Thermoregulation in monotremes: riddles in a mosaic

The three extant genera of the Monotremata have evolved, probably from a pre-Cretaceous Gondwanan origin, independently of the Theria to display a variety of ancestral and derived features. A comparison of their thermoregulation reveals a diversity of physiology that might represent both plesiomorphic and apomorphic elements within this mosaic. In the tachyglossids, the echidnas Tachyglossus and Zaglossus, body temperature is often labile, rising as a result of activity and allowed to decline during inactivity. This daily heterothermy, which is not necessarily torpor, may combine with typical mammalian hibernation to provide substantial energy economy in a wide variety of often unproductive habitats. Only when incubating do free-ranging echidnas display classic mammalian thermoregulation, the facultative nature of which suggests echidna-like physiology as an example of a protoendothermic stage in the evolution of endothermy. Similarly, physiological response to heat in Tachyglossus, at least, may be plesiomorphic, relying on the cyclic loss of heat stored during activity. Tachyglossids neither exhibit a panting response nor spread saliva to facilitate evaporative cooling and Tachyglossus, though not Zaglossus, lacks functional sweat glands. By contrast, the only extant ornithorhynchid, the platypus Ornithorhynchus, does not utilise heterothermy of any kind and maintains its body temperature more tightly than several semiaquatic eutherians. Although not necessarily required, it responds to heat via sweating, but not panting or saliva spreading. The classic nature of ornithorhynchid thermoregulation stands in marked contrast to the more diverse thermoregulatory responses shown by the tachyglossids, making it difficult to determine which aspects of monotreme thermoregulation are plesiomorphic and which are apomorphic.