Kudoa thyrsites is a myxosporean endoparasite that infects fish, frequently resulting in myoliquefaction, impacting the texture and eating quality of fish muscle. The aim of this study was to investigate the effects of K. thyrsites infection on the proximate composition and texture parameters of raw fillets from two South African economically important species: Cape hake (Merluccius capensis) and Cape dory (Zues capensis). Infection status was determined using real-time quantitative polymerase chain reaction. Association of Official Agricultural Chemists (AOAC) methods were used to determine moisture, protein, fat and ash of raw fillets. Texture Profile Analysis measured springiness and gumminess, using an Instron. The study revealed host species-specific impacts. For Cape hake, K. thyrsites infection showed no statistically significant effect on proximate composition or texture (springiness, gumminess). In contrast, Cape dory demonstrated a significant increase in moisture content in infected (76.48%) compared to uninfected samples (75.22%). Furthermore, infected Cape dory exhibited significantly lower springiness (5.98%) compared to uninfected samples (6.30%), correlating with a softer texture. These findings are the first for Cape dory and of interest to the fishing industry for developing quality assessment tools.

Kudoa thyrsites is a myxosporean parasite that infects the skeletal muscle of various teleost fish species globally. Severe infections lead to 'soft flesh' in fish fillets, resulting in food spoilage and subsequent discard. While K. thyrsites has previously been identified in migratory Atlantic mackerel in the northern Northeast Atlantic Ocean, it has not been observed in resident or farmed fish species in Norwegian waters until now. This study presents the first evidence of K. thyrsites infection and the associated 'soft flesh' condition in resident commercially important gadoid species from Norwegian waters, including Norwegian coastal cod (NCC), Northeast Arctic cod (NEA) and tusk. Molecular analyses confirmed the parasitic infection in 'soft flesh'-affected fish sampled from multiple coastal locations in Norway. The life cycle of Kudoa remains unknown but likely involves an alternating annelid host as in other myxosporeans. These findings in resident hosts suggest that the parasite completes its life cycle also at higher latitudes, in northern Norway. Consequently, there is a risk for the Norwegian fishing industry, as the effect of the parasite on fish fillet texture can occasionally occur and impact both consumer acceptance and industry revenues.

Small pelagic fishes represent one of the most important food resources off the Northwest coast of Africa. Despite their economic significance, little is known about the infections with flesh invading myxosporean parasites of genus Kudoa (Cnidaria, Myxozoa). Heavy infections in the flesh may be associated with post-mortem myoliquefaction, commonly known as ‘soft flesh’. This condition may reduce the quality and marketability of the fish fillet, resulting in both economic losses to the fishing industry and loss of consumer confidence. In this study, we investigated Kudoa-induced ‘soft flesh’ occurrence in European anchovy Engraulis encrasicolus, European pilchard Sardina pilchardus, and Atlantic chub mackerel Scomber colias caught in 2019 off the Moroccan Atlantic coast. Five hundred specimens of each fish species were examined for ‘soft flesh’ by texture testing and visual inspection 48 h post-catch. ‘Soft flesh’ occurred in 0.2 % of the European anchovies, 1.4 % of the European pilchard, and in 4.4 % of the Atlantic chub mackerel. Microscopic examination of muscle samples revealed that ‘soft flesh’-affected fish were infected with myxospores of K. thyrsites-like morphotype. Analysis of the kudoid SSU rDNA sequence obtained from European pilchard and the Atlantic chub mackerel identified these as K. thyrsites (100 % identity), whereas analysis of the sequence from European anchovy identified the presence of K. encrasicoli (100 % identity). Even if there are no known human health consequences associated with the ingestion of these Kudoa species, the unsightly appearance of some infected fillets is a food quality issue, that can eventually lead to reduced marketability and value.

