here, we describe two novel myxosporean species that parasitize economically important Bryconidae from the São Francisco River basin in Brazil. Myxospores morphometry, morphology, small-subunit ribosomal DNA - ssrDNA sequences, and other biological traits were used in the taxonomic analysis. Phylogenetic analysis was performed to assess the position of the new Myxobolus species among the closest Myxobolus/Henneguya. Myxobolus iecoris n. sp. was found infecting the liver of Salminus franciscanus (dourado). Myxospores were oval with the anterior region aculiform in frontal view and biconvex in lateral view and measured 11.4-14.2 (12.8 ±â€¯0.8) µm long, 7.7-9.9 (8.7 ±â€¯0.6) µm wide, 6.5-7.5 (6.9 ±â€¯0.4) µm thick. Two pyriform and equal-sized polar capsules measuring 4.9-7.4 (5.9 ±â€¯0.5) µm long and 2.3-3.5 (3.0 ±â€¯0.2) µm wide contained polar tubules with 8-9 turns. Myxobolus lienis n. sp. was found infecting the spleen of Brycon orthotaenia (matrinxã). Myxospores were round to oval in frontal view and biconvex in lateral view and measured 10.3-13.8 (12 ±â€¯0.6) µm long, 6.8-9.3 (8.3 ±â€¯0.5) µm wide, and 6.9-7.0 (7.0 ±â€¯0.6) µm thick. Two oval and equal-sized polar capsules measured 3.9-5.8 (4.6 ±â€¯0.5) µm long and 2.0-3.5 (2.8 ±â€¯0.3) µm wide contained polar tubules with 5-6 turns. Ultrastructural analysis revealed asynchronous sporogenesis with germinative cells and young sporogonic stages in the periphery of the plasmodia. A connective tissue capsule was observed surrounding Myxobolus lienis n. sp., but it was absent for Myxobolus iecoris n. sp. Maximum likelihood and Bayesian inferences showed the two novel species clustering in a well-supported subclade composed by Myxobolus spp. of bryconids. Myxobolus iecoris n. sp. appeared as a sister species of M. aureus and Myxobolus lienis n. sp. as sister to M. umidus.

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This study presents morphologic, molecular and phylogenetic data about two new species of the genus Myxobolus and of the previously described Myxobolus colossomatis, all which are found infecting the Colossoma macropomum, a fish whose natural habitat is the Amazon Basin of Brazil, from where the specimens for this study were caught. A total of 51 C. macropomum specimens were examined between October of 2014 and January of 2016. Plasmodia of the myxosporeans were found infecting several organs: Myxobolus matosi n. sp. and Myxobolus longissimus n. sp. were respectively found in the inner face of the operculum and in the wall external surface of the stomach and gill arch. M. matosi n. sp. were 9.6 ± 0.4 µm in length, 7.0 ± 0.3 µm in width and 5.0 ± 0.3 µm in thickness of the myxospore. M. longissimus n. sp. measured 19.1 ± 0.4 µm in length, 9.4 ± 0.3 µm in width and 8.3 ± 0.4 µm in thickness. The polar capsules, which were elongated, showed 4.3 ± 0.4 µm in length and 1.9 ± 0.1 µm in width for M. matosi n. sp. and 10.5 ± 0.2 µm in length and 2.5 ± 0.1 µm in width for M. longissimus n. sp. The Myxobolus colossomatis had two myxospore morphotypes: 1) Ellipsoidal myxospores measuring 11.6 ± 0.4 µm in length and 7.6 ± 0.2 µm in width. Their elongated polar capsules measured 5.6 ± 0.2 µm in length and 2.5 ± 0.2 µm in width; 2) Oval myxospores measuring 10.4 ± 0.5 µm in length and 7.7 ± 0.3 µm in width. Their polar capsules were 5.4 ± 0.2 µm in length and 2.4 ± 0.0 µm in width. The number of turns of the polar filament was 7-8 coils. The molecular comparison of the small subunit ribosomal DNA (ssrDNA) showed a genetic divergence of 10.3% between M. matosi n. sp. and M. colossomatis, 22.4% between M. matosi n. sp. and M. longissimus n. sp., and 23.2% between M. longissimus n. sp. and M. colossomatis. Myxobolus cf. colossomatis, a parasite of Piaractus mesopotamicus, showed 11.1% of genetic divergence to M. colossomatis, demonstrating them to be distinct species. Phylogenetic analysis, based on sequences of the ssrDNA, showed the M. matosi n. sp. to be a sister species of M. colossomatis, and it also showed M. longissimus n. sp. to be a sister branch in the lineage composed by Myxobolus cf. cuneus and Henneguya pellucida.

