Acoustic characteristics and variations in grunt vocalizations in the oyster toadfish <Emphasis Type="Italic">Opsanus tau</Emphasis>

Acoustic communication is critical for reproductive success in the oyster toadfish Opsanus tau. While previous studies have examined the acoustic characteristics, behavioral context, geographical variation, and seasonality of advertisement boatwhistle sound production, there is limited information on the grunt or other non-advertisement vocalizations in this species. This study continuously monitored sound production in toadfish maintained in an outdoor habitat for four months to identify and characterize grunt vocalizations, compare them with boatwhistles, and test for relationships between the incidence of grunt vocalizations, sound characteristics and environmental parameters. Oyster toadfish produced grunts in response to handling, and spontaneous single (70% of all grunts), doublet (10%), and trains of grunts (20%) throughout the May to September study period. Grunt types varied in pulse structure, duration, and frequency components, and were shorter and of lower fundamental frequency than the pulse repetition rate of boatwhistles. Higher water temperatures were correlated with a greater number of grunt emissions, higher fundamental frequencies, and shorter sound durations. The number of grunts per day was also positively correlated with daylength and maximum tidal amplitude differences (previously entrained) associated with full and new moons, thus providing the first demonstration of semilunar vocalization rhythms in the oyster toadfish. These data provide new information on the acoustic repertoire and the environmental factors correlated with sound production in the toadfish, and have important implications for seasonal acoustic communication in this model vocal fish.     

The meaning and function of grunt variants in baboons

Wild baboons Papio cynocephalus ursinus, give tonal, harmonically rich vocalizations, termed grunts, in at least two distinct, behavioural contexts: when about to embark on a move across an open area ('move' grunts); and when approaching mothers and attempting to inspect or handle their young infants ('infant' grunts). Grunts in these two contexts elicit different responses from receivers and appear to be acoustically distinct (Owren et al. 1997 Journal of the Acoustical Society of America101 2951-2963). Differences in responses to grunts in the two contexts may, then, be due to acoustic differences, reflecting at least a rudimentary capacity for referential signalling. Alternatively, responses may differ simply due to differences in the contexts in which the grunts are being produced. We conducted playback experiments to test between these hypotheses. Experiments were designed to control systematically the effects of both context and acoustic features so as to evaluate the role of each in determining responses to grunts. In playback trials, subjects differentiated between putative move and infant grunts. Their responses based only on the acoustic features of grunts were functionally distinct and mirrored their behaviour to naturally occurring move and infant grunts. However, subjects' responses were in some cases also affected by the context in which grunts were presented, and by an interaction between the context and the acoustic features of the grunts. Furthermore, responses to grunts were affected by the relative rank difference between the caller and the subject. These results indicate that baboon grunts can function in rudimentary referential fashion, but that the context in which grunts are produced and the social identity of callers can also affect recipients' responses. Copyright 1999 The Association for the Study of Animal Behaviour.     

Acoustic communication in the Bocon toadfish (<Emphasis Type="Italic">Amphichthys cryptocentrus</Emphasis>)

The soundscapes of many coastal habitats include vocalizations produced by species of the family Batrachoididae (toadfish and midshipman). We describe the calling and grunting behavior of male Amphichthys cryptocentrus, a tropical toadfish, and predict how these vocalizations are influenced by conspecifics. We recorded individual males, which produced broadband grunts and multi-note, harmonic “boatwhistle” calls. Grunts were either in combination with calls or stand-alone. We used a null model to test if these latter grunts were produced at random or in response to calls from conspecifics. The model supports the hypothesis that grunts were in response to calls from neighboring males, suggesting acoustic competition. Using the most conservative estimate of hearing abilities we predict that males responded to the second harmonic of neighbor’s calls (230 Hz) at amplitudes of approximately 100–125 dB re 1μPa²/Hz. We also observed that call and grunt rates increased when males were exposed to higher rates of acoustic activity from neighboring fish. Fish used grunts to respond to background calls that occurred at different amplitudes, suggesting they responded to the calls of multiple neighboring fish and not just the highest amplitude neighbor. This communication with multiple fish within hearing range suggests a communication network in which the spatial distribution of individual toadfish relative to one another will impact their vocal behavior. Thus, the density and distribution, and not just abundance, of these toadfish at a given site will influence the characteristics of the chorus and the role of this species in the local soundscape.     

