Acoustic correlates of caller identity and affect intensity in the vowel-like grunt vocalizations of baboons

Comparative, production-based research on animal vocalizations can allow assessments of continuity in vocal communication processes across species, including humans, and may aid in the development of general frameworks relating specific constitutional attributes of callers to acoustic-structural details of their vocal output. Analyses were undertaken on vowel-like baboon grunts to examine variation attributable to caller identity and the intensity of the affective state underlying call production. Six hundred six grunts from eight adult females were analyzed. Grunts derived from 128 bouts of calling in two behavioral contexts: concerted group movements and social interactions involving mothers and their young infants. Each context was subdivided into a high- and low-arousal condition. Thirteen acoustic features variously predicted to reflect variation in either caller identity or arousal intensity were measured for each grunt bout, including tempo-, source- and filter-related features. Grunt bouts were highly individually distinctive, differing in a variety of acoustic dimensions but with some indication that filter-related features contributed disproportionately to individual distinctiveness. In contrast, variation according to arousal condition was associated primarily with tempo- and source-related features, many matching those identified as vehicles of affect expression in other nonhuman primate species and in human speech and other nonverbal vocal signals.     

Perceptual classification based on the component structure of song in European starlings

The ability to recognize individuals based on their vocalizations is common among many species of songbirds. Examining the psychological and neural basis of this functionally relevant behavior can provide insight into the perceptual processing of acoustically complex, real-world, communication signals. In one species of songbird, European starlings (Sturnus vulgaris), males sing long and acoustically complex songs composed of small stereotyped note clusters called motifs. Previous studies demonstrate that starlings are capable of individual vocal recognition, and suggest that vocal recognition results from the association of specific motifs with specific individuals. The present study tests this possibility by examining how variation among the motifs that comprise a song affect its discrimination and classification. Starlings were trained, using operant techniques, to associate multiple songs from a single male starling with one response, and songs from four other male starlings with another response. The level of stimulus control exerted by motif variation was then measured by having subjects classify three sets of novel song bouts in which motifs from the training songs were systematically recombined. The results demonstrate a significant, and approximately linear, relationship between song classification and the relative proportions of familiar motifs from different singers that compose a bout. The results also indicate that the motif proportion effects on song classification are primary to retroactive interference in the recall for specific motifs, and independent of any biases due to the syntactic organization of motifs within a bout. Together, the results of this study suggest that starlings organize the complex vocalizations of conspecifics by memorizing large numbers of unique song components (i.e., motifs) that are then associated with different classes. Because individual starlings tend to possess unique motif repertoires, it is likely that under natural conditions such classes will correspond to individual identity. Thus, it is likely that perceptual processing mechanisms similar to those described by the results of the present study form the basis for individual vocal recognition in starlings.     

Learned vocalizations in budgerigars (Melopsittacus undulatus): the relationship between contact calls and warble song

The budgerigar (Melopsittacus undulatus) has an extraordinarily complex, learned, vocal repertoire consisting of both the long rambling warble song of males and a number of short calls produced by both sexes. In warble, the most common elements (>30%) bear a strong resemblance to the highly frequency-modulated, learned contact calls that the birds produce as single utterances. However, aside from this apparent similarity, little else is known about the relationship between contact calls and warble call elements. Here, both types of calls were recorded from four male budgerigars. Signal analysis and psychophysical testing procedures showed that the acoustic features of these two vocalizations were acoustically different and perceived as distinctive vocalizations by birds. This suggests that warble call elements are not simple insertions of contact calls but are most likely different acoustic elements, created de novo, and used solely in warble. Results show that, like contact calls, warble call elements contain information about signaler identity. The fact that contact calls and warble call elements are acoustically and perceptually distinct suggests that they probably represent two phonological systems in the budgerigar vocal repertoire, both of which arise by production learning.     

