Phoenix from the Ashes: Fire, Torpor, and the Evolution of Mammalian Endothermy

The evolution of endothermy in mammals and birds has been widely debated. Endothermy is characterized by high endogenous heat production via combustion of metabolic fuels. This differs from ectothermy in most living organisms, which generally do not produce substantial amounts of internal heat for thermoregulation (Tattersall et al., 2012; Withers et al., 2016). Endogenous heat production is energetically very costly. In comparison to ectothermic terrestrial vertebrates, namely the amphibians and reptiles, the minimum metabolic rate (MR) of normothermic or homeothermic (high constant body temperature, Tb) animals at rest is about 4–8-fold higher in the endotherms. This difference is even more pronounced at low ambient temperatures (Ta) at which the Tb of ectotherms follows Ta, and the MR decreases to even lower levels. In contrast, the Tb of homeothermic endotherms remains high and constant over a wide range of Ta. Therefore, to compensate for increased heat loss at low Ta, MR of especially small mammals and birds must increase substantially and can be 100-fold or more of that in ectotherms (Bartholomew, 1982). Of course this high MR requires a substantial uptake of food and in endotherms much of this chemical energy is simply converted into heat for thermoregulation rather than growth or reproduction as in ectotherms

The role of basking in the development of endothermy and torpor in a marsupial

Marsupials have a slow rate of development and this allows a detailed examination of thermoregulatory developmental changes and stages. We quantified the cool-ing rates of marsupial dunnarts (Sminthopsis crassicaudata) at 40–56 days (d) old, and torpor and basking behaviour in animals given the option to bask in four age groups from 60 to 150 d. The development of thermoregulation was a continuum, but was characterised by three major thermoregulatory stages: (1) at 40 d, animals were unable to maintain a constant high body temperature during short-term cold exposure; (2) at 60 d, animals could maintain a high Tb for the first part of the night at an ambient temperature of 15.0 ± 0.7 °C; later in the night, they entered an apparent torpor bout but could only rewarm passively when basking under a heat lamp; (3) from ~90 d, they expressed prolonged torpor bouts and were able to rewarm endogenously. Young newly weaned 60 d animals were able to avoid hypothermia by basking. In this case, basking was not an optional behavioural method of reducing the cost of rewarming from torpor, but was essential for thermoregu-lation independent of the nest temperature. Results from our study suggest that basking is a crucial behavioural trait that permits young marsupials and perhaps other juvenile altricial mammals to overcome the developmental stage between poikilothermy early in development and full endo-thermy later in life

Do small precocial birds enter torpor to conserve energy during development?

Precocial birds hatch feathered and mobile, but when they become fully endothermic soon after hatching, their heat loss is high and they may become energy depleted. These chicks could benefit from using energy-conserving torpor, which is characterised by controlled reductions of metabolism and body temperature (Tb). We investigated at what age the precocial king quail Coturnix chinensis can defend a high Tb under a mild thermal challenge and whether they can express torpor soon after achieving endothermy to overcome energetic and thermal challenges. Measurements of surface temperature (Ts) using an infrared thermometer showed that king quail chicks are partially endothermic at 2-10 days, but can defend high Tb at a body mass of ~13 g. Two chicks expressed shallow nocturnal torpor at 14 and 17 days for 4-5 h with a reduction of metabolism by >40% and another approached the torpor threshold. Although chicks were able to rewarm endogenously from the first torpor bout, metabolism and T40% and another approached the torpor threshold. Although chicks were able to rewarm endogenously from the first torpor bout, metabolism and Ts decreased again by the end of the night, but they rewarmed passively when removed from the chamber. The total metabolic rate increased with body mass. All chicks measured showed a greater reduction of nocturnal metabolism than previously reported in quails. Our data show that shallow torpor can be expressed during the early postnatal phase of quails, when thermoregulatory efficiency is still developing, but heat loss is high. We suggest that torpor may be a common strategy for overcoming challenging conditions during development in small precocial and not only altricial birds