Development of Echolocation Calls in the Mustached Bat,Pteronotus parnellii

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Abstract
Adult mustached bats employ Doppler-sensitive sonar to hunt fluttering prey insects in acoustically cluttered habitats. The echolocation call consists of 4–5 harmonics, each composed of a long constant frequency (CF) component flanked by brief frequency modulations (FM). The 2nd harmonic CF component (CF2) at 61 kHz is the most intense, and analyzed by an exceptionally sharply tuned auditory system. The maturation of echolocation calls and the development of Doppler-shift compensation was studied in Cuba where large maternity colonies are found in hot caves. In the 1st postnatal week, infant bats did not echolocate spontaneously but could be induced to vocalize CF-FM signals by passive body motion. The CF2frequency emitted by the smallest specimens was at 48 kHz (i.e., 0.4 octaves lower than the adult signal). CF-FM signals were spontaneously produced in the 2nd postnatal week at a CF2frequency of 52 kHz. The CF2frequencies of induced and spontaneous calls shifted upward to reach a value of 60.5 kHz in the 5th postnatal week. Standard deviations of CF2frequency were large (up to ±1.5 kHz) in the youngest bats and dropped to values of ±250 Hz at the end of the 3rd postnatal week. Some individuals in the 4th and 5th postnatal weeks emitted with adultlike frequency precision of about ±100 Hz. In the youngest bats, the 1st harmonic CF component (CF1) was up to 22 dB stronger than CF2. Adultlike relative levels of CF1(–28 dB relative to CF2) were reached in the 5th postnatal week. In spontaneously emitted CF-FM calls, the duration of the CF2component gradually increased with age from 5 ms to maximum values of 18 ms. Durations of the CF2component in induced calls averaged 7 ± 2.6 ms in the 1st postnatal week and 8.2 ± 1.5 ms in the 5th postnatal week. There were no age-related changes in duration of the terminal FM sweep (3 ± 0.4 ms) in both induced and spontaneous calls. The magnitude of the terminal FM sweep in spontaneous calls was not correlated with age (mean 13.5 ± 2 kHz). Values for induced calls slightly increased with age from 11 ± 2 to 13 ± 2 kHz. The emission rate of induced CF-FM signals increased with age from values of 2.5 ± 2 to 17 ± 5 pulses/s. Values for spontaneously emitted calls were 4.4 ± 3 and 9 ± 4.5 pulses/s, respectively. Doppler-shift compensation, as tested in the pendulum task, emerged during the 4th postnatal week in young bats that were capable of very brief active flights, but before the time of active foraging outside the cave.