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Flattening of coiled shells has occurred in several gastropod lineages, while the evolutionary process of shell flattening is little known. The subfamily Fossarininae of the top shell family (Trochidae) is unique, because it includes four genera at various stages of shell flattening.  Broderipia  and  Roya , have zygomorphic shells that has lost coiling, while the sister genera,  Fossarina  and  Synaptocochlea , have respectively turbiniform and auriform shells. Therefore, comparisons of biology, habitats and detailed morphology among these four genera may help us to detect selection pressure driving shell flattening and loss of coiling. Although  Broderipia  has recently been identified as living symbiotically in the pits of sea urchins, the habitats and biology of the other three Fossarininae species, especially  Roya  are poorly known. After an extensive search on rocky shores of the Japanese Archipelago, we found live  Roya eximia  snails on intertidal/subtidal rock surfaces exposed to strong waves.  Roya  snails crept on the bare rock surface to graze periphyton at low tide, and fled into vacant barnacle shells at high tide. Comparison of the morphology of soft bodies in Fossarininae revealed that the columellar muscle of flattened species has been drastically elongated and arranged in posterior semi-outer edge of the flattened shell as observed in true limpets. The flattering and loss of coiling of the shell in  Roya  caused acquisition of a zygomorphic flat body, retraction of coiled visceral mass, and expansion of the foot sole. All of these changes improved tolerance against strong waves and the ability to cling to rock surfaces, and thus enabled a lifestyle utilizing both wave-swept rock surfaces and the inside of vacant barnacle shells.

Morphological and ecological adaptation of limpet-shaped top shells (Gastropoda: Trochidae: Fossarininae) to wave-swept rock reef habitats