A crucial, common feature of speech and music is that they show non-random structures over time. It is an open question which of the other species share rhythmic abilities with humans, but in most cases the lack of knowledge about their behavioural displays prevents further studies. Indris are the only lemurs who sing. They produce loud howling cries that can be heard at several kilometres, in which all members of a group usually sing. We tested whether overlapping and turn-taking during the songs followed a precise pattern by analysing the temporal structure of the individuals' contribution to the song. We found that both dominants (males and females) and non-dominants influenced the onset timing one another. We have found that the dominant male and the dominant female in a group overlapped each other more frequently than they did with the non-dominants. We then focused on the temporal and frequency structure of particular phrases occurring during the song. Our results show that males and females have dimorphic inter-onset intervals during the phrases. Moreover, median frequencies of the unit emitted in the phrases also differ between the sexes, with males showing higher frequencies when compared to females. We have not found an effect of age on the temporal and spectral structure of the phrases. These results indicate that singing in indris has a high behavioural flexibility and varies according to social and individual factors. The flexible spectral structure of the phrases given during the song may underlie perceptual abilities that are currently unknown in other nonhuman primates, such as the ability to recognize particular pitch patterns.

Contextual variation in the loud calls of strepsirhine primates is poorly understood. To understand whether songs given by indris in different contexts represent acoustically distinct variants and have the potential to elicit context-specific behaviours in conspecific listeners, we investigated the acoustic variability of these songs and the distance travelled by vocalizers after their emissions. Songs of 41 individuals were recorded from 16 indri groups in four different forest sites in eastern Madagascar. We collected a total of 270 duets and choruses arising during territorial defence, advertisement and cohesion. We demonstrated that the structure of indri songs conveyed context-specific information through their overall duration, but shared the sequential pattern of harsh units (roars) followed by long notes and, finally, descending phrases. Analysing in detail individual contributions to advertisement songs and cohesion songs, we found that the acoustic structure of units could be classified correctly with a high degree of reliability (96.23% of long notes, 80.16% of the descending phrases, 72.54% of roars). Future investigations using playback stimuli could explore the relationship between acoustic features and the information transmitted by the song.

The complexity of primates’ singing behavior has long gathered the attention of researchers interested in understanding the selective pressures underpinning the evolution of language. Among these pressures, a link between territoriality, pair-living, and singing displays has been suggested. Historically, singing primates have been found in a few taxa that are not closely related to each other, and, in the last years, their phylogeny has dramatically changed. Hence, we aimed at understanding if the dogmatic association between territorial behavior and a monogamous social structure still holds in the light of current research. Moreover, singing behavior has often been considered a whole, but animals can perform different singing forms depending on how many individuals call simultaneously. Currently, it is unclear to which extent these singing forms are widespread among these primate groups. Given that there is no unique definition for a “song”, “solo”, “duet”, and “chorus”, we envisioned some of the most used descriptions. We then formulated some new definitions that we followed in our review of the presence/absence of these different forms of song organization among singing primates’ taxa. In particular, we suggested that tarsier species that are typically considered non-singers may indeed sing, and we pointed out that non-duetting gibbons may perform duet interactions. We found that, besides duets, chorusing behavior and solo songs are essential features of primates’ communication, but their study is still in a descriptive phase. Moreover, while territorial behavior seems to be conserved in these singing taxa, we highlighted that the monogamous social structure is not the rule. Pair-living plus multi-females groups displaying singing behavior are common too. We suggest that ending to consider these taxa as uniform in their sociality and vocal behavior might be a significant turning point to unravel the different selective pressures that influenced the emergence and organization of such peculiar vocal behavior.

The increasing interest in the evolution of human language has led several fields of research to focus on primate vocal communication. The ‘singing primates’, which produce elaborated and complex sequences of vocalizations, are of particular interest for this topic. Indris (Indri indri) are the only singing lemurs and emit songs whose most distinctive portions are “descending phrases” consisting of 2-5 units. We examined how the structure of the indris’ phrases varied with genetic relatedness among individuals. We tested whether the acoustic structure could provide conspecifics with information about individual identity and group membership. When analyzing phrase dissimilarity and genetic distance of both sexes, we found significant results for males but not for females. We found that similarity of male song-phrases correlates with kin in both time and frequency parameters, while, for females, this information is encoded only in the frequency of a single type. Song phrases have consistent individual-specific features, but we did not find any potential for advertising group membership. We emphasize the fact that genetic and social factors may play a role in the acoustic plasticity of female indris. Altogether, these findings open a new perspective for future research on the possibility of vocal production learning in these primates.

Territorial, socially monogamous species actively defend their home range against conspecifics to maintain exclusive access to resources such as food or mates. Primates use scent marks and loud calls to signal territory occupancy and limit the risk of intergroup encounters, maximizing their energetic balance. Indri indri is a little-studied territorial, socially monogamous singing primate living in family groups. The groups announce territory occupancy with long-distance calls, and actively defend their territories from conspecific intruders. This work includes data collected in three forests in Madagascar on 16 indri groups over up to 5 years. We aimed (1) to estimate the extent of territories using minimum convex polygon (MCP), implementing minimum sampling effort requirements; (2) to quantify territorial exclusivity, measuring the overlap between territories; and (3) to evaluate the intergroup encounter rate and to quantify the dynamics of group encounters. Our results showed that indris range evenly within exclusive small territories with no or little overlap. Intergroup encounters are rare (0.05 encounters per day), and are located on the periphery of the territories. Disputes were mostly solved with vocal confrontation and only in 13% of the cases ended in physical fights. This frame underlines a cost–benefit explanation of territoriality, favouring a strategy that efficiently limits overlap and avoids costly intergroup encounters. We hypothesize that territorial behaviour in indri is related to mate-guarding strategy and that vocal behaviour plays a fundamental role in regulating intergroup spacing dynamics.