Caller sex and orientation influence spectral characteristics of "two-voice" stereotyped calls produced by free-ranging killer whales

This study investigates how particular received spectral characteristics of stereotyped calls of sexually dimorphic adult killer whales may be influenced by caller sex, orientation, and range. Calls were ascribed to individuals during natural behavior using a towed beamforming array. The fundamental frequency of both high-frequency and low-frequency components did not differ consistently by sex. The ratio of peak energy within the fundamental of the high-frequency component relative to summed peak energy in the first two low-frequency component harmonics, and the number of modulation bands off the high-frequency component, were significantly greater when whales were oriented towards the array, while range and adult sex had little effect. In contrast, the ratio of peak energy in the first versus second harmonics of the low-frequency component was greater in calls produced by adult females than adult males, while orientation and range had little effect. The dispersion of energy across harmonics has been shown to relate to body size or sex in terrestrial species, but pressure effects during diving are thought to make such a signal unreliable in diving animals. The observed spectral differences by signaler sex and orientation suggest that these types of information may be transmitted acoustically by freely diving killer whales.     

Vocal development in captive harbor seal pups, Phoca vitulina richardii: age, sex, and individual differences

The goal of this research was to determine when harbor seal pup vocalizations become sufficiently distinctive to allow individual recognition. A total of 4593 calls were analyzed from 15 captive pups. Nineteen were harsh, broadband, staccato calls used in an aggressive context. The rest were tonal "mother attraction calls," having an inverted "v"- or "u"-shaped spectrogram with harmonics and a fundamental frequency around 200-600 Hz. Calls were individually distinctive even in pups less than 2 weeks old, suggesting that mothers may be able to recognize pup vocalizations at this early age. Classification rates from discriminant function analysis were generally comparable to those of other phocids and less than in otariids, supporting the theory that recognition is more highly developed in otariids. Significant differences were found between male and female pup calls, and there were significant interactions between pup sex and age. The results of this study should be interpreted with caution until the findings are verified in wild harbor seal pups.     

Age-related differences in the acoustic characteristics of male leopard seals, Hydrurga leptonyx

During the breeding season, the underwater vocalizations and calling rates of adult male leopard seals are highly stereotyped. In contrast, sub-adult males have more variable acoustic behavior. Although adult males produce only five stereotyped broadcast calls as part of their long-range underwater breeding displays the sub-adults have a greater repertoire including the adult-like broadcast calls, as well as variants of these. Whether this extended repertoire has a social function is unknown due to the paucity of behavioral data for this species. The broadcast calls of the sub-adults are less stereotyped in their acoustic characteristics and they have a more variable calling rate. These age-related differences have major implications for geographic variation studies, where the acoustic behavior of different populations are compared, as well as for acoustic surveying studies, where numbers of calls are used to indicate numbers of individuals present. Sampling regimes which unknowingly include recordings from sub-adult animals will artificially exaggerate differences between populations and numbers of calling animals. The acoustic behavior of sub-adult and adult male leopard seals were significantly different and although this study does not show evidence that these differences reflect vocal learning in the male leopard seal it does suggest that contextual learning may be present.     

Vocal production mechanisms in the budgerigar (Melopsittacus undulatus): the presence and implications of amplitude modulation.

In this paper acoustic evidence is presented for the presence of amplitude modulation in budgerigar (Melopsittacus undulatus) contact calls and learned English vocalizations. Previously, acoustic analyses of budgerigar vocalizations have consisted solely of visual inspection of spectrograms or power spectra (derived from Fourier transformation). Such analyses have led researchers to conclude that budgerigar vocalizations are primarily frequency-modulated, harmonic vocalizations. Although budgerigar calls have been shown to contain regions that are modulated in amplitude, the implications of this fact have been largely ignored. Amplitude modulation, the nonlinear interaction between two separate signals that results in the creation of new, heterodyne (sum and difference) frequencies, can produce a very complex Fourier spectrum that may resemble that produced by a harmonic vocalization. In this paper, the acoustic principles necessary for identifying amplitude modulation present in signals are outlined, and followed by data demonstrating that amplitude modulation is a prominent feature not only of natural budgerigar contact calls, but also of their learned English vocalizations. It is illustrated how analyzing a vocalization that contains amplitude modulation as if it were harmonic can result in misinterpretations of the acoustic and physical properties of the sound and sound source. The implications of amplitude modulation for studies of the ontogenetic, physical, and neural basis of budgerigar vocalizations are discussed, and a potential model for how the budgerigar syrinx may function to produce amplitude modulation is proposed.     

