A review of Fennoscandian arthropods living on and in snow

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.