Flexible embryonic shell allies large offspring size and anti-predatory protection in viviparous snails

Abstract

The evolutionary conflicts between viviparous reproductive mode and skeleton shape may occur whenever the space available for embryo development or delivery is limited by hard inflexible structures of a parent (bones, shell, etc.). In tetrapods, offspring size is at odds with female locomotion efficiency, which results in obstetric selection. We suggest a similar relationship for viviparous gastropods, where spacious canal needed for embryo delivery may interfere with anti-predatory role of narrow and toothed shell aperture. We explored this hypothesis in the group of viviparous land snails (Clausiliidae, subfamily Phaedusinae), known for complex apertural barriers protecting the shell interior. Most of the shell structure modifications we recorded facilitate the delivery of embryos but simultaneously reduce the safeguard of a narrow shell opening. However, we also observed highly flexible embryonic shells that may withstand squeezing between apertural barriers during birth. We investigated the microstructure of these flexible embryonic shells, compared to the typical hard shells of clausiliid embryos, which are rigid and unpliable already in the genital tract of the parent. Our results suggest that the unusual flexibility, which is related to a low number of organomineral layers in the shell, evolved in two phylogenetically distant lineages of Phaedusinae. This adaptation reduces mechanical constraints for birth of the neonates but allows to maintain the protective function of the apertural barriers.