Identification of four new Kudoa spp. (Myxozoa: Myxosporea: Multivalvulida) in commercial fishes collected from South China Sea, Atlantic Ocean, and Bering Sea by integrated taxonomic approach

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.