Bioacoustic characterization of the black-and-white ruffed lemur (Varecia variegata) vocal repertoire.

Ruffed lemurs (Varecia spp.) exhibit a unique suite of behavioral traits compared to other lemur species, which includes their fluid fission-fusion social dynamics, communal rearing of parked litters, and pronounced frugivory in their humid rainforest habitats. Given these traits, and the dense rainforests they inhabit, vocal communication may be key to maintaining social cohesion, coordinating infant care, and/or defending their high-quality food resources. Indeed, they are known for their raucous 'roar-shriek' calls. However, there has been surprisingly little research on vocal communication in Varecia species and only two previously published repertoires, both of which were qualitative descriptions of their calls. In this study, we quantitatively examined the vocal repertoire of wild black-and-white ruffed lemurs (Varecia variegata) at Mangevo, Ranomafana National Park, Madagascar. We characterized 11 call types using 33 bioacoustic parameters related to frequency, duration, tonality, and composition. We also used discriminant function analysis and hierarchical clustering to quantitatively and objectively classify call types within the black-and-white ruffed lemur vocal repertoire. The repertoire consists of both monosyllabic and multisyllabic calls that are individually given or emitted in contagious choruses. Eight of the 11 calls were also used in combination or in larger multi-call sequences. The discriminant function analysis correctly assigned call types with 87% success, though this varied greatly by call type (1-65%). Hierarchical clustering identified 3-4 robust clusters, indicating low clustering structure in the data and suggesting that V. variegata exhibits a graded vocal repertoire. Future work should consider the environmental and behavioral contexts in which calls are used to better understand the function of these call types and combinations.

Factors affecting call usage in wild black-and-white ruffed lemurs (Varecia variegata) at Mangevo, Ranomafana National Park.

Vocal communication is an important modality for group-living primates inhabiting dense forest habitats that can hinder visual and olfactory signals. Nevertheless, research on primate vocalizations has historically focused on a narrow subset of haplorhine taxa; comparatively few studies have been focused on strepsirrhines, despite facing similar ecological and social challenges. Ruffed lemurs (Varecia)-a taxon known for their raucous calls-are rainforest specialists that exhibit strong fission-fusion dynamics and communally rear large litters of young. However, surprisingly few studies have examined Varecia vocalizations in the wild, meaning virtually nothing is known about the call types or how they facilitate their unique social and reproductive strategies. Our goal for this study was to examine how various contextual factors such as weather, behavioral state, and subgroup size and composition affected vocal activity across call types in wild black-and-white ruffed lemurs. We conducted focal follows on 31 individuals (two communities) in Mangevo (Ranomafana National Park) from May-August 2019 to record behavioral and vocal activity. We distinguished 11 call types, although three (hum, roar-shriek, chatter) constituted the majority of vocal activity. Calling rates were consistent throughout the day, but decreased with high rainfall. We found sex- and subgroup-specific differences in call usage, likely related to female dominance and subgroup composition, respectively. We identified behavioral contexts that some call types were consistently given in; this can be used to help infer call function. This study provides some of the first quantitative analyses of ruffed lemur vocal communication and lays the groundwork for more systematic hypothesis testing in future studies.