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Two new species of myxosporeans are described from the gallbladders of estuarine stonefish, Synanceia horrida, and reef stonefish, Synanceia verrucosa, from localities off Cairns, in tropical north Queensland and in Moreton Bay in southern Queensland, Australia. Sphaeromyxa horrida n. sp. can be distinguished from congeners in the morphologically distinct "balbianii" species group within Sphaeromyxa on the basis of morphometric differences in length and width of mature spores, length and width of polar capsules, and unique small-subunit (SSU) ribosomal (rDNA) sequence composition relative to other taxa. Replicate SSU rDNA sequences generated from Sph. horrida n. sp. collected from Sy. horrida and Sy. verrucosa in tropical north Queensland and from Sy. horrida in Moreton Bay were identical, suggesting that this species is widely distributed along the east coast of Australia. Myxidium lapipiscis n. sp. can be distinguished from the majority of described Myxidium species on the basis of its relatively small mature spore size (6.1-7.9 μm long × 3.1-3.9 μm wide), and its unique SSU rDNA sequence. Specimens putatively identified as M. lapipiscis n. sp. were found in Sy. horrida from both tropical north Queensland and Moreton Bay, suggesting that this taxon is also widely distributed along the east coast of Australia. However, no molecular data were available for the specimens from tropical north Queensland for comparative genetic analyses. Bayesian inference and maximum likelihood analysis of the SSU rDNA sequences for these 2 new species revealed that Sph. horrida n. sp. formed a strongly supported clade with Sphaeromyxa zaharoni Diamant, Whipps, and Kent, 2004, which was described from the scorpaeniform, Pterois miles, from the Red Sea. This is the first report of myxozoans infecting stonefish (Synanceiidae).

During a parasitological survey, Myxidium-like spores were identified in the gall bladders of marine fishes from Australian waters. This paper describes four novel species of Ellipsomyxa Køie, 2003, three novel species of Myxidium Bütschli, 1882 and six novel species of Zschokkella Auerbach, 1910 from teleosts from Australian waters using a combination of morphological, biological and molecular characters. Phylogenetic analyses showed a monophyletic relationship of all Ellipsomyxa spp. sequences with Sigmomyxa sphaerica (Thélohan, 1895) and Myxidium queenslandicus Gunter & Adlard, 2008 as sister species to the clade. The validity of genus Sigmomyxa Karlsbakk & Køie, 2012 is discussed. In phylogenetic analyses, the novel species of Myxidium fell within the 'marine' clade of Fiala (2006). However, the novel species of Zschokkella fell within the 'freshwater' clade of Fiala (2006) and formed a distinct clade with all other sequences of Zschokkella spp. from the gall bladder of marine fish and a sequence of a species of Myxobolus Bütschli 1882, also from the gall bladder of a marine fish. This is the second distinct marine lineage to emerge within the freshwater clade.

Ceratomyxa parasites from the gall bladders of 23 species of cardinalfishes (family Apogonidae) from Australian waters were examined for their taxonomic identity and phylogenetic relatedness. We identified 15 of the 23 apogonid fish species infected with species of Ceratomyxa. Although the majority of apogonid species harboured only a single Ceratomyxa species, four were found with multiple species of Ceratomyxa. This study describes eight novel species using a combination of morphological, small subunit ribosomal DNA (SSU rDNA) and biological characters. Six Ceratomyxa species are reported from single apogonid species, while two are reported from multiple host species. Molecular data were critical in identifying several morphologically cryptic species. However, our results suggest that SSU rDNA was not capable of distinguishing all the species present in the current study system and alternative genetic markers should be investigated in the future.

