Reverberation of rapid and slow trills: implications for signal adaptations to long-range communication.

Many acoustic signals in animals include trills, i.e., rapid repetitions of similar elements. Elements within these trills usually are frequency modulated and are degraded by reverberation during long-range transmission. Reverberation primarily affects consecutive elements with the same frequency characteristics and thus imposes a major constraint in the evolution of design and perception of long-range signals containing trills. Here transmission of frequency-unmodulated trills with different element repetition rates was studied. Trills were generated at different frequencies to assess frequency dependence of reverberation and then broadcast under three acoustic conditions--an open field and to assess seasonal changes in transmission properties, a deciduous forest before and after foliage had emerged. Reverberation was quantified at different positions within trills. The results show strong effects of vegetation density (season), transmission distance, frequency, element repetition rate, and element position within the trill on effects of reverberation. The experiments indicate that fast trills transmit less well than slow trills and thus are less effective in long-range communication. They show in particular that selection on trills should not act only on element repetition rate within trills but also on the trill duration as effects of reverberation increased with trill duration.