The structure of vocal sounds produced with the mouth closed or with the mouth open in treefrogs

Frogs and toads mostly call with their mouths shut, unlike many other vertebrates. Sound is generated when air crosses the larynx, but there is no direct airflow to the external environment and radiation occurs at the skin. This study directly compares the acoustic output obtained from euthanized frogs with the mouth open against the output obtained with the mouth closed during activation of the larynx by airflow. With the mouth closed, the vocal sac was inflated and the acoustic energy was concentrated in the same harmonics as in the advertisement call, whereas with the mouth open, energy was spread in a wide range of harmonics. The acoustic output at the dominant frequency was more intense with the mouth closed than with the mouth open. More sound was radiated through the vocal sac and head than through the rest of the body. The spectral differences between open and closed mouth treatments matched the differences observed between natural advertisement calls, produced with the mouth closed, and distress calls, produced with the mouth open. By calling with the mouth closed, treefrogs can potentially produce advertisement calls with the energy concentrated in a narrower frequency range than with the mouth open.     

Nonoptimal propagation of advertisement calls of midwife toads in Iberian habitats

This study compares the efficiency of transmission of the advertisement calls of two species of midwife toads, Alytes cisternasii and A. obstetricans, in both native and non-native habitats in the Iberian Peninsula. Recorded calls of both species and pure tones were broadcast at ten sites native to either the relatively small A. cisternasii or the larger A. obstetricans. A large variation in the patterns of excess attenuation between localities was observed for calls measured at distances of 0.5 to 8 m from a loudspeaker. However, attenuation rates were higher for calls of both species in habitats of A. obstetricans relative to habitats of A. cisternasii. The calls of A. obstetricans experienced lower attenuation rates than those of A. cisternasii in both conspecific and heterospecific localities. Thus, although A. cisternasii occupies habitats more favorable for sound transmission, its advertisement call spectrum is not optimized for these habitats; the calls of A. obstetricans suffer less attenuation in A. cisternasii habitats. This result argues against the notion that spectral features of the calls are adapted to enhance transmission efficiency in natural habitats, and suggests that differences in call dominant frequency between the two species result from constraints imposed by selection on body size.     

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.     

Courtship and agonistic sounds by the cichlid fish Pseudotropheus zebra

Courtship and agonistic interactions in an African cichlid species present a richer diversity of acoustic stimuli than previously reported. Male cichlids, including those from the genus Pseudotropheus (P.), produce low frequency short pulsed sounds during courtship. Sounds emitted by P. zebra males in the early stages of courtship (during quiver) were found to be significantly longer and with a higher number of pulses than sounds produced in later stages. During agonistic intrasexual quiver displays, males produced significantly longer sounds with more pulses than females. Also, male sounds had a shorter duration and pulse period in courtship than in male-male interactions. Taken together, these results show that the acoustic repertoire of this species is larger than what was previously known and emphasize the importance of further research exploiting the role of acoustic stimuli in intra- and interspecific communication in African cichlids.     

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.     

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.     

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.     

Call recognition in the bullfrog, Rana catesbeiana: generalization along the duration continuum

Male bullfrogs emit multicroak, quasiharmonic advertisement calls that function in mate attraction and neighbor recognition. The degree of variability of acoustic features in these calls can influence perceptual decisions by conspecific receivers. Analysis of duration of individual croaks in spontaneous advertisement calls of a sample of males shows considerable intraindividual variability in this feature, even within short chorusing bouts. The influence of this intraindividual variability on behavior was examined in a series of evoked calling experiments. When presented with synthetic calls whose croak durations varied over the range of the natural variability in this feature, males responded similarly to intermediate and long duration croaks, but significantly less to short duration croaks. When presented with playbacks of calls with croak durations outside the natural range of variability, males again responded significantly less to shorter durations. The response gradient for duration is thus asymmetrical, with stimuli at the shorter end of the continuum evoking fewer responses than stimuli at the longer end. This asymmetry may be related to the biological demands of rejecting perception of heterospecific advertisement calls, and of mediating appropriate responses to conspecific aggressive calls. The shape of the response gradient for duration may reflect a process of stimulus generalization.     

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.     

Low-frequency whale sounds recorded on hydrophones moored in the eastern tropical Pacific

An array of autonomous hydrophones moored in the eastern tropical Pacific was monitored for one year to examine the occurrence of whale calls in this region. Six hydrophones which recorded from 0-40 Hz were placed at 8 degrees N, 0 degree, and 8 degrees S along longitudes 95 degrees W and 110 degrees W. Seven types of sounds believed to be produced by large whales were detected. These sound types were categorized as either moan-type (4) or pulse-type (3) calls. Three of the moan-type calls, and probably the fourth, may be attributed to blue whales. The source(s) of the remaining calls is unknown. All of the call types studied showed seasonal and geographical variation. There appeared to be segregation between northern and southern hemispheres, such that call types were recorded primarily on the northern hydrophones in the northern winter and others recorded primarily on the southern hemisphere hydrophones in the southern winter. More calls and more call types were recorded on the eastern hydrophones than on the western hydrophones.     