A survey of the myxosporean fauna of Australian marine fishes revealed the presence of three previously unreported species of Unicapsula (Multivalvulida: Trilosporidae) from sites off Southeast Queensland, off Lizard Island on the Great Barrier Reef, Queensland, and from Jurien Bay in Western Australia. Morphometric data (spore, polar capsule and caudal appendage dimensions) combined with Bayesian inference and maximum likelihood analyses of small subunit (SSU) and large subunit (LSU) ribosomal DNA (rDNA) were used for species identification and to explore relationships among these taxa. The four species of Unicapsula for which DNA data are now available for comparative purposes (Unicapsula andersenae n. sp., Unicapsula pflugfelderi, Unicapsula seriolae and Unicapsula pyramidata) formed a well-supported monophyletic sister clade to the other major multivalvulidan group, the Kudoidae. The combined morphometric and genetic diagnostic approach identified an undescribed taxon, U. andersenae n. sp., from the muscle of Argyrosomus japonicus, Acanthopagrus australis and Eleutheronema tetradactylum off the Southeast Queensland coast and in Lutjanus russellii and Sillago ciliata off Lizard Island. Intra-specific variation within U. andersenae n. sp. varied from 2-4 (0.2-0.4%) nucleotides over the SSU region to 2-20 (0.3-3.2%) over the LSU region. Inter-specific variation between U. andersenae n. sp. and the other three species for which genetic sequence data are now available ranged from 15-66 (3-6.5%) nucleotides over the SSU region to 103-120 (17.6-21.2%) nucleotides over the LSU region. The host distribution observed here for U. andersenae n. sp. (five fish species from five different fish families) represents the broadest specificity known for a single species of Unicapsula. U. pyramidata Naidjenova & Zaika 1970, whose spore morphology and presence of caudal appendages immediately distinguish it from other species, was recovered from the nemipterid, Scolopsis monogramma, off Lizard Island. U. seriolae Lester 1982 is reported here from Yellowtail Kingfish, Seriola lalandi, from sites off Queensland and from Jurien Bay, Western Australia. Comparative genetic analyses also revealed that an unidentified species of Unicapsula from Epinephelus septemfasciatus off Japan whose rDNA sequence data are available on GenBank is consistent with U. seriolae. This suggests that U. seriolae may also exhibit low host specificity and may be distributed widely throughout the Indo-West Pacific region. In comparison to other myxozoan genera, it is clear that the species richness of Unicapsula spp. falls well below that displayed by either Ceratomyxa spp. or Kudoa spp. The discovery of a further new species of Unicapsula in Australia now brings the total worldwide number of formally described Unicapsula species to a modest 11. Nonetheless, this taxon remains of significant interest to commercial and recreational fisheries through the potential production of macroscopic pseudocysts in fish muscle and post-mortem muscle liquefaction, both of which can render fish fillets unpalatable and unmarketable.

Kudoa spp. from the musculature and intestinal mucosa of species of the teleost family Apogonidae were examined for their taxonomic identity. Two novel species are characterised: Kudoa cheilodipteri n. sp. from the musculature of Cheilodipterus quinquelineatus Cuvier, Ostorhinchus cyanosoma (Bleeker) and O. aureus (Lacépède); and Kudoa cookii n. sp. from the submucosa of the intestines of O. cookii (Macleay) only. Both species are characterised using morphology, small subunit ribosomal DNA (SSU rDNA), large subunit ribosomal DNA (LSU rDNA), and biological characters. Three new host records, O. cyanosoma, O. aureus and Apogon doederleini, and associated geographical, morphological and genetic data are also provided for Kudoa whippsi Burger & Adlard, 2010. Morphological and molecular intra-specific variation of all isolates assigned to K. whippsi is also examined. Phylogenetic analyses further support the idea that tissue tropism is a distinguishing character between morphologically similar species; species reported here display close relatedness to morphologically similar species infecting the same tissue within their hosts.

Kudoa leptacanthae n. sp. was identified within the pericardial cavities of two apogonid species, Zoramia leptacantha and Zoramia viridiventer, from waters off Lizard Island on the Great Barrier Reef. The species shows a close taxonomic affinity, both morphologically and genetically, to Kudoa shiomitsui Egusa & Shiomitsu 1983 reported from a tetraodontid and scombrid, both from Japan. The infection was at high prevalences (average = 75.8% ± 0.63, n = 343) within schools of the two host species. Different sporogonic stages ('developing' and 'fully-developed') were observed in pseudocysts within the pericardium of the hosts. Kudoa leptacanthae n. sp. appears to be a persistent infection within the Zoramia species hosts. No seasonal variation was detected for this host/parasite system and the parasite did not affect host condition, when assessed using host length/weight ratio as a coarse indicator. We hypothesise that a source of infection in close proximity to the home patch reef of apogonid schools may lead to the high prevalence and intensity of infection reported in this study., (Copyright © 2012 Elsevier B.V. All rights reserved.)