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We describe two novel myxosporean parasites from Brachyplatystoma rousseauxii, an economically important freshwater catfish from the Amazon basin, Brazil. Myxobolus tapajosi n. sp., was found in the gill filaments of 23.5% of 17 fish, with myxospores round to oval in frontal view and biconvex in lateral view: length 15 (13.5-17) µm and width 10.7 (9.6-11.4) µm; polar capsules equal, length 5.8 (4.6-7.1) µm and width 3 (2.3-3.8) µm containing polar tubules with 6-7 turns. Ellipsomyxa amazonensis n. sp. myxospores were found floating freely or inside plasmodia in the gall bladder of 23.5% of fish. The myxospores were ellipsoidal with rounded extremities: length 12.8 (12.3-13.6) µm and width 7.6 (6.7-8.7) µm; with two equal, slightly pyriform polar capsules, length 3.8 (3.8-4.0) µm and width 3.1 (2.5-3.4) µm, containing polar tubules with 2-3 turns. We combined spore morphometry, small-subunit ribosomal DNA data, specific host, and phylogenetic analyses, to identify both of these parasites as new myxozoan species. Maximum likelihood and Bayesian analyses showed that Myxobolus tapajosi n. sp. clustered in a basal branch in a subclade of parasites from exclusively South American pimelodid fishes. Ellipsomyxa amazonensis n. sp. clustered within the marine Ellipsomyxa lineage, but we suspect that although the parasite was collected in freshwater, its hosts perform a large migration throughout the Amazon basin and may have become infected from a brackish/marine polychaete host during the estuary phase of its life.

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We investigated the involvement of oligochaetes in the life cycles of fresh water myxozoan parasites in Brazil. In a fish farm in the State of Mato Grosso do Sul, we examined 192 oligochaetes and found that two (1%) released Aurantiactinomyxon type actinospores. We identified infected oligochaetes by morphology: both were Pristina synclites, from family Naididae. This is the first report of the involvement of this species in the life cycle of myxozoans. Small-subunit ribosomal DNA sequences of Aurantiactinomyxon type 1 (1882 nt) and Aurantiactinomyxon type 2 (1900 nt) did not match any previously sequenced myxozoan in the NCBI database, with the highest BLAST search similarities of 83% with Myxobolus batalhensis MF361090 and 93% with Henneguya maculosus KF296344, respectively, and the two aurantiactinomyxons were only 75% similar to each other (over ~ 1900 bases). Phylogenetic analyses showed that Aurantiactinomyxon type 1 had closest affinities with myxozoans from fish hosts in Order Characiformes, and Aurantiactinomyxon type 2 had affinities with myxozoans from fish of Order Siluriformes.

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We describe three new Henneguya spp. (Myxobolidae) found parasitizing two species of cichlid fish from the Amazon basin, Brazil: H. tucunarei n. sp. from gill filaments of Cichla monoculus and H. tapajoensis n. sp. from gill filaments of Cichla pinima, both from the Tapajós River, Pará State and H. jariensis n. sp. in the fins of Cichla monoculus from the Jari River, Amapá State. We based descriptions on myxospore morphology and small subunit ribosomal DNA sequences, and used a phylogenetic analysis to compare the new Henneguya species with known relatives. Spores of the three species had similar morphology and morphometrics, but differed molecularly 5-7.5%, and were no more than 94% similar to any other sequence in GenBank. Together with having different hosts, these data supported the diagnosis of the parasites as distinct, novel species. Maximum likelihood and Bayesian analyses showed that H. tucunarei n. sp., H. tapajoensis n. sp., and H. jariensis n. sp. plus Henneguya paraensis (which parasitizes Cichla temensis) formed a well-supported sub-clade of Henneguya parasites of cichlids from the Amazon basin, in a lineage sister to those in characiforms hosts. Our analysis was consistent with previous studies that suggest that aquatic environment and vertebrate host group are the strongest correlates with phylogenetic signals in the Myxobolidae.

