Empirical studies have demonstrated that Ichthyophonus is transmitted among piscivores via consumption of infected prey; however, this unidirectional mode of transmission from small to large fish cannot sustain Ichthyophonus within a population. To circumvent this problem, Ichthyophonus evolved an infective waterborne cell that has been shown to infect both fresh-and saltwater fish. Successful transmission of Ichthyophonus via a waterborne stage is linked to the proximity of infected and susceptible individuals, a condition met when the two groups occupy the same habitat. It is posited that this occurs during annual inshore migrations when herring (Clupea spp.) enter areas occupied by infected demersal predators. A plausible transmission scenario is that during inshore excursions, planktivores are exposed to infective waterborne cells shed by demersal piscivores. Once planktivores are infected, the parasite is recycled when consumed by predators. This model is supported by reports of ichthyophoniasis increasing in herring populations at the time of inshore migration. The possible role of an intermediate host for Ichthyophonus, as well as evidence for a free-living stage in its life cycle, is presented.
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