We describe a new species of myxozoan, Henneguya mauritaniensis n. sp., extracted from the arterial bulb of the bluespotted seabream, Pagrus caeruleostictus (Valenciennes, 1830), collected in Mauritanian waters. Out of the 209 individuals examined, 30.1 % were infected with this new taxon. Spore total length ranged from 15.0 to 20.5 μm with a mean of 17.9 μm. The two polar capsules were equal in size, and pyriform and caudal appendages joined until mid-length. Morphometric analysis revealed significant differences between H. mauritaniensis n. sp. and morphologically similar species from this region as well as congeners known from other sparid hosts. Phylogenetic analysis of 18 S rDNA indicated that this new species is closely related to Henneguya pagri, reported recently from Pagrus major off Japan. Bayesian inference and maximum likelihood analyses of the 18 S rDNA dataset also revealed that species of marine Henneguya reported forming pseudocysts in the hearts of their fish hosts were closely related. Histological analysis of the H. mauritaniensis n. sp. pseudocysts embedded in the arterial bulb of P. caeruleostictus suggests that these parasites may cause considerable pathology, which may impact negatively on the health of the fish host. Finally, we discussed the importance of a combination of morphological and molecular analysis for species description because of high variability in size within the same taxa.

Six novel species of Chloromyxum Mingazzini, 1890 are described using a whole evidence approach combining morphometric and molecular data, together with features of their biology. Elasmobranchs were collected in Australian waters, from the Great Barrier Reef, Queensland, off Lizard and Heron Islands; from Moreton Bay, southeast Queensland; off Hobart, Tasmania; and from the Tamar River, Launceston, Tasmania. The novel species proposed here are: Chloromyxum hemiscyllii n.sp. from Hemiscyllium ocellatum; Chloromyxum kuhlii n.sp. from Neotrygon kuhlii; Chloromyxum lesteri n.sp. from Cephaloscyllium laticeps; Chloromyxum mingazzinii n.sp. from Pristiophorus nudipinnis; Chloromyxum myliobati n.sp. from Myliobatis australis; and Chloromyxum squali n.sp. from Squalus acanthias. A seventh species from Squalus acanthias is also reported but due to limited material is not formally described. Molecular phylogenetic analyses revealed that the genus Chloromyxum is polyphyletic, and species from elasmobranchs form a well-supported sister clade, with the type species Chloromyxum leydigi, to all other congeneric species clustering within the freshwater myxosporean clade. Morphological analysis showed that elasmobranch-infecting species are predominantly pyriform shaped, have clearly thickened spore apex and possess caudal filaments, compared to other Chloromyxum species which are generally spherical or subspherical, and lack caudal filaments. These morphological and phylogenetic data provide further support for the erection of new genera, but we conservatively consider the species described in this study and other elasmobranch-infecting Chloromyxum species as Chloromyxum sensu strictu, whilst the freshwater teleost infecting and amphibian infecting species we will assign as Chloromyxum sensu lato, until more comprehensive data are available., (Crown Copyright © 2011. Published by Elsevier Ireland Ltd. All rights reserved.)

