Tonal vocalizations in the red wolf (Canis rufus): potential functions of nonlinear sound production.

The purpose of this study is to further understanding of the function of nonlinear vocalizations in red wolves (Canis rufus) by examining the acoustic, structural, and contextual characteristics of nonlinear sounds as compared to linear sounds. Video recordings of captive wolves from a breeding facility were analyzed. The acoustic nature of sound units was consistent with that of other social canids. The sound units included high-frequency squeaks (2600-9500 Hz) and low-frequency wuhs (160-1600 Hz) occurring either as separate units or in combination as nonlinear units (squeak-wuh frequency jumps, biphonations, squeaks with sidebands) and frequency jumps within squeaks. These low-amplitude sounds occurred in trains of 1-30 units that were classified as squeak vocalizations (49%), wuh vocalizations (19%), and nonlinear vocalizations (any combination including one or more nonlinear units, 32%). Nonlinear vocalizations transitioned directionally from high-frequency units to mixed-frequency units which has implications for the study of sound production and function. Wolves squeaked most often when oriented toward others, implying a solicitation function, while wuh vocalizations were more common during social interactions. Nonlinear vocalizations occurred most often during penmate-play or when oriented toward neighbors, indicating that nonlinear sound production may signal an increase in arousal.