Cardiovascular contributions and energetic costs of thermoregulation in ectothermic vertebrates

Ectothermic vertebrates use a suite of physiological and behavioral mechanisms to thermoregulate, which result in various thermoregulatory strategies from thermoconformity to thermoregulation. Here, we present a novel synthesis of theoretical and empirical methods to determine cardiovascular contributions to heat transfer in free-living ectothermic vertebrates. We start by identifying the fundamental components of heat transfer and the cardiovascular mechanisms for physiological modulation of heat exchange, and then integrate these components into a single, integrative framework: the cardiovascular heat exchange framework (CHEF). We demonstrate that this framework can identify details of the thermoregulatory strategy in two turtle species, most notably the preponderance of instances where turtles use physiological mechanisms to avoid overheating, suggesting vulnerability to climate change. As modulated physiological contributions to heat flow incur a greater energy demand than relying on unmodulated passive heat transfer, we then asked whether we could characterize the energetic costs of thermoregulation. We measured field metabolic rate (FMR) in free-living turtles and used the CHEF to determine FMR while actively or passively thermoregulating. Comparing an individual's actual FMR to the rate calculated assuming absence of thermoregulation revealed that painted turtles, a partial thermoregulator, elevate their daily energy expenditure (DEE) by about 25%, while box turtles, a thermoconformer, have a DEE that is nearly unchanged as a result of thermoregulation. This integrative framework builds a new paradigm that provides a mechanism to explain correlations between energy demand and thermoregulatory strategy, quantifies the energetic costs of thermoregulation, and identifies the role of cardiovascular contributions to thermoregulation in free-living animals.