Insect cold tolerance: How many kinds of frozen?

Insect cold tolerance mechanisms are often divided into freezing tolerance and freeze intolerance. This division has been criticised in recent years; Bale (1996) established five categories of cold tolerance. In Bale's view, freezing tolerance is at the extreme end of the spectrum of cold tolerance, and represents insects which are most able to survive low temperatures. Data in the literature from 53 species of freezing tolerant insects suggest that the freezing tolerance strategies of these species are divisible into four groups according to supercooling point (SCP) and lower lethal temperature (LLT): (1) Partially Freezing Tolerant-species that survive a small proportion of their body water converted into ice, (2) Moderately Freezing Tolerant-species die less than ten degrees below their SCP, (3) Strongly Freezing Tolerant-insects with LLTs 20 degrees or more below their SCP, and (4) Freezing Tolerant Species with Low Supercooling Points which freeze at very low temperatures, and can survive a few degrees below their SCP. The last 3 groups can survive the conversion of body water into ice to an equilibrium at sub-lethal environmental temperatures. Statistical analyses of these groups are presented in this paper. However, the data set is small and biased, and there are many other aspects of freezing tolerance, for example proportion of body water frozen, and site of ice nucleation, so these categories may have to be revised in the future. It is concluded that freezing tolerance is not part of Bale's (1996) continuum, but rather a parallel, alternative strategy of cold tolerance.