In March and April 2018, we carried out a survey of myxosporean parasites in coastal fishes collected from Nha Trang Bay in Vietnam's East Sea. Of the 159 fish specimens, 18 fish species were represented, belonging to 10 families. In 8 host species, 7 myxosporean species were found, representing the genera Kudoa and Unicapsula. Two of these species were new to science: Kudoa igori n. sp. from the gallbladder of Longarm mullet Osteomugil cunnesius (Valenciennes, 1836) (prevalence 10%) and Kudoa borimiri n. sp. from skeletal muscles of Longarm mullet and Longfinned mullet Osteomugil perusii (Valenciennes, 1836) (prevalence 30% and 60%, respectively). Vegetative stages were not found. Spores of K. igori n. sp. were small, orbicular to quadrate in apical view, and with four equal valves. In lateral view, spores were shallowly ovoid with a slightly protruding anterior pole. The 4 polar capsules were slightly unequal sizes and were oriented toward the apex of the spore. Dimensions were as follows: spore length 4.56 ± 0.22 (4.18-4.56), thickness 4.42 ± 0.39 (3.55-5.13), width 5.74 ± 0.46 (4.66-6.50), length of biggest polar capsule 1.38 ± 0.14 (1.1-1.65), middle 1.28 ± 0.12 (1.03-1.53), smallest 1.13 ± 0.11 (0.94-1.30), width of biggest polar capsule 1.03 ± 0.14 (0.83-1.4), middle 0.92 ± 0.12 (0.76-1.08), smallest 0.74 ± 0.12 (0.55-0.94). Spores of K. borimiri n. sp. had four equal valves with slightly rounded ends and were quadrate in apical view. In lateral view, spores were broadly deltoid. Four equal-sized polar capsules opened at the apex of the spore. In O. cunnesius, spore dimensions were as follows: length 3.51 ± 0.15 (3.34-3.92), thickness 3.65 ± 0.38 (2.79-4.21), width 4.97 ± 0.37 (4.13-5.97), length of the polar capsules 1.23 ± 0.18 (0.99-1.57), width 0.88 ± 0.07 (0.70-1.00). Overlapping dimensions for K. borimiri n. sp. spores were also found in O. perusii. Other known myxozoan species we encountered were Kudoa thyrsites and Kudoa whippsi, which had not previously been reported from the East Sea and Vietnam. We also encountered Kudoa monodactyli, which had not previously been reported from Nha Trang Bay. In addition, we report 2 additional species, a Kudoa sp. and a Unicapsula sp., that were not attributable to previously described myxozoans and need further investigation to completely characterize.

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 encrasicolin. 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.

Kudoa thyrsites is a myxosporean parasite (Cnidaria, Myxozoa) that infects the skeletal and cardiac muscle of Northeast Atlantic (NEA) mackerel (Scomber scombrus). Heavy infections are associated with post-mortem myoliquefaction of the host skeletal muscle which reduces the quality of the fish product. The biological infection characteristics of the parasite in NEA mackerel are poorly known. This study examined the distribution of K. thyrsites in various organs of NEA mackerel from the northern North Sea, and elucidates the relationship between density of infection, developmental stage and parasite distribution in the musculature, and the extent of visible flesh myoliquefaction. Quantitative polymerase chain reaction (qPCR) data showed that K. thyrsites is unevenly distributed in the somatic musculature of the fish host, with highest density in the anterior ventral muscle sections—the belly flaps. A weak positive correlation was observed between the level of myoliquefaction and the parasite density in the fish host muscle. This relationship was also reflected by the amount and distribution of parasite developmental stages seen during histological examinations. Histological findings indicate an association between the dispersion of free myxospores and the level of myoliquefaction of the fish host muscle. Visceral organs were also found infected using qPCR, although at lower densities compared to the musculature.