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We describe a new freshwater myxosporean species Ceratomyxa gracillima n. sp. from the gall bladder of the Amazonian catfish Brachyplatystoma rousseauxii; the first myxozoan recorded in this host. The new Ceratomyxa was described on the basis of its host, myxospore morphometry, ssrDNA and internal transcribed spacer region (ITS-1) sequences. Infected fish were sampled from geographically distant localities: the Tapajós River, Pará State, the Amazon River, Amapá State and the Solimões River, Amazonas State. Immature and mature plasmodia were slender, tapered at both ends, and exhibited vermiform motility. The ribosomal sequences from parasite isolates from the three localities were identical, and distinct from all other Ceratomyxa sequences. No population-level genetic variation was observed, even in the typically more variable ITS-1 region. This absence of genetic variation in widely separated parasite samples suggests high gene flow as a result of panmixia in the parasite populations. Maximum likelihood and maximum parsimony analyses placed C. gracillima n. sp. sister to Ceratomyxa vermiformis in a subclade together with Ceratomyxa brasiliensis and Ceratomyxa amazonensis, all of which have Amazonian hosts. This subclade, together with other Ceratomyxa from freshwater hosts, formed an apparently early diverging lineage. The Amazonian freshwater Ceratomyxa species may represent a radiation that originated during marine incursions into the Amazon basin that introduced an ancestral lineage in the late Oligocene or early Miocene.

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A new species of Ceratomyxa parasitizing the gall bladder of Cichla monoculus, an endemic cichlid fish from the Amazon basin in Brazil, is described using morphological and molecular data. In the bile, both immature and mature myxospores were found floating freely or inside elongated plasmodia: length 304 (196-402) µm and width 35.7 (18.3-55.1) µm. Mature spores were elongated and only slightly crescent-shaped in frontal view with a prominent sutural line between two valve cells, which had rounded ends. Measurements of formalin-fixed myxospores: length 6.3±0.6 (5.1-7.5) µm, thickness 41.2±2.9 (37.1-47.6) µm, posterior angle 147°. Lateral projections slightly asymmetric, with lengths 19.3±1.4µm and 20.5±1.3µm. Two ovoid, equal size polar capsules, length 2.6±0.3 (2-3.3) µm, width 2.5±0.4 (1.8-3.7) µm, located adjacent to the suture and containing polar filaments with 3-4 turns. The small subunit ribosomal DNA sequence of 1605 nt was no more than 97% similar to any other sequence in GenBank, and together with the host, locality and morphometric data, supports diagnosis of the parasite as a new species, Ceratomyxa brasiliensis n. sp. Maximum parsimony and maximum likelihood analyses showed that C. brasiliensis n. sp. clustered within the marine Ceratomyxa clade, but was in a basally divergent lineage with two other freshwater species from the Amazon basin. Our results are consistent with previous studies that show Ceratomyxa species can cluster according to both geography and host ecotype, and that the few known freshwater species diverged from marine cousins relatively early in evolution of the genus, possibly driven by marine incursions into riverine environments.