A survey of the myxosporean fauna of Australian marine fishes revealed the presence of a number of putative species of Kudoidae (Multivalvulida) forming pseudocysts between the outer meningeal layer and the outer surface of the brains of the lutjanids Caesio cuning, Lutjanus carponotatus, Lutjanus ehrenbergii and Lutjanus fulviflamma and the mugilid Liza vaigiensis from Lizard Island on the Great Barrier Reef, Australia and Lutjanus lemniscatus off Ningaloo Reef, Western Australia. Morphometric data combined with Bayesian inference and maximum likelihood analyses of small subunit (SSU) and large subunit (LSU) ribosomal DNA (rDNA) was used for species identification and to explore relationships among these taxa. The brain-infecting taxa examined here formed a well-supported clade to the exclusion of non-brain infecting species in the phylogenetic analyses. The combined diagnostic approach identified an undescribed taxon, Kudoa lemniscati n. sp., from the brain of L. lemniscatus (Perciformes: Lutjanidae) off Ningaloo Reef, Western Australia, which we describe and characterise here. K. lemniscati n. sp. can be distinguished from all other species of Kudoa based on the combination of the distinct tropism for forming pseudocysts in the brain tissue, spores with 7 or 8 equal shell valves and 7 or 8 polar capsules, spore size and the differences in the SSU and LSU rDNA sequence data relative to other kudoids. Kudoa chaetodoni was found in the lutjanids C. cuning and L. carponotatus, expanding the known host range for this species to include chaetodontids and lutjanids. L. ehrenbergii and L. fulviflamma were infected with Kudoa lethrini off Lizard Island, a parasite previously known only from lethrinids. Specimens putatively identified as Kudoa yasunagai from Liza vaigiensis and Lutjanus ehrenbergii were morphologically similar and genetically identical over the SSU rDNA dataset to previously reported specimens, but differed by 4 to 11 nucleotides over the LSU dataset from the remaining isolates examined here. While these data are not definitive, they suggest the presence of a K. yasunagai complex., (Crown Copyright © 2012. Published by Elsevier Ireland Ltd. All rights reserved.)

Parvicapsula minibicornis is a myxozoan parasite implicated in mortalities of both juvenile and pre-spawning adult salmon in the Pacific Northwest of North America. Disease severity and presentation varies between salmon species and geographical localities. To better characterize population structure of the parasite, we sought genetic markers in the P. minibicornis ribosomal RNA gene. We compared samples from California with the type specimen from British Columbia, identified sequence variations, and then sequenced 197 samples from fish, river water and the parasite's polychaete worm host. Although DNA sequences of the parasite were >98·9% similar, there was enough variation to define 15 genotypes. All genotypes were detected in fish samples, although not in all species. A single genotype only was found in sockeye and pink salmon in the Fraser River Basin, but was not detected in sockeye from the adjacent Columbia River Basin. All coho salmon, irrespective of river basin, were infected with a unique mix of 2 genotypes. These data indicated that the P. minibicornis population exhibited strong signals of structuring by both geography and salmonid host species. Particular genotypes may correlate with disease differences seen in salmon populations in the Pacific Northwest.

Five novel species, Auerbachia scomberoidi n. sp., Auerbachia chaetodoni n. sp., Auerbachia caranxi n. sp., Coccomyxa colurodontidis n. sp. and Coccomyxa gobiodoni n. sp. are described from the gall bladders of marine teleosts. These species descriptions provide the first record of Auerbachia from Australian waters. Each species is characterized morphologically, including additional measurements for Auerbachia spp. and small subunit ribosomal DNA (SSU rDNA) sequences were determined for molecular phylogenetic analyses. All 5 species were each recovered from a single (and different) species of host. Phylogenetic analyses revealed a close genetic relatedness between members of Auerbachia and Coccomyxa. Based on these phylogenetic data, on obvious paraphyly displayed by the Myxidiidae and on priority, we propose the re-establishment of the family Coccomyxidae to house all species of the genera, Coccomyxa, Auerbachia and Globospora.

Kudoid parasites are known to infect a large variety of fish. A significant proportion of Kudoa species have relatively low host specificity, with a single species able to infect multiple host species representing various host families even from different host orders. Since DNA sequences have been associated with myxosporean species characterisations, it has become far easier to determine host range of new species and validate host records from earlier descriptions. This study investigated the host specificity of a kudoid parasite, Kudoa thalassomi Adlard, Bryant, Whipps et Kent, 2005, from the Great Barrier Reef in Australia using DNA sequence analysis and morphology. The results revealed the host specificity to be broad, with K. thalassomi identified in 18 different fish species representing six different fish families. This study also compares current genetic information from different host isolates of Kudoa Meglitsch, 1947 to their host ranges recorded in existing literature. From this analysis, only half of the Kudoa species with multiple host records (27 Kudoa species) have half or more isolates that are genetically characterised, and thus specifically identified with a high confidence, from their known hosts. Only five kudoid species have genetically characterised isolates from all of their recorded hosts.