Vocal development in vervet monkeys

Observations and playback experiments were used to study the development of grunts and alarm calls among free-ranging vervet monkeys. Results indicate that the production of vocalizations, their use in appropriate circumstances, and the response to the vocalizations of others emerge gradually during an individual's first 4 years. Particularly in the case of grunts, different acoustic components develop at different rates. Immatures respond appropriately to the calls of others before they produce appropriate vocalizations themselves. Finally, immature vocal development may be aided by cues received from adults. If an infant gives an alarm call to a genuine predator (as opposed to a non-predator), adults respond more strongly. When infants hear a playback of an alarm, they are more likely to respond appropriately if they first look at an adult.     

Vocal Tract Morphology Determines Species-Specific Features in Vocal Signals of Lemurs (Eulemur)

The source-filter theory describes vocal production as a two-stage process involving the generation of a sound source, with its own spectral structure, which is then filtered by the resonant properties of the vocal tract. This theory has been successfully applied to the study of animal vocal signals since the 1990s. As an extension, models reproducing vocal tract resonance can be used to reproduce formant patterns and to understand the role of vocal tract filtering in nonhuman vocalizations. We studied three congeneric lemur species —Eulemur fulvus, E. macaco, E. rubriventer— using morphological measurements to build computational models of the vocal tract to estimate formants, and acoustic analysis to measure formants from natural calls. We focused on call types emitted through the nose, without apparent articulation. On the basis of anatomical measurements, we modeled the vocal tract of each species as a series of concatenated tubes, with a cross-sectional area that changed along the tract to approximate the morphology of the larynx, the nasopharyngeal cavity, the nasal chambers, and the nostrils. For each species, we calculated the resonance frequencies in 2500 randomly generated vocal tracts, in which we simulated intraspecific length and size variation. Formant location and spacing showed significant species-specific differences determined by the length of the vocal tract. We then measured formants of a set of nasal vocalizations (“grunts”) recorded from captive lemurs of the same species. We found species-specific differences in the natural calls. This is the first evidence that morphology of the vocal tract is relevant in generating filter-related acoustic cues that potentially provide receivers with information about the species of the emitter.     

How vervet monkeys perceive their grunts: Field playback experiments

Free-ranging vervet monkeys grunt to each other in a variety of social situations: when approaching a dominant or subordinate individual, when moving into a new area of their range, or when observing another group. Like other non-human primate vocalizations, these grunts have traditionally been interpreted as a single, highly variable call that reflects the arousal state of the signaller. Field playback experiments suggest, however, that what humans initially perceive as one grunt the monkeys perceive as at least four. Each grunt carries a specific meaning that seems to depend more on its acoustic properties than on the context in which it occurs. Results suggest that the vocalizations given by monkeys during social interactions may function in a rudimentary representational manner, as if to designate objects or events in the external world.     

Saccular potentials of the vocal plainfin midshipman fish, <Emphasis Type="Italic">Porichthys notatus</Emphasis>

The plainfin midshipman fish, Porichthys notatus, is a vocal species of teleost fish that generates acoustic signals for intraspecific communication during social and reproductive behaviors. All adult morphs (females and males) produce single short duration grunts important for agonistic encounters, but only nesting males produce trains of grunts and growls in agonistic contexts and long duration multiharmonic advertisement calls to attract gravid females for spawning. The midshipman fish uses the saccule as the main acoustic endorgan for hearing to detect and locate vocalizing conspecifics. Here, I examined the response properties of evoked potentials from the midshipman saccule to determine the frequency response and auditory threshold sensitivity of saccular hair cells to behaviorally-relevant single tone stimuli. Saccular potentials were recorded from the rostral, medial and caudal regions of the saccule while sound was presented by an underwater speaker. Saccular potentials of the midshipman, like other teleosts, were evoked greatest at a frequency that was twice the stimulus frequency. Results indicate that midshipman saccular hair cells of non-reproductive adults had a peak frequency sensitivity that ranged from 75 (lowest frequency tested) to 145 Hz and were best suited to detect the low frequency components (≤105 Hz) of midshipman vocalizations.     