Affect cues in vocalizations of the bat, Megaderma lyra, during agonistic interactions

Some features of emotional prosody in human speech may be traced back to affect cues in mammalian vocalizations. The present study addresses the question whether affect intensity, as expressed by the intensity of behavioral displays, is encoded in vocal cues, i.e., changes in the structure of associated calls, in bats, a group evolutionarily remote from primates. A frame-by-frame video analysis of 109 dyadic agonistic interactions recorded in approach situations was performed to categorize displays into two intensity levels based on a cost-benefit estimate. M. lyra showed graded visual displays accompanied by specific calls and response calls of the second bat. A sound analysis revealed systematic changes of call sequence parameters with display level. At the high intensity level, total call duration, number of syllables within a call, and the number of calls within a sequence were increased, while intervals between call syllables were decreased for both call types. In addition, the latency of the response call was shorter, and its main syllable-type durations and fundamental frequency were increased. These systematic changes of vocal parameters with affect intensity correspond to prosodic changes in human speech, suggesting that emotion-related acoustic cues are a common feature of vocal communication in mammals.     

Acoustic correlates of individuality in the vocal repertoire of a nocturnal primate (Microcebus murinus)

In mammals individual distinctiveness in vocalizations provides the basis for individual recognition and thus plays an important role in social behavior. In this study, first evidence is provided for a nocturnal primate that variation in individual distinctiveness across the vocal repertoire is to some extent determined by the context and the acoustic structure of the call types. Individual distinctiveness was investigated across call types in the gray mouse lemur, a nocturnal primate, living in a dispersed multi-male multi-female social system. To explore to what degree context and acoustic structure predict variations in individual distinctiveness, four major call types were examined (grunts, tsaks, short whistles, and trills). Call types differed in context and acoustic structure and were recorded under controlled experimental conditions. A discriminant function analysis revealed that all call types are individually distinct, but not to the same degree. The findings suggest that variations in individual distinctiveness can to some extent be explained by the context and the acoustic structure of the call types.     

Vocal learning in Budgerigars (Melopsittacus undulatus): effects of an acoustic reference on vocal matching

Budgerigars were trained to produce specific vocalizations (calls) using operant conditioning and food reinforcement. The bird's call was compared to a digital representation of the call stored in a computer to determine a match. Once birds were responding at a high level of precision, we measured the effect of several manipulations upon the accuracy and the intensity of call production. Also, by differentially reinforcing other aspects of vocal behavior, budgerigars were trained to produce a call that matched another bird's contact call and to alter the latency of their vocal response. Both the accuracy of vocal matching and the intensity level of vocal production increased significantly when the bird could hear the template immediately before each trial. Moreover, manipulating the delay between the presentation of an acoustic reference and the onset of vocal production did not significantly affect either vocal intensity or matching accuracy. Interestingly, the vocalizations learned and reinforced in these operant experiments were only occasionally used in more natural communicative situations, such as when birds called back and forth to one another in their home cages.     

Within-individual variation in bullfrog vocalizations: implications for a vocally mediated social recognition system

Acoustic signals provide a basis for social recognition in a wide range of animals. Few studies, however, have attempted to relate the patterns of individual variation in signals to behavioral discrimination thresholds used by receivers to discriminate among individuals. North American bullfrogs (Rana catesbeiana) discriminate among familiar and unfamiliar individuals based on individual variation in advertisement calls. The sources, patterns, and magnitudes of variation in eight acoustic properties of multiple-note advertisement calls were examined to understand how patterns of within-individual variation might either constrain, or provide additional cues for, vocal recognition. Six of eight acoustic properties exhibited significant note-to-note variation within multiple-note calls. Despite this source of within-individual variation, all call properties varied significantly among individuals, and multivariate analyses indicated that call notes were individually distinct. Fine-temporal and spectral call properties exhibited less within-individual variation compared to gross-temporal properties and contributed most toward statistically distinguishing among individuals. Among-individual differences in the patterns of within-individual variation in some properties suggest that within-individual variation could also function as a recognition cue. The distributions of among-individual and within-individual differences were used to generate hypotheses about the expected behavioral discrimination thresholds of receivers.     