Analysis of pain-related vocalization in young pigs

The assessment of pain constitutes a major issue for animal welfare research. The objective of this study was to classify vocalizations during castration pain and to assess alterations in vocalizations under local anaesthesia. The alterations in vocalization were measured by multiparametric call analysis. A total of 4537 calls of 70 young pigs were evaluated. With the data of this study three call types are distinguishable (grunt, squeal, scream). A high percentage (94.64%) of calls that could be classified in one of the three call types during the castration process within the confidence level of 95% was found. The comparison of the occurrence of the call types during treatments gives evidence for pain-related use of screams. The piglets castrated without local anaesthesia produced almost double the number of screams as piglets castrated with anaesthesia. The comparison of the recorded sound parameters reveals the particular position of screams in the call repertoire of young pigs. Screams are significantly different in their sound parameters than grunts or squeals. Castration in comparison to mere restraint produced a comprehensive change in sound parameters, with castration calls becoming more extended and more powerful. The findings in this study also show differences in the effectiveness of the parameters which indicate pain. Parameters that describe a single event in a call, such as peak level or peak frequency give better results than parameters that describe an average, such as weighted frequency and main frequency. The research indicated that pain-related changes of calls in piglets can be identified. On the basis of the results, automatic classification of call types during management operations may be developed. This could contribute to objective animal welfare assessment.     

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.     

Finding good acoustic features for parrot vocalizations: the feature generation approach

A crucial step in the understanding of vocal behavior of birds is to be able to classify calls in the repertoire into meaningful types. Methods developed to this aim are limited either because of human subjectivity or because of methodological issues. The present study investigated whether a feature generation system could categorize vocalizations of a bird species automatically and effectively. This procedure was applied to vocalizations of African gray parrots, known for their capacity to reproduce almost any sound of their environment. Outcomes of the feature generation approach agreed well with a much more labor-intensive process of a human expert classifying based on spectrographic representation, while clearly out-performing other automated methods. The method brings significant improvements in precision over commonly used bioacoustical analyses. As such, the method enlarges the scope of automated, acoustics-based sound classification.     

Cepstral coefficients and hidden Markov models reveal idiosyncratic voice characteristics in red deer (Cervus elaphus) stags

Bouts of vocalizations given by seven red deer stags were recorded over the rutting period, and homomorphic analysis and hidden Markov models (two techniques typically used for the automatic recognition of human speech utterances) were used to investigate whether the spectral envelope of the calls was individually distinctive. Bouts of common roars (the most common call type) were highly individually distinctive, with an average recognition percentage of 93.5%. A "temporal" split-sample approach indicated that although in most individuals these identity cues held over the rutting period, the ability of the models trained with the bouts of roars recorded early in the rut to correctly classify later vocalizations decreased as the recording date increased. When Markov models trained using the bouts of common roars were used to classify other call types according to their individual membership, the classification results indicated that the cues to identity contained in the common roars were also present in the other call types. This is the first demonstration in mammals other than primates that individuals have vocal cues to identity that are common to the different call types that compose their vocal repertoire.     

Developmental changes in ultrasonic vocalizations by infant Japanese echolocating bats, Pipistrellus abramus

Developmental changes in vocalizations by Pipistrellus abramus were investigated during the first post-natal month. Vocalizations by pups on the day of birth were frequency-modulated ultrasounds from 30.0 ± 4.0 kHz to 19.3 ± 1.9 kHz with multiple harmonics. The terminal frequency of the second harmonic (TF(2)) of pup vocalizations corresponded to that of the fundamental (TF(1)) in adult bats (41.4 ± 2.6 kHz), suggesting that pup vocalizations can be easily detected by the mother. In addition, there are two types of infant vocalization: short duration echolocation precursor and long duration isolation calls, which showed separate developmental patterns over time.     