Structural Classification of Wild Boar (Sus scrofa) Vocalizations

Determining whether a species' vocal communication system is graded or discrete requires definition of its vocal repertoire. In this context, research on domestic pig (Sus scrofa domesticus) vocalizations, for example, has led to significant advances in our understanding of communicative functions. Despite their close relation to domestic pigs, little is known about wild boar (Sus scrofa) vocalizations. The few existing studies, conducted in the 1970s, relied on visual inspections of spectrograms to quantify acoustic parameters and lacked statistical analysis. Here, we use objective signal processing techniques and advanced statistical approaches to classify 616 calls recorded from semi‐free ranging animals. Based on four spectral and temporal acoustic parameters—quartile Q25, duration, spectral flux, and spectral flatness—extracted from a multivariate analysis, we refine and extend the conclusions drawn from previous work and present a statistically validated classification of the wild boar vocal repertoire into four call types: grunts, grunt‐squeals, squeals, and trumpets. While the majority of calls could be sorted into these categories using objective criteria, we also found evidence supporting a graded interpretation of some wild boar vocalizations as acoustically continuous, with the extremes representing discrete call types. The use of objective criteria based on modern techniques and statistics in respect to acoustic continuity advances our understanding of vocal variation. Integrating our findings with recent studies on domestic pig vocal behavior and emotions, we emphasize the importance of grunt‐squeals for acoustic approaches to animal welfare and underline the need of further research investigating the role of domestication on animal vocal communication.     

Intended Receivers and Functional Significance of Grunt and Girney Vocalizations in Free-Ranging Female Rhesus Macaques

We investigated the intended receivers and contexts of occurrence of grunt and girney vocalizations in rhesus macaques (Macaca mulatta) to assess whether these calls are best interpreted as signals of benign intent or as calls that may function to attract the attention of other individuals or induce arousal. We focally observed 19 free‐ranging adult female rhesus macaques. Female calls increased dramatically after infants were born, and most were directed toward mother–infant dyads. When infants were physically separated from their mothers, callers visually oriented toward infants in over 90% of the cases, suggesting that infants were the intended receivers of grunts and girneys. Approaches followed by vocalizations were more likely to lead to the caller grooming the mother, less likely to elicit a submissive response, and more likely to result in infant handling than approaches without calls. Infant handling, however, was not necessarily benign. Vocalizations were often emitted from a distance >1 m and were rarely followed by approaches or social interactions. Our results suggest that grunts and girneys are unlikely to have evolved as signals that encode information about the caller's intention or subsequent behavior. Whereas girneys may be acoustically designed to attract infants’ attention and elicit arousal, grunts may have no adaptive communicative function. Mothers, however, may have learned that other females’ grunts and girneys are unlikely to be associated with significant risk and, therefore, are generally tolerant of the caller's proximity and behavior.     

Vocalizations and Behaviour of Pacific Humpback Dolphins Sousa chinensis

Very little is known about the acoustic repertoire of the Pacific humpback dolphin Sousa chinensis . This study, off eastern Australia, used concurrent observations of surface behaviour and acoustic recordings to gain an insight into the behavioural significance of humpback dolphin vocalizations. Humpback dolphins exhibit five different vocalization categories: broad band clicks; barks; quacks; grunts; and whistles. Broad band clicks were high in frequency (8 kHz to > 22 kHz), were directly related to foraging behaviour and may play a role in social behaviour. Barks and quacks were burst pulse sounds (frequency: 0.6 kHz to > 22 kHz, duration: 0.1–8 s) and were associated with both foraging and social behaviour. The grunt vocalization is a low frequency narrow band sound (frequency 0.5–2.6 kHz, duration 0.06–2 s) and was only heard during socializing. There were 17 different types of whistles, ranging widely in frequency (0.9–22 kHz) and vocal structure (n=329). The predominant whistle types used by the groups were type 1 (46%) and type 2 (17%). Most whistles were heard during both socializing and foraging. The number of whistles recorded in a group increased significantly as the number of mother–calf pairs increased, suggesting that whistles may be used as contact calls. Few vocalizations were heard during either travelling or milling behaviours. Broad band clicks, barks and whistle type 1 were the only vocalizations recorded during either travelling or milling.     