Myxosporean parasites are significant parasites of fishes not only for their apparent high diversity but also for their potential impact on fish health and/or marketability. Regardless, our knowledge of most myxosporeans, especially those found in elasmobranch hosts, is superficial. A study of multivalvulidan diversity in a range of elasmobranchs from Queensland, Western Australia and the Northern Territory (Australia) was conducted to address this knowledge gap. Specimens were collected from a total of 3 orders, 9 families and 31 species of elasmobranchs. Myxosporean infections referable to the genus Kudoa were discovered in host muscle and characterized morphologically and genetically. Both small subunit (SSU) and large subunit (LSU) rDNA sequences were used in molecular phylogenetic analyses. Kudoa spp. infected 27 of the 31 species of elasmobranchs examined, representing new records of this parasite genus in 26, of the 27, host species. Kudoids were observed in all 3 orders, and 7 out of the 9 families of elasmobranchs investigated. This paper reports the first 2 multivalvulidan species to be formally described from elasmobranchs, Kudoa hemiscylli n.sp. characterized from Hemiscyllium ocellatum (and 8 other host species) and Kudoa carcharhini n. sp. characterized from Carcharhinus cautus (and 2 other host species). Phylogenetic analyses revealed that kudoids from elasmobranchs form a separate lineage to those of teleosts, but are anchored within the overall kudoid clade.

Some Kudoa species display variations in the number of polar capsules in spores within an individual pseudocyst. Nonetheless, there is usually a dominant morphotype which forms a significant element of diagnosis. In 2007, a Kudoa isolate from whiting (spores with 5 (dominant) or 6 (minor) polar capsules) was characterized by Burger et al. (2007) as being 100% identical in SSU rDNA to Kudoa yasunagai (spores with 7 polar capsules) from a halibut, despite its obvious morphological differences. The authors hypothesized that either SSU rDNA had reached its level of resolution or that the genetic identity revealed conspecificity. To further investigate these hypotheses, SSU and LSU rDNA sequence data were coupled with principal components, correlation, and regression analyses of morphometric data from different kudoid isolates that infect brain tissue to determine the relationships between spore morphotypes and different kudoid isolates. The trends in morphometrics between the spores of particular isolates were so similar that it was concluded that the molecular results did indicate conspecificity rather than SSU reaching its level of resolution. This phenotypic influence on a significant diagnostic character within the Kudoidae has a major impact on the diagnosis of this, and potentially other, pathogenic species.

Kudoid parasites are significant pathogens of marine fish. In the past, specific identification has been difficult due to a paucity of detailed morphological and biological information provided in the original description of some species. However, the introduction of DNA analysis has dramatically improved diagnosis. For morphological characterization, this paper proposes a new set of schematics including additional measurements to give a more thorough description of spore morphology, and provides evidence for uniqueness of 4 new species: Kudoa gunterae n. sp. (from 10 pomacentrid species and 1 apogonid species), K. kenti n. sp. (from 4 pomacentrid species), K. paraquadricornis n. sp. (from 4 carangid species), and K. whippsi n. sp. (from 8 pomacentrid species and 1 apogonid species). Subtle morphological differences found between closely related species were reflected in their genetics, with increased resolution provided by the large subunit, compared with that of the small subunit, of the ribosomal DNA gene region. This article proposes comprehensive requirements for species descriptions within the Kudoidae that incorporate biology, morphology, and genetic sequence.

