Surviving the big chill: overwintering strategies of aquatic and terrestrial insects.
The purpose of this paper is to describe the cold-hardiness of aquatic insects and to use the literature to compare physiological and behavioral strategies that aquatic and terrestrial insects use to cope with minimum winter temperatures. In sharp contrast to terrestrial insects, aquatic insects from seven different orders had limited ability to supercool and did so to temperatures of only −3 to −7°C. Inability to supercool may be due to inoculative freezing—the penetration of external ice crystals through pores or orifices of the insect's cuticle. Furthermore, our results suggest that terrestrial adult stages of aquatic insects may have greater capacity to supercool than aquatic stages of the same taxon. Our results and others' suggested that few aquatic species are freeze tolerant, and those that are appear to be restricted to the order Diptera. Consequently, behavioral avoidance of ice or the capacity to remain unfrozen while encased in ice may be particularly important for overwintering aquatic insects. Ecological implications of insect coldhardiness at the individual, population, and community level are discussed for both terrestrial and aquatic insects.