Individual variation in the pup attraction call produced by female Australian fur seals during early lactation

Otariid seals (fur seals and sea lions) are colonial breeders with large numbers of females giving birth on land during a synchronous breeding period. Once pups are born, females alternate between feeding their young ashore and foraging at sea. Upon return, both mother and pup must relocate each other and it is thought to be primarily facilitated by vocal recognition. Vocalizations of thirteen female Australian fur seals (Arctocephalus pusillus doriferus) were recorded during the breeding seasons of December 2000 and 2001, when pups are aged from newborns to one month. The pup attraction call was examined to determine whether females produce individually distinct calls which could be used by pups as a basis for vocal recognition. Potential for individual coding, discriminant function analysis (DFA), and classification and regression tree analysis were used to determine which call features were important in separating individuals. Using the results from all three analyses: F0, MIN F and DUR were considered important in separating individuals. In 76% of cases, the PAC was classified to the correct caller, using DFA, suggesting that there is sufficient stereotypy within individual calls, and sufficient variation between them, to enable vocal recognition by pups of this species.     

Influence of anabolic steroid treatment associated to physical exercise in the ultrasonic vocalization of rats

Unlike humans, who communicate in frequency bands between 250 Hz and 6 kHz, rats can communicate in frequencies above 18 kHz. Their vocalization types depend on the context and are normally associated to subjective or emotional states. It was reported significant vocal changes due to administration of replacement testosterone in a trained tenor singer with hypogonadism. Speech-Language Pathology clinical practices are being sought by singers who sporadically use anabolic steroids associated with physical exercise. They report difficulties in reaching and keeping high notes, "breakage" in the passage of musical notes and post singing vocal fatigue. Those abnormalities could be raised by the association of anabolic steroids and physical exercise. Thus, in order to verify if this association could promote vocal changes, maximum, minimum and fundamental frequencies and call duration in rats treated with anabolic steroids and physically trained (10 weeks duration) were evaluated. The vocalizations were obtained by handling the animals. At the end of that period, rats treated and trained showed significant decrease in call duration, but not in other parameters. The decrease in call duration could be associated to functional alterations in the vocal folds of treated and trained animals due to a synergism between anabolic steroids and physical training.     

Acoustic analysis of vocal development in a New World primate, the common marmoset (Callithrix jacchus)

In contrast to humans and songbirds, there is limited evidence of vocal learning in nonhuman primates. While previous studies suggested that primate vocalizations exhibit developmental changes, detailed analyses of the extent and time course of such changes across a species' vocal repertoire remain limited. In a highly vocal primate, the common marmoset (Callithrix jacchus), we studied developmental changes in the acoustic structure of species-specific communication sounds produced in a social setting. We performed detailed acoustic analyses of the spectral and temporal characteristics of marmoset vocalizations during development, comparing differences between genders and twin pairs, as well as with vocalizations from adult marmosets residing in the same colony. Our analyses revealed significant changes in spectral and temporal features as well as variability of particular call types over time. Infant and juvenile vocalizations changed progressively toward the vocalizations produced by adult marmosets. Call types observed early in development that were unique to infants disappeared gradually with age, while vocal exchanges with conspecifics emerged. Our observations clearly indicate that marmoset vocalizations undergo both qualitative and quantitative postnatal changes, establishing the basis for further studies to delineate contributions from maturation of the vocal apparatus and behavioral experience.     

Acoustics and behavioral contexts of "gecker" vocalizations in young rhesus macaques (Macaca mulatta)

Loud, pulsed "gecker" vocalizations are commonly produced by young rhesus macaques in distressful circumstances. The acoustics, usage, and responses associated with these calls were examined using audio recordings and observational data from captive, socially living rhesus up to 24 months old. One-hundred-eleven gecker bouts were recorded from ten individuals (six males, four females), with most geckers produced during the first 6 months of age. A gecker call consisted of a bout of up to 28 pulses of spectrally structured noise with a single prominent frequency peak. Nine contexts of calling were identified, but little evidence of context-specific acoustic variation was found. While geckering often triggered responses by the vocalizer's mother, the most common outcome was the absence of any reaction. Females geckered longer and at higher rates than did males, while also showing acoustic evidence of greater vocal effort. Mothers nonetheless responded more often and more positively to males. Overall, results show that gecker acoustics vary somewhat with vocalizer sex, age, and likely arousal level, but do not reflect detailed aspects of behavioral context. Circumstances of production suggest that geckers function primarily to draw the attention of mothers, who in turn are selective in responding.     

