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.     

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.     

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.     

Perceptual salience of individually distinctive features in the calls of adult king penguins

In the king penguin, Aptenodytes patagonicus, incubation and brooding duties are undertaken alternately by both partners of a pair. Birds returning from foraging at sea find their mate in the crowded colony using acoustic signals. Acoustic recognition of the mate maintains and strengthens the mate's fidelity and favors synchronization in the different stages of reproduction. In this study it was found that the king penguin vocalizes in response to the mate's playback calls, but not to those of neighbors or unfamiliar conspecific individuals. To study individual features used by the birds for individual recognition of mates, various experimental signals consisting of synthesized modifications of the mate' s call were played back to the incubating bird. Results indicated that birds attend to the FM profile of the call, in particular its initial inflexion. The frequency modulation shape of the syllable can be assimilated to a vocal signature repeated though the different syllables of the call. King penguins pay little attention to the call' s AM envelope or its absolute frequency.     

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.     

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.     

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.     

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.     

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.     

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.     

Call repertoire of infant African elephants: first insights into the early vocal ontogeny

African savannah elephants (Loxodonta africana) have a complex acoustic communication system, but very little is known about their vocal ontogeny. A first approach in ontogenetic studies is to define the call repertoire of specific age groups. Twelve hundred calls of 11 infant elephants from neonatal to 18 months of age recorded at the Vienna Zoo in Austria and at the Daphne Sheldrick's orphanage at the Nairobi National Park, Kenya were analyzed. Six call types were structurally distinguished: the rumble, the bark, the grunt, the roar (subdivided into a noisy-, tonal-, and mixed-roar), the snort, and the trumpet. Generally, within-call-type variation was high in all individuals. In contrast to adult elephants, the infants showed no gender-dependent variation in the structure or in the number of call types produced. Male infants, however, were more vocally adamant in their suckle behavior than females. These results give a first insight to the early vocal ontogeny and should promote further ontogenetic studies on elephants. Due to their vocal learning ability in combination with the complex fission-fusion society, elephants could be an interesting model to study the role of imitation in the vocal ontogeny of a nonprimate terrestrial mammal.     

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.     

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.     

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.     

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.     

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.     

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.     

Nonlinear acoustics in the pant hoots of common chimpanzees (Pan troglodytes): vocalizing at the edge

Common chimpanzee (Pan troglodytes) "pant hoots" are multi-call events that build from quiet, consistently harmonic introductory sounds to loud, screamlike "climax" calls with acoustic irregularities known as "nonlinear phenomena" (NLP). Two possible functions of NLP in climax calls are to increase direct auditory impact on listeners and to signal physical condition. These possibilities were addressed by comparing climax calls from 12 wild chimpanzee males with "screams" and pant hoot "introduction" calls from the same individuals. Climax calls that included NLP were found to have higher fundamental frequencies (F0s) than introduction or climax calls that were purely harmonic. NLP onsets within climax calls were also specifically associated with local F0 maxima, suggesting vocalizers are vibrating their vocal folds at the upper limits of stability. Furthermore, climax calls showed far fewer NLP than did screams recorded from the same individuals, while showing equivalent or higher F0 values. Overall, the results are consistent with the hypothesis that the relative prevalence of NLP is a signal of physical condition, with callers "vocalizing at the edge" of regular, stable production while producing few NLP. The results are discussed in light of the initial hypotheses.     

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.