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.     

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.     

Sound production by North Atlantic right whales (Eubalaena glacialis) in surface active groups

The surface active group (SAG) is the most commonly observed surface social behavior of North Atlantic right whales. Recordings were made from 52 SAGs in the Bay of Fundy, Canada between July and September, from 1999 to 2002. The call types recorded from these groups were similar to those described previously for Southern right whales (Eubalaena australis), with six major call types being termed scream, gunshot, blow, upcall, warble, and downcall. The percentage of total calls of each call type depended on the group size and composition. The most common call type recorded was the scream call. The scream calls were produced by the focal female in a SAG. Production of other sound types can be attributed to whales other than the focal female, with gunshot and upcalls produced by males, and warble calls produced by female calves. The source levels for these sounds range from 137 to 162 dB rms re 1 ,tPa-m for tonal calls and 174 to 192 dB rms for broadband gunshot sounds.     

Sound characteristics of big-snout croaker, Johnius macrorhynus (Sciaenidae)

The sounds of a big-snout croaker, Johnius macrorhynus, produced under hand-held and voluntary conditions (in a large aquarium and in the field) were compared. Voluntary calls included "purr" and "dual-knocks", only purrs were produced when the fish was hand-held. The purr is composed of pulses in which the first interpulse interval was six to nine times longer than the other interpulse intervals, which were approximate in duration, and is a unique sound type in the coastal water of Taiwan. Purrs emitted under these conditions did not differ significantly, suggesting that the hand-held sound can be employed to match the sound in the field. These sounds contained energy reaching 5 kHz, with two peaks at about 1 and 2 kHz-the former being the dominant frequency. First interpulse interval, main interpulse interval, repetition rate of pulse, and pulse duration may serve as the diagnostic characters for the species-specific sound (i.e., purrs). One specimen survived in a large aquarium for 6 months and its vocal activity was monitored. The fish produced fewer dual-knocks than purrs, and purrs were about 11 dB louder than dual-knocks emitted in the aquarium. The temporal sequential relationship of these types in the sound of this individual was described.     

Doppler-shift compensation in the Taiwanese leaf-nosed bat (Hipposideros terasensis) recorded with a telemetry microphone system during flight

Biosonar behavior was examined in Taiwanese leaf-nosed bats (Hipposideros terasensis; CF-FM bats) during flight. Echolocation sounds were recorded using a telemetry microphone mounted on the bat's head. Flight speed and three-dimensional trajectory of the bat were reconstructed from images taken with a dual high-speed video camera system. Bats were observed to change the intensity and emission rate of pulses depending on the distance from the landing site. Frequencies of the dominant second harmonic constant frequency component (CF2) of calls estimated from the bats' flight speed agreed strongly with observed values. Taiwanese leaf-nosed bats changed CF2 frequencies depending on flight speed, which caused the CF2 frequencies of the Doppler-shifted echoes to remain constant. Pulse frequencies were also estimated using echoes returning directly ahead of the bat and from its sides for two different flight conditions: landing and U-turn. Bats in flight may periodically alter their attended angles from the front to the side when emitting echolocation pulses.     

Low frequency vocalizations attributed to sei whales (Balaenoptera borealis)

Low frequency (<100 Hz) downsweep vocalizations were repeatedly recorded from ocean gliders east of Cape Cod, MA in May 2005. To identify the species responsible for this call, arrays of acoustic recorders were deployed in this same area during 2006 and 2007. 70 h of collocated visual observations at the center of each array were used to compare the localized occurrence of this call to the occurrence of three baleen whale species: right, humpback, and sei whales. The low frequency call was significantly associated only with the occurrence of sei whales. On average, the call swept from 82 to 34 Hz over 1.4 s and was most often produced as a single call, although pairs and (more rarely) triplets were occasionally detected. Individual calls comprising the pairs were localized to within tens of meters of one another and were more similar to one another than to contemporaneous calls by other whales, suggesting that paired calls may be produced by the same animal. A synthetic kernel was developed to facilitate automatic detection of this call using spectrogram-correlation methods. The optimal kernel missed 14% of calls, and of all the calls that were automatically detected, 15% were false positives.     

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.     

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.     

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.