Singing out of tune: sexual and developmental differences in the occurrence of nonlinear phenomena in primate songs

Animal vocalizations contain a varying degree of nonlinear phenomena (NLP) caused by irregular or chaotic vocal organ dynamics. Several hypotheses have been proposed to explain NLP presence, from unintentional by-products of poor vocal technique to having a functional communicative role. We aimed to disentangle the role of sex, age and physiological constraints in the occurrence of NLP in the songs of the lemur Indri indri, which are complex harmonic vocal displays organized in phrases. Age and sex affected the presence and type of NLP in songs. In particular, the proportion of the phenomena considered decreased with age, except for subharmonics. Subharmonics potentially mediate the perception of lower pitch, making signallers appear larger. Subharmonics and frequency jumps occurred in lower-pitched notes than regular units, while chaos and sidebands occurred in higher-pitched units. This suggests that different types of NLP can be associated with different vocal constraints. Finally, indris might present short-term vocal fatigue, with units occurring in the last position of a phrase having the highest probability of containing NLP. The presence of NLP in indris might result from proximate causes, such as physiological constraints, and ultimate causes, such as evolutionary pressures, which shaped the communicative role of NLP.