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.     

Variation in chick-a-dee calls of tufted titmice, Baeolophus bicolor: note type and individual distinctiveness

The chick-a-dee call of chickadee species (genus Poecile) has been the focus of much research. A great deal is known about the structural complexity and the meaning of variation in notes making up calls in these species. However, little is known about the likely homologous "chick-a-dee" call of the closely related tufted titmouse, Baeolophus bicolor. Tufted titmice are a prime candidate for comparative analyses of the call, because their vocal and social systems share many characteristics with those of chickadees. To address the paucity of data on the structure of chick-a-dee calls of tufted titmice, we recorded birds in field and aviary settings. Four main note types were identified in the call: Z, A, D(h), and D notes. Several acoustic parameters of each note type were measured, and statistical analyses revealed that the note types are acoustically distinct from one another. Furthermore, note types vary in the extent of individual distinctiveness reflected in their acoustic parameters. This first step towards understanding the chick-a-dee call of tufted titmice indicates that the call is comparable in structure and complexity to the calls of chickadees.     

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.     

Feature weighting in "chick-a-dee" call notes of Poecile atricapillus

Artificial neural networks were trained to discriminate between different note types from the black-capped chickadee (Poecile atricapillus) "chick-a-dee" call. Each individual note was represented as a vector of summary features taken from note spectrograms and networks were trained to respond to exemplar notes of one type and to fail to respond to exemplar notes of another type. Following initial network training, the network was presented novel notes in which individual acoustic features had been modified. The strength of the response of the network to each novel and shifted note was recorded. When network responses were plotted as a function of the degree of acoustic feature modification and training context, it became clear that modifications of some acoustic features had significant effects on network responses, while others did not. Moreover, the training context of the network also played a role in the responses of networks to manipulated test notes. The implications of using artificial neural networks to generate testable hypotheses for animal research and the role of context are discussed.     

Alarm signals of the great gerbil: acoustic variation by predator context, sex, age, individual, and family group

The great gerbil, Rhombomys opinus, is a highly social rodent that usually lives in family groups consisting of related females, their offspring, and an adult male. The gerbils emit alarm vocalizations in the presence of diverse predators with different hunting tactics. Alarm calls were recorded in response to three predators, a monitor lizard, hunting dog, and human, to determine whether the most common call type, the rhythmic call, is functionally referential with regard to type of predator. Results show variation in the alarm calls of both adults and subadults with the type of predator. Discriminant function analysis classified an average of 70% of calls to predator type. Call variation, however, was not limited to the predator context, because signal structure also differed by sex, age, individual callers, and family groups. These variations illustrate the flexibility of the rhythmic alarm call of the great gerbil and how it might have multiple functions and communicate in multiple contexts. Three alarm calls, variation in the rhythmic call, and vibrational signals generated from foot-drumming provide the gerbils with a varied and multi-channel acoustic repertoire.     

Sounds produced by Norwegian killer whales, Orcinus orca, during capture

To date very little is still known about the acoustic behavior of Norwegian killer whales, in particular that of individual whales. In this study a unique opportunity was presented to document the sounds produced by five captured killer whales in the Vestfjord area, northern Norway. Individuals produced 14 discrete and 7 compound calls. Two call types were used both by individuals 16178 and 23365 suggesting that they may belong to the same pod. Comparisons with calls documented in Strager (1993) showed that none of the call types used by the captured individuals were present. The lack of these calls in the available literature suggests that call variability within individuals is likely to be large. This short note adds to our knowledge of the vocal repertoire of this population and demonstrates the need for further studies to provide behavioural context to these sounds.     

Acoustic Analyses of Developmental Changes and Emotional Expression in the Preverbal Vocalizations of Infants

The nonverbal vocal utterances of seven normally hearing infants were studied within their first year of life with respect to age- and emotion-related changes. Supported by a multiparametric acoustic analysis it was possible to distinguish one inspiratory and eleven expiratory call types. Most of the call types appeared within the first two months; some emerged in the majority of infants not until the 5th (“laugh”) or 7th month (“babble”). Age-related changes in acoustic structure were found in only 4 call types (“discomfort cry,” “short discomfort cry,” “wail,” “moan”). The acoustic changes were characterized mainly by an increase in harmonic-to-noise ratio and homogeneity of the call, a decrease in frequency range and a downward shift of acoustic energy from higher to lower frequencies. Emotion-related differences were found in the acoustic structure of single call types as well as in the frequency of occurrence of different call types. A change from positive to negative emotional state was accompanied by an increase in call duration, frequency range, and peak frequency (frequency with the highest amplitude within the power spectrum). Negative emotions, in addition, were characterized by a significantly higher rate of “crying,” “hic” and “ingressive vocalizations” than positive emotions, while positive emotions showed a significantly higher rate of “babble,” “laugh,” and “raspberry.”     