Atlantic mackerel Scomber scombrus is an important migratory fish in Faroese waters and the Northeast Atlantic at large. Similar to other fish species from marine environments, mackerel can be infected by the myxozoan parasite Kudoa thyrsites, leading to myoliquefaction that renders the fish commercially unviable. Despite the ecological and economic significance of K. thyrsites as a parasite, little is known regarding its prevalence in the Faroese mackerel fishery. Prior to analysing field samples, we examined 104 samples of Atlantic mackerel selected from a Faroese processing plant on the basis of visible soft-tissue. Using microscope smears we observed K. thyrsites in 98% of the soft-tissue mackerel specimens and a direct comparison with qPCR demonstrated strong agreement between the two techniques (Φ = 0.429, p<0.01; Fisher’s exact test). We used qPCR to analyze a total of 594 Atlantic mackerel specimens, collected from Faroese fishing grounds during 2017 and 2018. Overall prevalence was 4.1% (95% confidence interval; 2.5-5.7%) and ranged from 0-14% at different locations. Infection by K. thyrsites was documented in fish as young as 2 years (195g) and up to 14 years (615g) of age. A logistic regression model indicated the odds of infection were statistically associated with sampling location and month, but not gender, age, weight or length, although a Chi-squared test on age categories showed the oldest fish (>10 years) had highest prevalence (10.8%, CI: 4.2-17.5%). In order to investigate potential infection pathways we performed qPCR analysis on eDNA in water samples collected from the Faroese Plateau. However, we found no molecular evidence for K. thyrsites spores in Faroese plankton assemblages. Our data support the prevailing hypothesis that both the alternating invertebrate host and infection of Atlantic mackerel by K. thyrsites occurs in more southern waters.

Northeast Atlantic (NEA) mackerel (Scomber scombrus, Scombridae) represents an economically important target for the Norwegian pelagic fishing industry. Despite the commercial significance of NEA mackerel, little is known about the infections with the myxosporean parasite Kudoa thyrsites (Kudoidae). The parasite may cause post-mortem myoliquefaction of the fish skeletal muscle and therefore reduce the quality of the fish product. In this study, we examined ‘soft flesh’ occurrence in commercial size groups of NEA ‘autumn mackerel’ caught between 2007 and 2020, and investigated the prevalence and density of K. thyrsites (qPCR) and how they related to the occurrence of ‘soft flesh’. The present study is the first long-term investigation of the occurrence of K. thyrsites-induced ‘soft flesh’ in NEA mackerel. After appearing stable for over a decade, the ‘soft flesh’ occurrence increased three- to six-fold in 2019 and 2020. This increase, together with the findings that ‘soft flesh’ seems primarily to affect the commercially most valuable mackerel size group (>400 g), may have important implications for the fishing industry and the fishery management. Molecular analysis (qPCR) suggests that the prevalence of K. thyrsites is substantially higher than ‘soft flesh’ occurrence. The majority (87.4%, n = 76/87) of infected mackerel did not develop ‘soft flesh’ and only individuals with high parasite density in the musculature (12.6%, n = 11/87) showed the condition. Therefore, qPCR analyses should be used for estimating the prevalence of K. thyrsites in fish. The method may also be used to assess the risk of NEA mackerel to develop post-mortem ‘soft flesh’.

Kudoa thyrsites is a myxozoan parasite of marine fish with a global distribution. In British Columbia (BC), Canada, severe infections are associated with an economically significant degradation of fillet quality in farmed Atlantic salmon. Exposures to naturally occurring actinospores at a coastal research laboratory were used to test the hypothesis that the prevalence and severity of K. thyrsites infections acquired by exposure of Atlantic salmon to seawater (SW) of various depths are not different. In Expt 1, fish were exposed to SW from 1, 7 or 13 m below the surface. Following exposure to deeper-sourced SW, the prevalence of K. thyrsites, determined from microscopic examination of muscle histology sections, was greater in all 4 trials and the severity of infection was greater in 2 trials. In Expt 2, infections were compared over time among salmon held in tanks supplied with deep-sourced SW (raw or UV-irradiated) or in a surface net-pen. The infection was observed in 35 of 40 fish sampled between 3 and 6 mo after tank exposure to raw SW. Coincidentally, the parasite was observed in 4 of 40 fish maintained in the net-pen. No consistent association of the parasite infection was observed with temperature; however, reduced salinity and solar radiation were not ruled out as factors which may reduce the risk of infection from surface SW. These findings require verification at commercial aquaculture sites in BC, as they will inform considerations related to farm siting and net-pen configuration.

