Snail shell shape, force of attachment, and metabolic rate together cope with the intertidal challenge

Inhabitants of rocky intertidal shores, including gastropods, require specific adaptations to cope with numerous challenges that vary across the intertidal levels. We collected Stramonita biserialis snails from upper (wave-protected and intense predation) and lower (wave-exposed and low predation) intertidal sites to compare the following traits: shell skeleton (ventral and abaxial lateral views of shell shape, thickness, and mass), foot size, energy metabolism, and attachment strength to determine whether the trait values of snails from each zone fit the environmental challenges they face. We used a Principal Component Analysis to reduce the dimensionality of the data. Multivariate Analysis of Covariance (MANCOVA) for comparing characteristics between the two intertidal zones, and Partial Least Squares (PLS) analyses for testing the integration of overall snail characteristics. The traits of the snails of the two intertidal sites matched with the adaptations expected to allow them to cope with their contrasting challenges. The snails from the lower intertidal had more streamlined shells (which reduces drag forces) and a larger aperture and foot extension (which increase the strength of their attachment to the substrate) compared to snails from the upper sites. Snails from the lower intertidal also had a high mass-specific metabolism and soft body proportion, indicating that these snails from the wave-exposed sites have an energetically active musculature that matches their strong substrate attachment. The thin shell walls of the snails of the lower intertidal match the relatively low predatory pressure there.