Supercool snails

In the last couple of months, I've regularly come across active Girdled Snails on my way to work. Damp, but often very cold even during frosty mornings I see these small snails on they way back to their day retreats on the pavement by a front garden, presumably after having been active, feeding? during the night (above, on the 23rd of January at 8:48 am). This nonchalant cold hardiness is in stark contrast to common or garden snails (Cornu aspersum), which have been dormant for a good while, and won't become active until March or April. Why is that? how cold resistant are snails? do they differ in their cold hardiness?
  Land snails are a very useful indicator species for ancient environments, as their shells fossilise very well and can be often identified to species level. Despite this, surprisingly little is known on their cold tolerance. Amazingly, 35 snail species live north of the arctic circle and 44 species over 2,000 m of altitude. How do they survive ice-cold temperatures? As other invertebrates unable to migrate snails have two strategies to survive sub-zero temperatures: freezing avoidance and freezing tolerance. Species that engage in freezing avoidance can actually be active in sub-zero conditions by supercooling. A supercooled snail will be at a temperature under 0 oC, but it won't be frozen, that is cool indeed! They can do this by producing large amounts of small sugar molecules that bind water and make their tissues more dehydrated, and also large antifreeze proteins, which inhibit ice formation even further, allowing them to remain active at sub-zero temperatures. Smaller snails (of shells up to 15 mm) appear to be more freeze avoidant than tolerant, and therefore, they are better at supercooling.

A favourite overwintering spot, with dozens of garden snails of various sizes under a tile lined against a wall in my garden.

We know a bit more about the cold tolerance of garden snails, thanks to the research of Armelle Ansart, from Rennes University and her colleagues. The garden snail has limited supercooling abilities, it is a partial frost tolerant species (they can only survive to a minimum of -5 oC). The are partially freeze-tolerant, avoiding freezing by emptying their guts - gut contents can start the formation of deadly ice crystals, reducing the water content of their body (which makes soluble chemicals more concentrated and decreases the temperature at which ice crystals form) and producing an epiphragm, a hard, thick calcareous layer of mucus that seals their shells shut, keeping the deadly moisture out. As an aside, the epiphragm is also produced in very dry weather, during aestivation, another dormant state in snails, but then the epiphragm keeps the moisture in.

An early waking young garden snail (28 Feb 2011), still carrying its epiphragm attached to its shell.

The preparation for overwintering seems to be kickstarted by the decreasing photoperiod of autumn, rather than temperatures dropping. Garden snails also seek high and dry microhabitats to overwinter and congregate, sometimes in very large numbers in favourable spots, such as the underside of logs, stones or holes in tree trunks. Large garden snails are more resistant to the cold than small ones, as they are better at avoiding the formation of ice crystals, so adults are more likely to survive a hard winter than immature snails.
 As for the Girdled snail, sadly I found nothing, although a comparative analysis of cold hardiness by Ansart and colleagues found out that its congeneric species Hygromia limbata freezes at -7 oC, not too impressive when compared to the tiny Columella edentula, also a British species, which doesn't freeze until the temperature descends to -17 oC, but probably enough to allow it to survive in the mild frosts of Hull.

References
Armelle Ansart, Annie Guiller, Olivier Moine, Marie-Claire Martin, Luc Madec. 2014. Is cold hardiness size-constrained? A comparative approach in land snails. Evolutionary Ecology, 28: 471-493. Here.

Ansart, Armelle, and Philippe Vernon (2003) Cold hardiness in molluscs. Acta Oecologica 24.2: 95-102.

Ansart, A. & Vernon, P. (2004). Cold hardiness abilities vary with the size of the land snail Cornu aspersum. Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology, 139: 205-211.