A permanent snow cover for several months is typical for large parts of Norway, Sweden and Finland. Snow layers thicker than about 20 cm insulate the soil surface and stabilize the ground temperature close to 0°C. Many ground-living invertebrates are active at this temperature in the subnivean air space. From this "base camp", some invertebrates migrate upwards to use the snow as a substrate. The intranivean fauna consists of springtails (Collembola) and mites (Acari) that are small enough to move within the narrow pores between snow crystals. The supranivean fauna consists of various invertebrates that are active on the snow surface. Some of them are Collembola that have migrated through the snow layers. However, most of them are larger insects and spiders which migrate between the subnivean and supranivean habitats following air channels which are naturally created along tree stems, bushes etc. penetrating the snow. Likewise, certain Chironomidae and Plecoptera, hatching from winter-open rivers and brooks, are active on the snow surface. The supranivean arthropod fauna has the following characteristics: 1. It is a weather dependent assemblage of species, coming and going with changes in air temperature, cloud cover, and wind. Below ca. -6°C animals are absent, but at temperatures around or above zero, many groups can be simultaneously active on snow. 2. The snow surface fauna shows phenological changes throughout the winter, as certain species and groups are mainly active during certain months. 3. Some invertebrates are highly specialized and take advantage of the snow surface as an arena in their life cycle. Examples are Hypogastrura socialis (Collembola), and the two wingless insects Chionea sp. (Diptera: Limoniidae) and Boreus sp. (Mecoptera). They use the smooth snow surface for efficient migration. Chionea sp. and Boreus sp. lay their eggs during the snow-covered period, while H. socialis migrates to create new colonies. The cold tolerant spider Bolephthyphantes index is unique in constructing webs in small depressions on the snow, to catch migrating Collembola. Various adaptations for using the snow as a substrate are discussed. Besides physiological and morphological adaptations, snow surface arthropods show special behavioural adaptations. Most conspicuous is the ability of several Collembola species to navigate during migration, using the position of the sun for orientation. Furthermore, in Collembola and Mecoptera, jumping as an original mechanism to escape predators has independently evolved into a migrating mechanism. An evolutionary potential exists for more invertebrate groups to take advantage of snow as a substrate in their life cycle. For instance, several more cold tolerant spiders might evolve the ability to catch migrating Collembola on snow.
Ceratophysella sigillata (Collembola, Hypogastruridae) has a life cycle which may extend for >2 years in a temperate climate. It exists in two main morphs, a winter-active morph and a summer-dormant morph in central European forests. The winter-active morph often occurs in large aggregations, wandering on leaf litter and snow surfaces and climbing on tree trunks. The summer-dormant morph is found in the upper soil layers of the forest floor. The cryobiology of the two morphs, sampled from a population near Bern in Switzerland, was examined using Differential Scanning Calorimetry to elucidate the roles of body water and the cold tolerance of individual springtails. Mean (SD) live weights were 62 ± 16 and 17 ± 6 µg for winter and summer individuals, respectively. Winter-active springtails, which were two feeding instars older than summer-dormant individuals, were significantly heavier (by up to 4 times), but contained less water (48% of fresh weight [or 0.9 g g-1 dry weight]) compared with summer-dormant animals (70% of fresh weight [or 2.5 g g-1 dry weight]). Summer-dormant animals had a slightly greater supercooling capacity (mean (SD) -16 ± 6°C) compared with winter-active individuals (-12 ± 3°C), and they also contained significantly larger amounts of both total body water and osmotically inactive (unfrozen) water. In the summer morph, the unfrozen fraction was 26%, compared to 11% in the winter morph. The ratio of osmotically inactive to osmotically active (freezable) water was 1 : 1.7 (summer) and 1 : 3.3 (winter); thus unfrozen water constituted 59% of the total body water during summer compared with only 30% in winter. Small, but significant, levels of thermal hysteresis were detected in the winter-active morph (0.15°C) and in summer-dormant forms (0.05°C), which would not confer protection from freezing. However, the presence of antifreeze proteins may prevent ice crystal growth when feeding on algae with associated ice crystals during winter. It is hypothesised that in summer animals a small decrease in freezable water results in a large increase in haemolymph osmolality, thereby reducing the vapour pressure gradient between the springtail and the surrounding air. A similar decrease in freezable water in winter animals will not have such a large effect. The transfer of free water into the osmotically inactive state is a possible mechanism for increasing drought survival in the summer-dormant morph. The ecophysiological differences between the summer and winter forms of C. sigillata are discussed in relation to its population ecology and survival.