Vocal cues indicate level of arousal in infant African elephant roars

Arousal-based physiological changes influence acoustic features of vocalizations in mammals. In particular, nonlinear phenomena are thought to convey information about the caller's arousal state. This hypothesis was tested in the infant African elephant (Loxodonta africana) roar, a call type produced in situations of arousal and distress. Ninety-two percent of roars exhibited nonlinear phenomena, with chaos being the most common type. Acoustic irregularities were strongly associated with elevated fundamental frequency values. Roars produced in situations of highest urgency, based on the occurrence of behavioral indicators of arousal, were characterized by the lowest harmonics-to-noise ratio; this indicates low tonality. In addition, roars produced in these situations lasted longer than those produced in contexts of lower presumed urgency. Testing the infant roars for individual distinctiveness revealed only a moderate classification result. Combined, these findings indicate that infant African elephant roars primarily function to signal the caller's arousal state. The effective communication of this type of information may allow mothers to respond differentially based on their infant's degree of need and may be crucial for the survival of infant African elephants in their natural environment.     

Characteristics of gunshot sound displays by North Atlantic right whales in the Bay of Fundy

North Atlantic right whales (Eubalaena glacialis) produce a loud, broadband signal referred to as the gunshot sound. These distinctive sounds may be suitable for passive acoustic monitoring and detection of right whales; however, little is known about the prevalence of these sounds in important right whale habitats, such as the Bay of Fundy. This study investigates the timing and distribution of gunshot sound production on the summer feeding grounds using an array of five marine acoustic recording units deployed in the Bay of Fundy, Canada in mid-summer 2004 and 2005. Gunshot sounds were common, detected on 37 of 38 recording days. Stereotyped gunshot bouts averaged 1.5 h, with some bouts exceeding 7 h in duration with up to seven individuals producing gunshots at any one time. Bouts were more commonly detected in the late afternoon and evening than during the morning hours. Locations of gunshots in bouts indicated that whales producing the sounds were either stationary or showed directional travel, suggesting gunshots have different communication functions depending on behavioral context. These results indicate that gunshots are a common right whale sound produced during the summer months and are an important component in the acoustic communication system of this endangered species.     

Preliminary evidence for signature vocalizations among free-ranging narwhals (Monodon monoceros)

Animal signature vocalizations that are distinctive at the individual or group level can facilitate recognition between conspecifics and re-establish contact with an animal that has become separated from its associates. In this study, the vocal behavior of two free-ranging adult male narwhals (Monodon monoceros) in Admiralty Inlet, Baffin Island was recorded using digital archival tags. These recording instruments were deployed when the animals were caught and held onshore to attach satellite tags, a protocol that separated them from their groups. The signature content of two vocal categories was considered: (1) combined tonal/pulsed signals, which contained synchronous pulsatile and tonal content; (2) whistles, or frequency modulated tonal signals with harmonic energy. Nonparametric comparisons of the temporal and spectral features of each vocal class revealed significant differences between the two individuals. A separate, cross-correlation measure conducted on the whistles that accounted for overall contour shape and absolute frequency content confirmed greater interindividual compared to intraindividual differences. These data are consistent with the hypothesis that narwhals produce signature vocalizations that may facilitate their reunion with group members once they become separated, but additional data are required to demonstrate this claim more rigorously.     

The social vocalization repertoire of east Australian migrating humpback whales (Megaptera novaeangliae)

Although the songs of humpback whales have been extensively studied, other vocalizations and percussive sounds, referred to as "social sounds," have received little attention. This study presents the social vocalization repertoire of migrating east Australian humpback whales from a sample of 660 sounds recorded from 61 groups of varying composition, over three years. The social vocalization repertoire of humpback whales was much larger than previously described with a total of 34 separate call types classified aurally and by spectrographic analysis as well as statistically. Of these, 21 call types were the same as units of the song current at the time of recording but used individually instead of as part of the song sequence, while the other 13 calls were stable over the three years of the study and were not part of the song. This study provides a catalog of sounds that can be used as a basis for future studies. It is an essential first step in determining the function, contextual use and cultural transmission of humpback social vocalizations.     