Quantitative acoustic analysis of the vocal repertoire of the golden rocket frog (Anomaloglossus beebei)

This study describes the vocal repertoire of the Guyanan golden rocket frog, Anomaloglossus beebei, a bromeliad specialist with biparental care. Using multivariate analyses of nine call properties, as well as the occurrence of nonlinear phenomena, three signal types were distinguished-advertisement, courtship, and aggressive calls. Although all three call types were composed of a short series of rapidly repeated pulses, advertisement calls were produced at higher amplitudes and had longer pulse durations than both courtship calls and aggressive calls. Courtship calls exhibited lower dominant frequencies than both advertisement and aggressive calls, which had similar dominant frequencies. Aggressive calls had more pulses per call, had longer intervals between calls, and occasionally contained one or two introductory pulses preceding the pulsed call. Several acoustic properties predicted aspects of the signaler's body size and condition. Our study demonstrates the reliability of human observers to differentiate the multiple call types of A. beebei based on hearing calls and observing the social context in which they are produced under field conditions.     

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.     

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.     

Divergence of a stereotyped call in northern resident killer whales

Northern resident killer whale pods (Orcinus orca) have distinctive stereotyped pulsed call repertoires that can be used to distinguish groups acoustically. Repertoires are generally stable, with the same call types comprising the repertoire of a given pod over a period of years to decades. Previous studies have shown that some discrete pulsed calls can be subdivided into variants or subtypes. This study suggests that new stereotyped calls may result from the gradual modification of existing call types through subtypes. Vocalizations of individuals and small groups of killer whales were collected using a bottom-mounted hydrophone array in Johnstone Strait, British Columbia in 2006 and 2007. Discriminant analysis of slope variations of a predominant call type, N4, revealed the presence of four distinct call subtypes. Similar to previous studies, there was a divergence of the N4 call between members of different matrilines of the same pod. However, this study reveals that individual killer whales produced multiple subtypes of the N4 call, indicating that divergence in the N4 call is not the result of individual differences, but rather may indicate the gradual evolution of a new stereotyped call.     

Sources of acoustic variation: implications for production specificity and call categorization in chacma baboon (Papio ursinus) grunts

Elucidating the information content of vocal signals is fundamental to the understanding of animal communication. Acoustically distinct calls produced in specific contexts allow listeners to predict future events and choose adequate responses. However, the vocal repertoires of most terrestrial mammals consist of a limited number of call types that vary within and between categories. These "graded signaling systems" are thought to be rich in information, at the cost of increasing uncertainty regarding call categorization. In this study, patterns of acoustic variation in grunts of wild chacma baboons (Papio ursinus) were assessed in relation to different contexts, callers' arousal, the presence of listeners, and individual identity. Although overall production specificity was low, and sensitive to the number of contexts under consideration, grunts given in three contexts could be statistically distinguished from each other. Contextual differences remained when controlling for caller arousal, suggesting that these differences cannot be explained by variation in arousal. No audience effect was detected, but individual identity was found to have an influence on acoustic structure. Overall, these results support the view that, in comparison to other signaling systems associated with hazardous conditions, lower production specificity might evolve under relaxed circumstances where unambiguous signaling is less important.     

Vocal behavior of resident killer whale matrilines with newborn calves: the role of family signatures

Studies of the vocal behavior of resident killer whales or orcas, Orcinus orca, in British Columbia have shown that matrilines have unique call repertoires consisting of up to 17 different call types. These call types cannot be attributed exclusively to specific behaviors, and their function in social contexts is poorly understood. This study investigated the change in call patterns of three resident matrilines in a changed social environment, before and up to one year after the birth of a calf. Acoustic data were collected with a network of hydrophones and were supplemented by visual observations. Call use changed distinctly after the birth of a calf in all three observed matrilines. All call types that were recorded in control situations were also recorded in postbirth situations; however, aberrant versions of discrete calls and excitement calls made up a higher proportion of calls after birth. Most conspicuously, family-specific call types occurred significantly more frequently in the days following a birth in two of the three matrilines and gradually returned to prebirth values within 2 weeks. Their increased use after a calf's birth may facilitate the learning process of this "acoustic family badge" and thereby help to recognize and maintain cohesion with family members.     

Quantifying complex patterns of bioacoustic variation: use of a neural network to compare killer whale (Orcinus orca) dialects

A quantitative measure of acoustic similarity is crucial to any study comparing vocalizations of different species, social groups, or individuals. The goal of this study was to develop a method of extracting frequency contours from recordings of pulsed vocalizations and to test a nonlinear index of acoustic similarity based on the error of an artificial neural network at classifying them. Since the performance of neural networks depends on the amount of consistent variation in the training data, this technique can be used to assess such variation from samples of acoustic signals. The frequency contour extraction and the neural network index were tested on samples of one call type shared by nine social groups of killer whales. For comparison, call similarity was judged by three human subjects in pairwise classification tasks. The results showed a significant correlation between the neural network index and the similarity ratings by the subjects. Both measures of acoustic similarity were significantly correlated with the groups' association patterns, indicating that both methods of quantifying acoustic similarity are biologically meaningful. An index based on neural network analysis therefore represents an objective and repeatable means of measuring acoustic similarity, and allows comparison of results across studies, species and time.     

