Many animals have evolved adept sensory systems that enable dexterous mobility in complex environments. Echolocating bats hunting in dense vegetation represent an extreme case of this, where all necessary information about the environment must pass through a parsimonious channel of pulsed, 1D echo signals. We have investigated whether certain bats (rhinolophids and hipposiderids) actively create Doppler shifts with their pinnae to encode additional sensory information. Our results show that the bats' active pinna motions are a source of Doppler shifts that have all attributes required for a functional relevance: (i) the Doppler shifts produced were several times larger than the reported perception threshold; (ii) the motions of the fastest moving pinna portions were oriented to maximize the Doppler shifts for echoes returning from the emission direction, indicating a possible evolutionary optimization; (iii) pinna motions coincided with echo reception; (iv) Doppler-shifted signals from the fast-moving pinna portion entered the ear canal of a biomimetic pinna model; and (v) the time-frequency Doppler shift signatures were found to encode target direction in an orderly fashion. These results indicate that instead of avoiding or suppressing all self-produced Doppler shifts, rhinolophid and hipposiderid bats actively create Doppler shifts with their own pinnae. These bats could hence make use of a previously unknown nonlinear mechanism for the encoding of sensory information, based on Doppler signatures. Such a mechanism could be a source for the discovery of sensing principles not only in sensory physiology but also in the engineering of sensory systems.biosonar | ear motions | Doppler shifts | time-frequency signatures | nonlinear sensing C onspicuous pinna motions (1-5) are an integral part of biosonar behaviors in horseshoe bats and Old World leafnosed bats [families Rhinolophidae and Hipposideridae (6, 7)]. These motions have been demonstrated to enhance sensing and navigation performance (8-10), but the functional role of these dynamic features and the underlying mechanisms have yet to be fully understood (7). In the acoustic domain, source or receiver motions can result in Doppler shifts, i.e., nonlinear scaling of signals in time and frequency. The possibility of pinna-induced Doppler shifts had been mentioned as an aside in early work by Pye and coworkers (2, 3, 11) but has not been considered further-let alone investigated in any depthin the literature since. This complete neglect is regrettable, because ear-generated Doppler shifts could constitute a previously unknown, nonlinear mechanism for the active encoding of sensory information. To test this hypothesis, we have carried out a quantitative experimental investigation of the hypothesis that pinna motions cause functionally relevant Doppler shifts in bats. For pinna-generated Doppler shifts to have a functional role, five necessary conditions must be met: (i) pinna surface speeds must be high enough to produce Doppler shifts that exceed the animal's per...