Classification of behavior using vocalizations of Pacific white-sided dolphins (Lagenorhynchus obliquidens)

Surface behavior and concurrent underwater vocalizations were recorded for Pacific white-sided dolphins in the Southern California Bight (SCB) over multiple field seasons spanning 3 years. Clicks, click trains, and pulsed calls were counted and classified based on acoustic measurements, leading to the identification of 19 key call features used for analysis. Kruskal-Wallis tests indicated that call features differ significantly across behavioral categories. Previous work had discovered two distinctive click Types (A and B), which may correspond to known subpopulations of Pacific white-side dolphins in the Southern California Bight; this study revealed that animals producing these different click types also differ in both their behavior and vocalization patterns. Click Type A groups were predominantly observed slow traveling and milling, with little daytime foraging, while click Type B groups were observed traveling and foraging. These behavioral differences may be characteristic of niche partitioning by overlapping populations; coupled with differences in vocalization patterns, they may signify that these subpopulations are cryptic species. Finally, random forest decision trees were used to classify behavior based on vocalization data, with rates of correct classification up to 86%, demonstrating the potential for the use of vocalization patterns to predict behavior.     

Acoustic identification of female Steller sea lions (Eumetopias jubatus)

Steller sea lion (Eumetopias jubatus) mothers and pups establish and maintain contact with individually distinctive vocalizations. Our objective was to develop a robust neural network to classify females based on their mother-pup contact calls. We catalogued 573 contact calls from 25 females in 1998 and 1323 calls from 46 females in 1999. From this database, a subset of 26 females with sufficient samples of calls was selected for further study. Each female was identified visually by marking patterns, which provided the verification for acoustic identification. Average logarithmic spectra were extracted for each call, and standardized training and generalization datasets created for the neural network classifier. A family of backpropagation networks was generated to assess relative contribution of spectral input bandwidth, frequency resolution, and network architectural variables to classification accuracy. The network with best overall generalization accuracy (71%) used an input representation of 0-3 kHz of bandwidth at 10.77 Hz/bin frequency resolution, and a 2:1 hidden:output layer neural ratio. The network was analyzed to reveal which portions of the call spectra were most influential for identification of each female. Acoustical identification of distinctive female acoustic signatures has several potentially important conservation applications for this endangered species, such as rapid survey of females present on a rookery.     

Intra- and intergroup vocal behavior in resident killer whales, Orcinus orca

Vocal communication within and between groups of individuals has been described extensively in birds and terrestrial mammals, however, little is known about how cetaceans utilize their sounds in their natural environment. Resident killer whales, Orcinus orca, live in highly stable matrilines and exhibit group-specific vocal dialects. Single call types cannot exclusively be associated with particular behaviors and calls are thought to function in group identification and intragroup communication. In the present study call usage of three closely related matrilines of the Northern resident community was compared in various intra- and intergroup contexts. In two out of the three matrilines significant changes in vocal behavior depending both on the presence and identity of accompanying whales were found. Most evidently, family-specific call subtypes, as well as aberrant and variable calls, were emitted at higher rates, whereas "low arousal" call types were used less in the presence of matrilines from different pods, subclans, or clans. Ways in which the observed changes may function both in intra- and intergroup communication.     

Gaussian mixture model classification of odontocetes in the Southern California Bight and the Gulf of California

A method for the automatic classification of free-ranging delphinid vocalizations is presented. The vocalizations of short-beaked and long-beaked common (Delphinus delphis and Delphinus capensis), Pacific white-sided (Lagenorhynchus obliquidens), and bottlenose (Tursiops truncatus) dolphins were recorded in a pelagic environment of the Southern California Bight and the Gulf of California over a period of 4 years. Cepstral feature vectors are extracted from call data which contain simultaneous overlapping whistles, burst-pulses, and clicks from a single species. These features are grouped into multisecond segments. A portion of the data is used to train Gaussian mixture models of varying orders for each species. The remaining call data are used to test the performance of the models. Species are predicted based upon probabilistic measures of model similarity with test segment groups having durations between 1 and 25 s. For this data set, 256 mixture Gaussian mixture models and segments of at least 10 s of call data resulted in the best classification results. The classifier predicts the species of groups with 67%-75% accuracy depending upon the partitioning of the training and test data.