Finding the right thermal limit: a framework to reconcile ecological, physiological and methodological aspects of CTmax in ectotherms.

Upper thermal limits (CTmax) are frequently used to parameterize the fundamental niche of ectothermic animals and to infer biogeographical distribution limits under current and future climate scenarios. However, there is considerable debate associated with the methodological, ecological and physiological definitions of CTmax. The recent (re)introduction of the thermal death time (TDT) model has reconciled some of these issues and now offers a solid mathematical foundation to model CTmax by considering both intensity and duration of thermal stress. Nevertheless, the physiological origin and boundaries of this temperature-duration model remain unexplored. Supported by empirical data, we here outline a reconciling framework that integrates the TDT model, which operates at stressful temperatures, with the classic thermal performance curve (TPC) that typically describes biological functions at permissive temperatures. Further, we discuss how the TDT model is founded on a balance between disruptive and regenerative biological processes that ultimately defines a critical boundary temperature (Tc) separating the TDT and TPC models. Collectively, this framework allows inclusion of both repair and accumulation of heat stress, and therefore also offers a consistent conceptual approach to understand the impact of high temperature under fluctuating thermal conditions. Further, this reconciling framework allows improved experimental designs to understand the physiological underpinnings and ecological consequences of ectotherm heat tolerance.