How aquatic insects live in cold climates

In cold climates most aquatic habitats are frozen for many months. Nevertheless, even in such regions the conditions in different types of habitat, in different parts of one habitat, and from one year to the next can vary considerably; some water bodies even allow winter growth. Winter cold and ice provide challenges for aquatic insects, but so do high spring flows, short, cool summers, and unpredictable conditions. General adaptations to cope with these constraints, depending on species and habitat, include the use of widely available foods, increased food range, prolonged development (including development lasting more than one year per generation), programmed life cycles with diapause and other responses to environmental cues (often enforcing strict univoltinism), and staggered development. Winter conditions may be anticipated not only by diapause and related responses but also by movement for the winter to terrestrial habitats, to less severe aquatic habitats, or to different parts of the same habitat, and by construction of shelters. Winter itself is met by various types of cold hardiness, including tolerance of freezing in at least some species, especially chironomid midges, and supercooling even when surrounded by ice in others. Special cocoons provide protection in some species. A few species move during winter or resist anoxia beneath ice. Spring challenges of high flows and ice scour may be withstood or avoided by wintering in less severe habitats, penetrating the substrate, or delaying activity until after peak flow. However, where possible species emerge early in the spring to compensate for the shortness of the summer season, a trait enhanced (at least in some lentic habitats) by choosing overwintering sites that warm up first in spring. Relatively low summer temperatures are offset by development at low temperatures, by selection of warm habitats and microhabitats, and in adults by thermoregulation and modified mating activity. Notwithstanding the many abiotic constraints in cold climates, aquatic communities are relatively diverse, though dominated by taxa that combine traits such as cold adaptation with use of the habitats and foods that are most widely available and most favourable. Consequently, except in the most severe habitats, food chains and community structure are complex even at high latitudes and elevations, including many links between aquatic and terrestrial habitats. Despite the complex involvement of aquatic insects in these cold-climate ecosystems, we know relatively little about the physiological and biochemical basis of their cold hardiness and its relationship to habitat conditions, especially compared with information about terrestrial species from the same regions.