Relationship between visual counts and call detection rates of gray whales (Eschrichtius robustus) in Laguna San Ignacio, Mexico

Daily acoustic calling rates of Eastern North Pacific (ENP) gray whales were measured on 6 days during 1 mo of their 2008 breeding season in the sheltered coastal lagoon of Laguna San Ignacio in Baja California, Mexico. Visual counts of whales determined that the numbers of single animals in the lower lagoon more than tripled over the observation period. All call types showed production peaks in the early morning and evening with minimum rates generally detected in the early afternoon. For four of the five observation days, the daily number of "S1"-type calls increased roughly as the square of the number of the animals in the lower lagoon during both daytime and nighttime. This relationship persisted when raw call counts were adjusted for variations in background noise level, using a simple propagation law derived from empirical measurements. The one observation day that did not fit the square-law relationship occurred during a week when the group size in the lagoon increased rapidly. These results suggest that passive acoustic monitoring does not measure gray whale group size directly but monitors the number of connections in the social network, which rises as roughly M(2)/2 for a group size M.     

American and Swedish children's acquisition of vowel duration: effects of vowel identity and final stop voicing

Vowel durations typically vary according to both intrinsic (segment-specific) and extrinsic (contextual) specifications. It can be argued that such variations are due to both predisposition and cognitive learning. The present report utilizes acoustic phonetic measurements from Swedish and American children aged 24 and 30 months to investigate the hypothesis that default behaviors may precede language-specific learning effects. The predicted pattern is the presence of final consonant voicing effects in both languages as a default, and subsequent learning of intrinsic effects most notably in the Swedish children. The data, from 443 monosyllabic tokens containing high-front vowels and final stop consonants, are analyzed in statistical frameworks at group and individual levels. The results confirm that Swedish children show an early tendency to vary vowel durations according to final consonant voicing, followed only six months later by a stage at which the intrinsic influence of vowel identity grows relatively more robust. Measures of vowel formant structure from selected 30-month-old children also revealed a tendency for children of this age to focus on particular acoustic contrasts. In conclusion, the results indicate that early acquisition of vowel specifications involves an interaction between language-specific features and articulatory predispositions associated with phonetic context.     

Vocalization control in Mongolian gerbils (Meriones unguiculatus) during locomotion behavior

The vocalization behavior of Mongolian gerbils, a model animal of auditory physiology, was examined. A pair of gerbils was placed in a chamber, and their species-specific vocalizations and locomotive behaviors were recorded and analyzed. Two types of calls were predominantly produced: high-frequency upward frequency-modulated (HU-FM) calls and low-frequency multi-harmonic frequency-modulated (LM-FM) calls. Emission rates of HU-FM calls significantly decreased as the distance between the two gerbils increased, and playback of simulated HU-FM calls increased the emission rates. Acoustic analysis of HU-FM calls showed that the calls exhibited a stereotypic spectro-temporal structure including a fixed inter-onset interval (100-175 ms) and that individual differences in the frequency could convey the body size of the callers. The timing of HU-FM calls was highly synchronized with jump movements when an animal vocalized while jumping, suggesting the existence of tight locomotor-vocal coupling. Conversely, LM-FM calls were observed only when the gerbils tactilely contacted with each other while fighting over a food. These results suggest that Mongolian gerbils change the rates of call emissions and call types (e.g., LM-FM or HU-FM calls) in response to changes in visual and possibly tactile and auditory information. The functions of both calls are discussed in terms of their acoustic structures.     

Dynamics of communal vocalizations in a social songbird, the zebra finch (Taeniopygia guttata)

Colonies or communities of animals such as fishes, frogs, seabirds, or marine mammals can be noisy. Although vocal communication between clearly identified sender(s) and receiver(s) has been well studied, the properties of the noisy sound that results from the acoustic network of a colony of gregarious animals have received less attention. The resulting sound could nonetheless convey some information about the emitting group. Using custom-written software for automatic detection of vocalizations occurring over many hours of recordings, this study reports acoustic features of communal vocal activities in a gregarious species, the zebra finch (Taeniopygia guttata). By biasing the sex ratio and using two different housing conditions (individual versus communal housing), six groups of zebra finches were generated, with six different social structures that varied both in terms of sex-composition and proportion of paired individuals. The results showed that the rate of emission and the acoustic dynamic both depended on the social structure. In particular, the vocal activity of a group of zebra finches depended mainly on the number of unpaired birds, i.e., individuals not part of a stably bonded pair.