Reciprocal Symbiont Sharing in the Lodging Mutualism between Walking Corals and Sipunculans

Solitary scleractinian corals of the genera Heterocyathus and Heteropsammia inhabit soft marine bottoms without attaching to hard substrata. The corallums of these genera contain a coiled cavity inhabited by a sipunculan that roams the seafloor, carrying the host coral with it. The coral serves as a sturdy shelter that protects the sipunculan from possible predators. At the same time, the sipunculan maintains the coral in an upright position on the soft bottom. This coral-sipunculan association is unique because two phylogenetically distant coral genera have developed convergent associations with sipunculans. We investigate the process of convergent evolution of two coral species, Hc. aequicostatus and Hp. cochlea, in Okinawa, Japan, with their symbiotic sipunculans, using phylogenetic and morphological analyses. Phylogenetic analyses clarified that the symbiotic sipunculans comprise two distinct clades, surprisingly both of which are associated with both coral species. The bodily habitus of the sipunculan differed between coral species and fit the morphologies of the coiled cavities of their respective host corals. Our results suggest that the two coral species share two sipunculan clades and that sipunculan morphology is plastic and determined by the internal structure of their host corals.