A conceptual framework to integrate cold-survival strategies: torpor, resistance and seasonal migration

Freezing temperatures are inherently challenging for life, which is water based. How species cope with these conditions fundamentally shapes ecological and evolutionary processes. Despite this, there is no comprehensive conceptual framework for cold-survival strategies-seasonal migration, cold resistance and torpor. Here, I propose a framework with four components for conceptualizing and quantifying cold-survival strategies. Cold-survival strategies are (i) collectively encompassed by the proposed framework, and that this full breadth of strategies should be considered in focal species or systems (comprehensive consideration). These strategies also (ii) exist on a spectrum, such that species can exhibit partial use of strategies, (iii) are non-exclusive, such that some species use multiple strategies concurrently (combined use) and (iv) should collectively vary inversely and proportionally with one another when controlling for the external environment (e.g. when considering species that occur in sympatry in their summer range), such that use of one strategy reduces, collectively, the use of others (proportional use). This framework is relevant to understanding fundamental patterns and processes in evolution, ecology, physiology and conservation biology.