Four new species of Ceratomyxa Thélohan, 1892 are described from the gall bladders of fishes collected off Lizard Island, Australia. These species are characterised using a combination of morphometric and molecular data. Ceratomyxa bartholomewae sp. n. is described from Hyporhamphus dussumieri (Valenciennes) (family Hemirhamphidae); C. koieae sp. n. is described from Sphyraena forsteri Cuvier (family Sphyraenidae); C. pantherini sp. n. is described from Bothus pantherinus (Rüppell) (family Bothidae) and C. reidi sp. n. is described from Chaetodon vagabundus Linnaeus (family Chaetodontidae). A fifth species from Zebrasoma veliferum (Bloch) (family Acanthuridae) is also reported but due to limited material is not formally described here.

The genus Ceratomyxa (Myxozoa: Myxosporea: Bivalvulida) contains parasites that typically infect the gall bladders of marine teleosts. Species of this genus have also been recorded from elasmobranchs, while the best known species (Ceratomyxa shasta) is a systemic pathogen of fresh water salmonid fishes. Here we characterise 10 new species of Ceratomyxa from marine teleosts using morphometric and rDNA sequence data. A phylogeny of all Ceratomyxa species for which ssrDNA sequence is available was estimated by parsimony, maximum likelihood and Bayesian analyses. Mapping host fish taxonomy, geographic locality and morphology onto the phylogenetic tree provided some concordance of these characters to groups of Ceratomyxa species, but in no case was it consistent throughout the inferred phylogeny. The position of C. shasta as a sister species to the Ceratomyxa clade contradicts previous estimates of marine myxozoan phylogeny which suggested C. shasta was an unrelated lineage. Comparative DNA sequence data is available for more than 17% of some 200 described Ceratomyxa species and the genus now represents one of the most cohesive lineages within the Myxozoa. The independent branching of all atypical Ceratomyxa species and Palliatusindecorus, indicates a review of the diagnostic characters and possible division into more genera is warranted when further data are available.

Ceratomyxa spp. from the gall-bladder of five members of the family Serranidae were examined for their taxonomic identity. This paper describes seven new ceratomyxid species, i.e. C. brayi n. sp. and C. whippsi n. sp from Cephalopholis boenak (Bloch); C. cutmorei n. sp. from Epinephelus fasciatus (Forsskål); C. gleesoni n. sp. from Plectropomus leopardus (Lacépède); C. hooperi n. sp. and C. nolani n. sp. from E. quoyanus (Valenciennes); and C. yokoyamai n. sp. from E. maculatus (Bloch). Each species is characterised morphologically and small subunit (SSU) rDNA sequences. All seven new species have so far been found in only a single host species.

Parasites of the genus Kudoa (Phylum Myxozoa) have long been known to cause considerable losses to finfish aquaculture. One such parasite species, Kudoa amamiensis, causes unsightly white cysts in the skeletal muscle of yellowtail kingfish, Seriola quinqueradiata, in Japan rendering the fillets unmarketable. The authors who characterized K. amamiensis, Egusa & Nakajima, 1980, hypothesized that yellowtail kingfish, as non-natives to the area, were accidental hosts of the parasite and that it normally infects native reef fish (damselfish, Family Pomacentridae). Since then, we have found parasites that are consistent with the description of K. amamiensis in two species of damselfish and one species of carangid fish in Australia, and it has been recorded previously in another species of reef-associated fish. Our morphometric, histological and DNA results suggest that these specimens are K. amamiensis, and are new host records for that species. Furthermore, our observations show that reef fish may act as a reservoir of myxozoan infection for commercial species, and as such should be considered an infection pathway for species in aquaculture.

Bivalvulidan parasites from the gall bladder of 31 species of damselfishes (family Pomacentridae) were examined for their taxonomic identity and their relatedness to other species of myxozoans. This paper describes 11 novel ceratomyxid species and a novel Myxidium sp. Each species is characterized morphologically and small subunit (SSU) rDNA sequences were used in molecular phylogenetic analyses. Five pomacentrid species were found to harbour multiple infections of bivalvulidan species. One species of Ceratomyxa and Myxidium were found to infect more than a single species of damselfish. Phylogenetic analyses revealed there has been no radiation of ceratomyxids that can be associated with the fish host taxon and that Myxidium queenslandicus n.sp. was more closely related to Zschokkella mugilis and Ellipsomyxa gobii than other members of the genus Myxidium.