Rapid expansion of salmon aquaculture has resulted in high-density populations that host diverse infectious agents, for which surveillance and monitoring are critical to disease management. Screening can reveal infection diversity from which disease arises, differential patterns of infection in live and dead fish that are difficult to collect in wild populations, and potential risks associated with agent transmission between wild and farmed hosts. We report results from a multi-year infectious-agent screening program of farmed salmon in British Columbia, Canada, using quantitative PCR to assess presence and load of 58 infective agents (viruses, bacteria, and eukaryotes) in 2931 Atlantic salmon (Salmo salar). Our analysis reveals temporal trends, agent correlations within hosts, and agent-associated mortality signatures. Multiple agents, most notably Tenacibaculum maritimum, were elevated in dead and dying salmon. We also report detections of agents only recently shown to infect farmed salmon in BC (Atlantic salmon calicivirus, Cutthroat trout virus-2), detection in freshwater hatcheries of two marine agents (Kudoa thyrsites and Tenacibaculum maritimum), and detection in the ocean of a freshwater agent (Flavobacterium psychrophilum). Our results provide information for farm managers, regulators, and conservationists, and enable further work to explore patterns of multi-agent infection and farm/wild transmission risk.

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

Myxosporean parasites of the genus Kudoa are fish parasites of great economic importance, as some species can affect the fish fillet quality by producing macroscopic cysts or generating post mortem myoliquefaction, commonly referred to as ‘soft flesh’. Kudoa mirabilis is a ‘soft flesh’-inducing species originally described based on morphology in the musculature of Trichiurus lepturus from the Indian Ocean. An integrative morphological and genetic characterization of K. mirabilis from the type host caught off the coast of Tanzania is here provided. The spores were stellate with four unequal polar capsules, showing similarities to Kudoa thyrsites. For comparative and validation purpose, K. mirabilis was compared morphologically and genetically with K. thyrsites reference isolates, including new obtained samples from the type host Thyrsites atun caught in the SE Atlantic Ocean. Morphological analyses of spores revealed key diagnostic characters clearly distinguishing the two Kudoa species. Phylogenetic analyses based on SSU and LSU rRNA genes demonstrated that K. mirabilis is a distinct and valid species, representing a sister group to a K. thyrsites subclade that comprises several isolates from Japan and one single isolate from South Africa. This finding raises questions about the true diversity likely hidden in the K. thyrsites complex.

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

Prevalence of infection with the myxozoan parasite Kudoa thyrsites is not well documented in sardine Sardinops sagax, the Cape hakes Merluccius capensis and M. paradoxus, and kingklip Genypterus capensis. Quantitative polymerase chain reaction assays were used to determine the prevalence of K. thyrsites in wild-caught S. sagax (n = 296), Merluccius spp. (n = 162) and G. capensis (n = 70), harvested in 2015. Sardinops sagax were harvested from the east, south and west coasts of South Africa, whereas M. capensis, M. paradoxus and G. capensis were taken from the west coast only. Relationships between K. thyrsites infection prevalence and fish capture locality, season, sex and size were determined with Chi-square tests. Prevalence was highest in S. sagax (91%), followed by M. capensis and M. paradoxus (87%) and G. capensis (40%). Prevalence was independent of sex of the host for all fish species. In the case of S. sagax only, the prevalence of infection was independent of locality of capture but depended on season, with prevalence lowest during spring and highest during autumn. No significant relationship (p > 0.05) was found between infection prevalence and host size for any of the fish species.

