Elements of cold hardiness in a littoral population of the land snail Helix aspersa (Gastropoda: Pulmonata)

The land snail Helix aspersa can be considered partially tolerant to freezing, in the sense it can survive some ice formation within its body for a limited time, and possesses a limited ability to supercool. This study aimed at understanding what factors are responsible for the variation of the temperature of crystallization (Tc) in a littoral temperate population. The ability to supercool was maximal (ca. -5 degreesC) during dormancy periods (hibernation and aestivation) and minimal (ca. -3 degreesC) during spring and autumn, in relation with the decrease of water mass and the increase of osmolality. Tc decreased in October to remain stable through late autumn and winter; it increased quickly with the awakening of animals in April. Snails with an epiphragm had a significantly higher ability to supercool (ca. -4.8 degreesC) than snails which did not form an epiphagm (ca. -4.2 degreesC). The animals' size had a weak but significant influence on the realization of the Tc. It appeared that there was not a real cold-hardiness strategy in this population; rather a sum of parameters, varying in consequences of the external conditions and of the activity cycle, which are responsible for the enhancement of the supercooling ability during winter.