The Lombard effect, an involuntary rise in call amplitude in response to masking ambient noise, represents one of the most efficient mechanisms to optimize signal-to-noise ratio. The Lombard effect occurs in birds and mammals, including humans, and is often associated with several other vocal changes, such as call frequency and duration. Most studies, however, have focused on noisedependent changes in call amplitude. It is therefore still largely unknown how the adaptive changes in call amplitude relate to associated vocal changes such as frequency shifts, how the underlying mechanisms are linked, and if auditory feedback from the changing vocal output is needed. Here, we examined the Lombard effect and the associated changes in call frequency in a highly vocal mammal, echolocating horseshoe bats. We analyzed how bandpassfiltered noise (BFN; bandwidth 20 kHz) affected their echolocation behavior when BFN was centered on different frequencies within their hearing range. Call amplitudes increased only when BFN was centered on the dominant frequency component of the bats' calls. In contrast, call frequencies increased for all but one BFN center frequency tested. Both amplitude and frequency rises were extremely fast and occurred in the first call uttered after noise onset, suggesting that no auditory feedback was required. The different effects that varying the BFN center frequency had on amplitude and frequency rises indicate different neural circuits and/or mechanisms underlying these changes.A ny transmission of signals between sender and receiver faces the challenge of being subjected to masking by noise. For acoustic signals, for example, animals have evolved several strategies that aid in increasing the signal-to-noise ratio, thus facilitating signal transmission. One of the most efficient mechanisms is the socalled Lombard effect, i.e., the involuntary rise in call amplitude in response to masking ambient noise (1). This effect was first described in human communication a century ago (2) and has since been found in several species of birds (3-11) and various mammals (12-17), including bats (18). In human speech, several vocal changes, such as a rise in fundamental frequency (19) or lengthening in word duration (20), are often accompanied with the Lombard effect; combined, these changes are referred to as Lombard speech (21). Vocal changes associated with the Lombard effect have rarely been analyzed in animal species. So far, noisedependent changes in call frequency have been observed in birds (8,11,22), and changes in call duration in birds and monkeys (8,12).Most animal studies, however, have focused on noise-dependent changes in call amplitude and do not examine other possible vocal changes (2-5, 7, 9, 10, 15-17). It is therefore still largely unknown how the adaptive changes in call amplitude relate to frequency shifts or call elongation and how the underlying mechanisms are linked. It is also unknown if auditory feedback from the changing vocal output is needed to drive the Lombard effect in general.Over...