Cold-hardiness and related adaptations of insects in the Arctic corres
pond to characteristic climatic constraints. Some species are long-liv
ed and are cold-hardy in several stages. In the Arctic, diapause and c
old-hardiness are less likely to be linked than in temperature regions
, because life-cycle timing depends as much on the need to coincide de
velopment with the short summer as on the need to resist winter cold.
Winter habitats of many species are exposed rather than sheltered from
cold so that development in spring can start earlier. Several feature
s of cold-hardiness in arctic species differ from the characteristics
of temperate species: these include very cold-hardy insects with low s
upercooling points that are not freezing tolerant; freezing-tolerant s
pecies that supercool considerably rather than freezing at relatively
high subfreezing temperatures; mitochondrial degradation linked with t
he accumulation of cryoprotectants; and the possibly limited occurrenc
e of thermal hysteresis proteins in winter. Several interesting relati
onships between cold-hardiness and water have been observed, including
different types of dehydration. Winter mortality in arctic insects ap
pears to be relatively low. Adaptations to cold in summer include rete
ntion of cold-hardiness, even freezing tolerance; selection of warm si
tes; and behaviour such as basking that allows elevated body temperatu
res. Studies especially on the high-arctic moth Gynaephora groenlandic
a show that various factors including cold-hardiness and other summer
and winter constraints dictate the structure of energy budgets and the
timing of life cycles. Future work should focus on the biological and
climatic differences between arctic and other areas by addressing hab
itat conditions, life-cycle dynamics, and various aspects of cryoprote
ctant production at different times of year. Even in the Arctic cold-h
ardiness is complex and involves many simultaneous adaptations.