Syntax denotes a rule system that allows one to predict the sequencing of communication signals. Despite its significance for both human speech processing and animal acoustic communication, the representation of syntactic structure in the mammalian brain has not been studied electrophysiologically at the single-unit level. In the search for a neuronal correlate for syntax, we used playback of natural and temporally destructured complex species-specific communication calls-socalled composites-while recording extracellularly from neurons in a physiologically well defined area (the FM-FM area) of the mustached bat's auditory cortex. Even though this area is known to be involved in the processing of target distance information for echolocation, we found that units in the FM-FM area were highly responsive to composites. The finding that neuronal responses were strongly affected by manipulation in the time domain of the natural composite structure lends support to the hypothesis that syntax processing in mammals occurs at least at the level of the nonprimary auditory cortex.From a linguist's point of view (1-3), ''syntax'' denotes a rule system that accounts for the ability to produce an infinite variety of sequences (i.e., words and sentences) from a fixed number of phonemes (i.e., vowels and consonants). Apart from human speech, rule systems for the sequencing of speciesspecific vocalizations have been found repeatedly in both birds (4, 5) and nonhuman mammals (6, 7). Thus, more generally, ''syntax'' can be understood as any system of rules that allows one to predict the sequencing of communication signals (3).Our present report of syntax processing by auditory cortical neurons in the mustached bat Pteronotus parnellii is founded on several previous studies in this species by employing both neurophysiological and behavioral methods. First, in the context of intraspecific acoustic communication, mustached bats frequently combine otherwise independently emitted simple syllables to form either isosyllabic trains or heterosyllabic composites. The syntactical rules for the generation of the last-mentioned higher order constructs have been revealed in detail (8). Second, the auditory cortex of P. parnellii is arguably the most intensively studied and best understood of all mammals (9). Thus, established representational maps (e.g., ref. 10) can be used as a reference to record from defined areas and functional subtypes of auditory cortical neurons. Third, neurons in the FM-FM area of the mustached bat auditory cortex were shown recently to respond facilitatively to isosyllabic pairs (11). Presumably, these neurons mediate acoustic communication (11) in addition to their primary (i.e., first discovered) function in echolocation (12).FM-FM neurons exhibit heteroharmonic combinationsensitivity for paired stimuli mimicking the FM components of the bats' echolocation sounds (pulse and echo) (12) and often respond well to communication calls (11). Thus, we hypothesized that FM-FM neurons may also show combination-sensit...