Kudoa thyrsites is a cosmopolitan myxozoan parasite of marine fish. The infection causes an economically important myoliquefaction in farmed Atlantic salmon in British Columbia, Canada. Laboratory exposure of Atlantic salmon smolts to infectious seawater was used to test the hypothesis that infection with K. thyrsites is more severe in age-matched, smaller salmon. In each of 2 trials approximately 4 mo apart, smolts were graded into small (80 and 68 g), medium (117 and 100 g) and large (142 and 157 g) initial weight groups (IWGs) and concurrently exposed to infectious seawater. The effects of IWG and time on fish size and infection severity were assessed by linear mixed-effects models. The fish were screened for infection by histological examination at intervals following exposure. Increases in mean length and weight were statistically significant in all IWG during both trials. The infection was detected in fish in both trials, and in Trial 2, the prevalence was significantly greater in larger fish 1000 degree-days (DD) after exposure. The severity of infection (plasmodia mm-2 muscle) was significantly higher in larger smolts: between medium and large IWGs at 2500 DD in Trial 1 and between small and medium IWGs at 1500 and 2000 DD in Trial 2. The hypothesis is rejected and possible explanations for the greater occurrence of K. thyrsites in larger smolts are discussed.

In response to concerns that novel infectious agents were introduced through the movement of eggs as Atlantic salmon aquaculture developed in British Columbia (BC), Canada, we estimated the prevalence of infectious agents in archived return-migrating Sockeye salmon, from before and during aquaculture expansion in BC (1985-94). Of 45 infectious agents assessed through molecular assays in 652 samples, 23 (7 bacterial, 2 viral and 14 parasitic) were detected in liver tissue from six regions in BC. Prevalence ranged from 0.005 to 0.83 and varied significantly by region and year. Agent diversity ranged from 0 to 12 per fish (median 4), with the lowest diversity observed in fish from the Trans-Boundary and Central Coast regions. Agents known to be endemic in Sockeye salmon in BC, including Flavobacterium psychrophilum, Infectious haematopoietic necrosis virus, Ceratonova shasta and Parvicapsula minibicornis, were commonly observed. Others, such as Kudoa thyrsites and Piscirikettsia salmonis, were also detected. Surprisingly, infectious agents described only recently in BC salmon, Ca. Branchiomonas cysticola, Parvicapsula pseudobranchicola and Paranucleospora theridion, were also detected, indicating their potential presence prior to the expansion of the aquaculture industry. In general, our data suggest that agent distributions may not have substantially changed because of the salmon aquaculture industry.

Heart and skeletal muscle inflammation (HSMI) is an emerging disease of marine-farmed Atlantic Salmon (Salmo salar), first recognized in 1999 in Norway, and later also reported in Scotland and Chile. We undertook a longitudinal study involving health evaluation over an entire marine production cycle on one salmon farm in British Columbia (Canada). In previous production cycles at this farm site and others in the vicinity, cardiac lesions not linked to a specific infectious agent or disease were identified. Histologic assessments of both live and moribund fish samples collected at the farm during the longitudinal study documented at the population level the development, peak, and recovery phases of HSMI. The fish underwent histopathological evaluation of all tissues, Twort’s Gram staining, immunohistochemistry, and molecular quantification in heart tissue of 44 agents known or suspected to cause disease in salmon. Our analysis showed evidence of HSMI histopathological lesions over an 11-month timespan, with the prevalence of lesions peaking at 80–100% in sampled fish, despite mild clinical signs with no associated elevation in mortalities reported at the farm level. Diffuse mononuclear inflammation and myodegeneration, consistent with HSMI, was the predominant histologic observation in affected heart and skeletal muscle. Infective agent monitoring identified three agents at high prevalence in salmon heart tissue, including Piscine orthoreovirus (PRV), and parasites Paranucleospora theridion and Kudoa thyrsites. However, PRV alone was statistically correlated with the occurrence and severity of histopathological lesions in the heart. Immunohistochemical staining further localized PRV throughout HSMI development, with the virus found mainly within red blood cells in early cases, moving into the cardiomyocytes within or, more often, on the periphery of the inflammatory reaction during the peak disease, and reducing to low or undetectable levels later in the production cycle. This study represents the first longitudinal assessment of HSMI in a salmon farm in British Columbia, providing new insights on the pathogenesis of the disease.