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Myxobolus prochilodus and Myxobolus porofilus are parasites of Prochilodus lineatus, an economically important South American fish found in La Plata and Paraiba do Sul river basins. This study focusing on parasite-host interaction provides an ultrastructural and phylogenetic analysis, the latter based on ssrDNA sequencing of these parasites respectively infecting the gill filaments and fins of P. lineatus taken from the Mogi Guaçu River, São Paulo, Brazil. A total of 13 adult specimens were examined in this study. The prevalence of infection was 7.69 % for M. prochilodus and 15.38 % for M. porofilus. Phylogenetic analysis showed M. prochilodus and M. porofilus clustered in a subclade composed of parasites of the Prochilodontidae family. In M. prochilodus infecting gill filaments, where cellular degeneration in the epithelium was observed, the plasmodia were surrounded by a capsule composed of layers of fibrocyte-like cells, with cellular projections joined to the projections of other fibrocyte-like cells by desmosomes, and more externally typical fibroblast layers. Some granular leukocytes were seen interspersed among these layers. In M. porofilus infecting the fins, the capsule of connective tissue was represented only by loosely arranged collagen fibers, and no granular leucocytes were observed. Finally, several unusual vacuoles with filamentous content and some characteristics usually described as degenerative alterations, as myelin figure, were noted in plasmodia and pansporoblasts of both myxosporean species. The possible influence of inflammatory response and xenobiotics was considered to be the explanation for the alterations observed in Myxobolus species and its host.

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Thelohanellus kitauei Egusa & Nakajima, 1981 is a common parasite infecting the intestine of common carp Cyprinus carpio L., resulting in mass mortality or loss of economic value of cultured carp. In the present study, T. kitauei infecting host skin was detected. The morphological, molecular and histological data of this parasite in the new organ record are presented. Morphological analysis showed the current specimen morphologically similar to T. kitauei from the intestine. Despite the spore length and polar capsule length of the current specimen larger than those of T. kitauei from the intestine, ranges of dimensions overlap, which is more suggestive of intraspecific variation than distinct species. BLAST search revealed that the present small subunit ribosomal DNA gene sequence is identical to those of T. kitauei. Histologically, most of spores distributed in the stratum spongiosum of dermis, and some spores in the strata compactum of host skin were also observed. Above all, both morphology and molecular analysis indicated that the current species from the skin of common carp is conspecific with T. kitauei from the intestine of carp and organ habitats transfer of T. kitauei from host intestine to skin may have occurred.

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RESUMEN

Novel molecular data are presented to resolve the long-standing issue of the non-monophyly of the elasmobranch-hosted tapeworm order Tetraphyllidea relative to the other acetabulate eucestode orders. Bayesian inference analyses of various combinations of full ssrDNA, and full or partial lsrDNA (D1-D3), sequence data, which included 134 species representing 97 genera across the 15 eucestode orders, were conducted. New ssrDNA data were generated for 82 species, partial lsrDNA data for 53 species, and full lsrDNA data for 29 species. The monophyly of each of the elasmobranch-hosted orders Cathetocephalidea, Litobothriidea, Lecanicephalidea and Rhinebothriidea was confirmed, as was the non-monophyly of the Tetraphyllidea. Two relatively stable groups of tetraphyllidean taxa emerged and are hereby designated as new orders. The Onchoproteocephalidea n. ord. is established to recognise the integrated nature of one undescribed and 10 described genera of hook-bearing tetraphyllideans, previously placed in the family Onchobothriidae, with the members of the order Proteocephalidea. The Phyllobothriidea n. ord. is established for a subset of 12 non-hooked genera characterised by scoleces bearing four bothridia each with an anterior accessory sucker; most parasitise sharks and have been assigned to the Phyllobothriidae at one time or another. Tentative ordinal placements are suggested for eight additional genera; placements for the remaining tetraphyllidean genera have not yet emerged. We propose that these 17 genera remain in the "Tetraphyllidea". Among these, particularly labile across analyses were Anthobothrium, Megalonchos, Carpobothrium, Calliobothrium and Caulobothrium. The unique association of Chimaerocestus with holocephalans, rather than with elasmobranchs, appears to represent a host-switching event. Both of the non-elasmobranch hosted clades of acetabulate cestodes (i.e. Proteocephalidea and Cyclophyllidea and their kin) appear to have had their origins with elasmobranch cestodes. Across analyses, the sister group to the clade of "terrestrial" cestode orders was found to be an elasmobranch-hosted genus, as was the sister to the freshwater fish- and tetrapod-hosted Proteocephalidea. Whilst further data are required to resolve outstanding nomenclatural and phylogenetic issues, the present analyses contribute significantly to an understanding of the evolutionary radiation of the entire Cestoda. Clearly, elasmobranch tapeworms comprise the backbone of cestode phylogeny.

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