Sex differences in the acoustic structure of vowel-like grunt vocalizations in baboons and their perceptual discrimination by baboon listeners

This study quantifies sex differences in the acoustic structure of vowel-like grunt vocalizations in baboons (Papio spp.) and tests the basic perceptual discriminability of these differences to baboon listeners. Acoustic analyses were performed on 1028 grunts recorded from 27 adult baboons (11 males and 16 females) in southern Africa, focusing specifically on the fundamental frequency (F0) and formant frequencies. The mean F0 and the mean frequencies of the first three formants were all significantly lower in males than they were in females, more dramatically so for F0. Experiments using standard psychophysical procedures subsequently tested the discriminability of adult male and adult female grunts. After learning to discriminate the grunt of one male from that of one female, five baboon subjects subsequently generalized this discrimination both to new call tokens from the same individuals and to grunts from novel males and females. These results are discussed in the context of both the possible vocal anatomical basis for sex differences in call structure and the potential perceptual mechanisms involved in their processing by listeners, particularly as these relate to analogous issues in human speech production and perception.     

Responses of squirrel monkeys to their experimentally modified mobbing calls

Previous acoustic analyses suggested emotion-correlated changes in the acoustic structure of squirrel monkey (Saimiri sciureus) vocalizations. Specifically, calls given in aversive contexts were characterized by an upward shift in frequencies, often accompanied by an increase in amplitude. In order to test whether changes in frequencies or amplitude are indeed relevant for conspecific listeners, playback experiments were conducted in which either frequencies or amplitude of mobbing calls were modified. Latency and first orienting response were measured in playback experiments with six adult squirrel monkeys. After broadcasting yaps with increased frequencies or amplitude, squirrel monkeys showed a longer orienting response towards the speaker than after the corresponding control stimuli. Furthermore, after broadcasting yaps with decreased frequencies or amplitude, squirrel monkeys showed a shorter orienting response towards the speaker than after the corresponding manipulated calls with higher frequencies or amplitude. These results suggest that changes in frequencies or amplitude were perceived by squirrel monkeys, indicating that the relationship between call structure and the underlying affective state of the caller agreed with the listener's assessment of the calls. However, a simultaneous increase in frequencies and amplitude did not lead to an enhanced response, compared to each single parameter. Thus, from the receiver's perspective, both call parameters may mutually replace each other.     

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.     

Laryngeal and respiratory activity during vocalization in macaque monkeys

The present study describes the laryngeal and respiratory muscle activity associated with vocalizations in macaque monkeys. During the bark vocalization, a short, aperiodic call, the cricothyroid, thyroarytenoid, rectus abdominis, and intercostals were active while the posterior cricoarytenoid and diaphragm were quiet. During the coo vocalization, a longer, clear call, the cricothyroid, thyroarytenoid, intercostals, rectus abdominis, and diaphragm were active. In one monkey, the posterior cricoarytenoid was also active during the call, while in another monkey it was not. Laryngeal muscle activity was correlated with the amplitude and duration of the coo call. Results suggest that the amplitude and duration differences between calls are determined primarily by laryngeal modification of the airflow, and that the differences in posterior cricoarytenoid activity may be due to differences in voice intensity.     

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.     

Antimasking aspects of harp seal (Pagophilus groenlandicus) underwater vocalizations

Underwater sounds are very important in social communication of harp seals (Pagophilus groenlandicus) because they are the main means of long- and short-distance communication. Individual harp seals must try to avoid being masked and emit only those calls that will benefit them. Underwater vocalizations of harp seals were recorded during the breeding season. The physical characteristics associated with antimasking attributes of 16 call types were examined. Rising frequency or increasing amplitude within calls were not common. Most of the calls ended abruptly (range 145-966 dB/s), but call onset was more gradual. At high calling rates (95.1-135 calls/min) there were significantly more calls overlapping temporally than at medium (75.1-95 calls/min) or low (35-75 calls/min) calling rates, but even at the highest calling rates, 79.1% of the calls were not overlapped. When 2, 3, or 4 calls overlapped, there were significantly fewer frequency separations of less than 1/3 octave than would be expected by chance. This is important because sounds that are separated by less than 1/3 octave likely mask each other. When 2-4 calls are occurring simultaneously, only 4.5% to 14.2% are masked by virtue of being within 1/3 octave from their nearest neighbor. None of the overlappping calls was of the same type. This suggests that the seals are actively listening to each other's calls and are not randomly using the different call types. Harp seals use frequency and temporal separation in conjunction with a wide vocal repertoire to avoid masking each other.     

