Endothermy ("warm-bloodedness") underpins the ecological dominance of mammals and birds in diverse environmental settings^1-3. However, it is unclear when this crucial feature emerged during mammalian evolutionary history, as most fossil evidence is ambiguous^4-25. Here, we show that new information on this key evolutionary transition can be obtained from the morphology of the endolymph-lled semicircular ducts of the inner ear that monitor head rotations and are essential for motor coordination, navigation, and spatial awareness^26-31. Increased body temperature during the ectotherm-endotherm transition of mammal ancestors would decrease endolymph viscosity, negatively impacting the biomechanics of the semicircular ducts^32,33, while simultaneously increasing activity levels^34,35 required improved performance^36. Speci c morphological changes to the membranous ducts and enclosing bony canals were, therefore, necessary to maintain optimal functionality. We track these morphological changes in 341 vertebrates, including 56 extinct synapsids, and show that canals with relatively thin cross-sections and small radii of curvature are indicative of mammalian endothermy. This inner ear morphotype evolved abruptly ~233 million years ago, during the Late Triassic, in Mammaliamorpha. Our conclusion differs from previous suggestions3-17, and we interpret most stem-mammals as ectotherms. Endothermy as a crucial physiological characteristic joins other distinctive mammalian features that arose during this period of climatic instability^37-39. *Ricardo Araújo and Romain David contributed equally to this work.
Main TextEndotherms can maintain high and nearly constant body temperature through metabolic heat production, allowing them to optimize chemical reactions and sustain aerobic activity for long periods of time while remaining relatively independent from external conditions 1,2,40 . Compared to ectotherms, endotherms are more active, travel farther, and achieve higher locomotor speeds, all at the expense of higher energy costs 34,35 (Supplementary Data 1). Consequently, extant mammals and birds occupy a variety of ecological niches unrivalled by other vertebrates. Endothermy is a quintessentially mammalian feature, intimately related to other hallmarks such as lactation, sweat glands and fur 2,3 . However, its evolution remains one of the great unsolved mysteries of palaeontology 2,3 . Lines of evidence invoked to identify the emergence of mammalian endothermy rely mostly on skeletal anatomical features, but also on ichnological, histological and isotopic information, which have been used as correlates for aerobic