We developed a PCR assay targeting the 28S rDNA of Kudoa iwatai (Multivalvulida: Myxozoa) and investigated the prevalence of infection in rock bream Oplegnathus fasciatus, which is commercially an important aquaculture species in Korea, with this assay. Detection limit of the PCR assay was 2.5fg/μl with plasmid DNA and 8.6×103 spores/ml with purified spores, respectively. This PCR assay did not amplify DNA of other Kudoa species (Kudoa septempunctata, Kudoa lateolabracis, Kudoa thyrsites) tested. Sliced muscles of whole body from 318 rock bream (wild and cultured) were examined by this PCR assay and also with the naked eyes. All of the wild fish did not produce amplicons nor did harbor visible Kudoa cysts (0/70). Three of the cultured fish were PCR-positive and also harbored visible Kudoa cysts (3/248, 1.2%). The sequences of amplicons (574bp) were 100% identical with those of the K. iwatai already registered in Genbank. When the visceral organs of these three fish were examined, visible cysts were not found, but one stomach sample was found to be PCR-positive. There was no difference in the prevalence of infection estimated by PCR assay and the presence of visible Kudoa cysts in our samples. This is thought to be because the development of K. iwatai is already completed and only mature Kudoa cysts existed in our samples.

Kudoa septempunctata (Myxosporean: Multivalvulida) is known as a cause of foodborne disease associated with consumption of raw flesh of the olive flounder (Paralichthys olivaceus). Knowledge of its life cycle, particularly alternate annelid hosts and reservoirs or susceptible fish hosts in natural waters, may facilitate disease control in aquaculture farms. Our recent survey of myxosporean infection in monacanthid fish in natural waters around Japan revealed infection with three kudoid species prevalent in the olive flounder, i.e., K. septempunctata, Kudoa thyrsites, and Kudoa shiomitsui. Of the 51 black scrapers (Thamnaconus modestus) examined, five fish were infected: two fish with K. septempunctata and three with K. thyrsites. One of the fish infected with K. septempunctata was also infected with a K. thyrsites-like species. One of the 17 threadsail filefish (Stephanolepis cirrhifer) and two of four unicorn leatherjackets (Aluterus monoceros) were parasitized with K. shiomitsui. Three modest filefish (Thamnaconus modestoides) had no kudoid infection. K. septempunctata from a black scraper fished in the Inland Sea of Japan off Yamaguchi had 6-8 (predominantly 7) shell valves/polar capsules, whereas K. septempunctata found in another black scraper from the Sea of Japan off Tottori had 5 or 6 (predominantly 6). However, the two isolates displayed identical 18S and 28S ribosomal RNA gene (rDNA) nucleotide sequences, which were also identical to the isolates from the olive flounder. K. thyrsites from the Inland Sea of Japan off Yamaguchi and Sea of Japan off Tottori and K. shiomitsui from the Sea of Japan off Shimane and western Pacific Ocean off Kochi were also morphologically and genetically characterized. They were found to be coincident with the previous reports from olive flounders. Furthermore, the K. thyrsites-like species found in a black scraper from the Inland Sea of Japan off Yamaguchi was morphologically and genetically characterized; a new species, Kudoa parathyrsites n. sp., is erected for this species. The relationships of the new species with K. thyrsites and related species as well as those of K. shiomitsui with Kudoa pericardialis and related species parasitizing the pericardium are briefly discussed.

Resolution of Kudoa thyrsites infection is associated with infiltration of MHIIβ+ cells in Atlantic salmon, Salmo salar