Source of the North Pacific "boing" sound attributed to minke whales

During a recent cetacean survey of the U.S. waters surrounding the Hawaiian Islands, the probable source of the mysterious "boing" sound of the North Pacific Ocean was identified as a minke whale, Balaenoptera acutorostrata. Examination of boing vocalizations from three research surveys confirms previous work that identified two distinct boing vocalization types in the North Pacific. The eastern boing (n = 22) has a pulse repetition rate of 92 s(-1) and a duration of 3.6 s and was found only east of 138 degrees W. The central boing (n = 106) has a pulse repetition rate of 115 s(-1) and a duration of approximately 2.6 s and was found only west of 135 degrees W. Central boing vocalizations produced by a single source (n = 84) indicate that variation in repetition rate and duration of the calls of the individual were not significantly different than the variation among individuals of the same boing type. Despite a slight latitudinal overlap in the vocalizations, pulse repetition rates of the eastern and central boings were distinct.     

Perception of complex sounds in budgerigars (Melopsittacus undulatus) with temporary hearing loss

Songbirds and parrots deafened as nestlings fail to develop normal vocalizations, while birds deafened as adults show a gradual deterioration in the quality and precision of vocal production. Beyond this, little is known about the effect of hearing loss on the perception of vocalizations. Here, we induced temporary hearing loss in budgerigars with kanamycin and tested several aspects of the hearing, including the perception of complex, species-specific vocalizations. The ability of these birds to discriminate among acoustically distinct vocalizations was not impaired but the ability to make fine-grain discriminations among acoustically similar vocalizations was affected, even weeks after the basilar papilla had been repopulated with new hair cells. Interestingly, these birds were initially unable to recognize previously familiar contact calls in a classification task-suggesting that previously familiar vocalizations sounded unfamiliar with new hair cells. Eventually, in spite of slightly elevated absolute thresholds, the performance of birds on discrimination and perceptual recognition of vocalizations tasks returned to original levels. Thus, even though vocalizations may initially sound different with new hair cells, there are only minimal long-term effects of temporary hearing loss on auditory perception, recognition of species-specific vocalizations, or other aspects of acoustic communication in these birds.     

The effects of articulation on the acoustical structure of feline vocalizations

Feline isolation calls were analyzed, and a model was developed to relate the acoustical features of these calls to the physical processes used in their production. Fifty isolation calls were recorded from each of five cats for a total sample of 250 vocalizations. By combinations of Fourier transform, autocorrelation, and linear prediction methods, the fundamental frequency (glottal-pulse period) F0, the energy of F0, the frequency having maximum energy Fmax (not always F0), and the energy at this frequency were computed. Mean F0 ranged from 400-600 Hz for individual cats. For some cats F0 was consistent within calls, but for other cats sudden shifts in F0 occurred within calls. Here, Fmax was almost a harmonic of F0 and generally ranged from 1-2 kHz. For individual cats, the energy ratio E = (energy of Fmax/energy of F0) varied from 1 to 60 and the grand average E over the time course of the call varied from about 12 to 38. The mean rms call intensity was an inverted-U function of time. Measured jaw opening was strongly correlated with acoustical features of call. A Bessel-horn model with time-varying flare gave a good account of acoustical parameters such as Fmax. The presence of formantlike resonances in cat vocalizations and the important role of jaw movements (vocal gestures) in the production of these calls suggest that cats may provide a useful model for some aspects of human vocal behavior.     

Repetition patterns in Weddell seal (Leptonychotes weddellii) underwater multiple element calls

Many vocalizations produced by Weddell seals (Leptonychotes weddellii) are made up of repeated individual distinct sounds (elements). Patterning of multiple element calls was examined during the breeding season at Casey and Davis, Antarctica. Element and interval durations were measured from 405 calls all > 3 elements in length. The duration of the calls (22+/-16.6 s) did not seem to vary with an increasing number of elements (F4,404=1.83,p = 0.122) because element and interval durations decreased as the number of elements within a call increased. Underwater vocalizations showed seven distinct timing patterns of increasing, decreasing, or constant element and interval durations throughout the calls. One call type occurred with six rhythm patterns, although the majority exhibited only two rhythms. Some call types also displayed steady frequency changes as they progressed. Weddell seal multiple element calls are rhythmically repeated and thus the durations of the elements and intervals within a call occur in a regular manner. Rhythmical repetition used during vocal communication likely enhances the probability of a call being detected and has important implications for the extent to which the seals can successfully transmit information over long distances and during times of high level background noise.