Biology of Bothrimonus (= Diplocotyle) (Pseudophyllidea: Cestoda): Ecology, Life Cycle, and Evolution; a Review and Synthesis

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Abstract
An integrated account of the ecology and life cycle of Bothrimonus is synthesized from the literature, experiment, and observation of larval and adult stages of the parasite. Bothrimonus has a circumpolar distribution in boreal or subarctic conditions. As an adult it is found in a wide variety of fish hosts which feed in littoral or estuarine waters. Adult worms have a short life-span in the fish host and occur only seasonally. A single intermediate host appears to be involved in the life cycle and this is most commonly a brackish-water gammarid. The operculate eggs, which contain unciliated larvae, possess characteristic polar filaments which have an adhesive function. Entry is almost certainly by ingestion of eggs by the gammarid host; the larval forms often attain sexual maturity within the gammarid host with accompanying production of apparently normal eggs. Although it has been possible to infect fish hosts experimentally with larval stages, it has not yet been possible to infect gammarids with eggs obtained either from neotenic individuals in gammarids or from adults in the fish host.The life cycle of Bothrimonus indicates closer affinities of the Cyathocephalidae with caryophyllaeid than with other pseudophyllidean tapeworms. Distribution is discussed in terms of its evolution and a freshwater origin in the Ponto–Aralo–Caspian province is proposed.

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