1 Evolution, Origins and Diversification of Parasitic Cnidarians Beth Okamura*, Department of Life Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom. Email:
[email protected] Alexander Gruhl, Department of Symbiosis, Max Planck Institute for Marine Microbiology, Celsiusstraße 1, 28359 Bremen, Germany *Corresponding author 12th August 2020 Keywords Myxozoa, Polypodium, adaptations to parasitism, life‐cycle evolution, cnidarian origins, fossil record, host acquisition, molecular clock analysis, co‐phylogenetic analysis, unknown diversity Abstract Parasitism has evolved in cnidarians on multiple occasions but only one clade – the Myxozoa – has undergone substantial radiation. We briefly review minor parasitic clades that exploit pelagic hosts and then focus on the comparative biology and evolution of the highly speciose Myxozoa and its monotypic sister taxon, Polypodium hydriforme, which collectively form the Endocnidozoa. Cnidarian features that may have facilitated the evolution of endoparasitism are highlighted before considering endocnidozoan origins, life cycle evolution and potential early hosts. We review the fossil evidence and evaluate existing inferences based on molecular clock and co‐phylogenetic analyses. Finally, we consider patterns of adaptation and diversification and stress how poor sampling might preclude adequate understanding of endocnidozoan diversity. 2 1 Introduction Cnidarians are generally regarded as a phylum of predatory free‐living animals that occur as benthic polyps and pelagic medusa in the world’s oceans. They include some of the most iconic residents of marine environments, such as corals, sea anemones and jellyfish. Cnidarians are characterised by relatively simple body‐plans, formed entirely from two tissue layers (the ectoderm and endoderm), and by their stinging cells or nematocytes.