The biology and functional morphology of Modiolarca subpicta (Bivalvia: Mytilidae: Musculinae), epizoically symbiotic with Ascidiella aspersa (Urochordata: Ascidiacea), from the Kattegat, northern Jutland, Denmark

This study of the little known north-eastern Atlantic mytilid Modiolarca subpicta suggests a more intimate relationship with ascidians and, especially, Ascidiella aspersa, than has hitherto been appreciated. Both live for similar to 18 months and the reproductive cycle of both is approximately co-ordinated so that juvenile ascidians become available as hosts to the settling spat of the symbiotically epizoic M. subpicta each summer. Settling spat of M. subpicta are thought to be attracted to the exhalant flow from the ascidian's anal siphon. After the ascidian's death, the liberated mussels may adopt a wandering, free-living, lifestyle they have occasionally been reported to pursue. Anatomically, M. subpicta is of the typical mytilid plan, albeit simplified commensurate upon its normal lifestyle of protective envelopment within the host's tunic. The foot, however, is highly mobile and in addition to being used for locomotion, especially in juveniles, is responsible for the planting of byssal threads, typically dorsally above the shell, so that each individual stimulates the ascidian to produce an epidermal pocket into which it manoeuvres itself and reposes, securely attached, but dorsal side down Modiolarca subpicta, epizoic in the tests of ascidians, could have evolved from a more isomyarian, equivalve, ancestor via a nestling, epibyssate, Trichomusculus-like shaped intermediary with ventral flattening and, hence, stability characteristic of the more familiar heteromyarian mytilids of rocky shores and lotic freshwaters being achieved through living dorsal side down.