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Relatedness, host specificity and richness of the genus Ceratomyxa (Myxozoa: Myxosporea) in teleost fishes

The genus Ceratomyxa Thélohan, 1892 is one of the largest myxozoan genera, the species of which almost always infect the gall bladder of marine teleosts. Although there are over 180 species known globally, prior to this study only three had been described from Australia. This study explores the systematics, host specificity and species richness of Ceratomyxa species from a diverse range of Queensland marine fishes. The first part of this thesis deals with the issues associated with spore based taxonomy and incorporates the first molecular studies to investigate host specificity and radiation in three common families of Great Barrier Reef (GBR) fishes. Twenty-two new species were described in these chapters, and although they were superficially similar in morphology all were genetically distinct. A focus on Ceratomyxa spp. infecting labrid, pomacentrid and serranid fish revealed tight host specificity of parasite species, confirmed the presence of multiple parasite species in a single host and revealed that no parasite radiation had occurred that could be associated with co-evolution with host families. Exploration of intra- and inter-specific variation by sequencing of multiple replicates from host/parasite combinations allowed interpretation of species boundaries within the system. Ten additional species described in Chapter 5 supported the findings from the labrid, pomacentrid and serranid studies but broadened the host range studied. In total 32 Ceratomyxa species were described from Queensland fishes increasing the number of described species in the genus by 15%. This study also reports on undescribed species collected from a further 70 host species and broadens the known host range to seven families of fishes. A checklist of bivalvulidans from marine teleosts in Australia is compiled of described and undescribed species and highlights the abundance of Ceratomyxa relative to other gall bladder dwelling myxosporeans. A phylogeny of the Ceratomyxa based on SSU rDNA analyses explored the taxonomic integrity of the genus. In general, the morphological diagnostic characters that divide the Myxozoa into genera are not well supported by molecular phylogenetic analyses. The relationship between 42 Ceratomyxa spp. and 36 other marine myxosporeans was examined using Bayesian inference, maximum likelihood and maximum parsimony. Results indicate Ceratomyxa is one of the most cohesive lineages within the Myxozoa and that the freshwater fish parasite, C. shasta, does not represent an independent lineage as suggested in previous studies. The strict host specificity of Ceratomyxa species prompted the investigation of species richness on the GBR. Species accumulation curves were used to explore species richness by using Choa2 and Jackknife1 estimators. The estimates suggested Ceratomyxa is almost as rich as their teleost hosts and that 1,600 species could be present on the GBR and as many as 15,000 species may exist globally. As an unexpected result, Ceratomyxa may be the richest metazoan genus in the sea. In light of what the study revealed, a revision of a second genus within the family Ceratomyxidae, Leptotheca, is presented. The boundaries between Ceratomyxa and Leptotheca were unclear. The diagnostic characters that separate these genera (length to thickness ratios) were found to be plastic and the type species of Leptotheca ultimately fitted the diagnosis for Ceratomyxa. To eliminate confusion between these genera and also between Sphaerospora and Leptotheca, the genus Leptotheca was considered invalid, with all species appropriately assigned to other genera based on morphological and biological characters and supported by genetic evidence.