Individual recognition through olfactory–auditory matching in lemurs

Individual recognition can be facilitated by creating representations of familiar individuals, whereby information from signals in multiple sensory modalities become linked. Many vertebrate species use auditory–visual matching to recognize familiar conspecifics and heterospecifics, but we currently do not know whether representations of familiar individuals incorporate information from other modalities. Ring-tailed lemurs (Lemur catta) are highly visual, but also communicate via scents and vocalizations. To investigate the role of olfactory signals in multisensory recognition, we tested whether lemurs can recognize familiar individuals through matching scents and vocalizations. We presented lemurs with female scents that were paired with the contact call either of the female whose scent was presented or of another familiar female from the same social group. When the scent and the vocalization came from the same individual versus from different individuals, females showed greater interest in the scents, and males showed greater interest in both the scents and the vocalizations, suggesting that lemurs can recognize familiar females via olfactory–auditory matching. Because identity signals in lemur scents and vocalizations are produced by different effectors and often encountered at different times (uncoupled in space and time), this matching suggests lemurs form multisensory representations through a newly recognized sensory integration underlying individual recognition.     

The Perception of Sex-specificity in Long Calls of the Tamarin (Saguinus labiatus labiatus)

Long calls given by red-chested moustached tamarins (Saguinus l. labiatus), typically 1–2 s in duration and made up of individual syllables, reveal distinctive sex-specificity as well as population and individual differences in their acoustic structure. Distributions of male calls and female vocalizations are discrete with regard to two acoustic parameters, i.e. number of syllables and mean inter-syllable intervals. Mean number of syllables produced in a given time is much greater in male than in female calls. I played back synthetic versions of the long calls varying in mean inter-syllable intervals to captive tamarins and found that the sex differences were encoded in a relatively simple acoustic form. The tamarin is able to identify the sex of unfamiliar calling conspecifics by voice alone.     

Interspecific Responses of Ringtailed Lemurs to Playback of Antipredator Alarm Calls Given by Verreaux's Sifakas

The ringtailed lemur, Lemur catta, and Verreaux's sifaka, Propithecus verreauxi verreauxi, are diurnal prosimians living sympatrically in Madagascar. Species-specific alarm calls emitted by each of these two species in response to aerial and terrestrial predators differ acoustically. Behavioural responses of ringtailed lemurs evoked by playbacks of conspecific alarm calls differ when the vocalizations were produced in response to aerial predators as opposed to terrestrial predators. We conducted playback experiments on two populations of ringtailed lemurs, using two types of sifaka alarm calls. One population consisted of free-ranging groups which lived sympatrically with sifakas, the other was a colony group which had no contact with sifakas. The results illustrate that the former group of lemurs can perceive what type of predators the sifaka calls refer to, whereas the latter group was not able to recognize the difference in the calls.     

Spectral call features provide information about the aggression level of greater mouse-eared bats (Myotis myotis) during agonistic interactions

Bats rely heavily on acoustic signals in order to communicate with each other in a variety of social contexts. Among those, agonistic interactions and accompanying vocalizations have received comparatively little study. Here, we studied the communicational behaviour between male greater mouse-eared bats (Myotis myotis) during agonistic encounters. Two randomly paired adult males were placed in a box that allowed us to record video and sound synchronously. We describe their vocal repertoire and compare the acoustic structure of vocalizations between two aggression levels, which we quantified via the bats’ behaviour. By inspecting thirty, one-minute long encounters, we identified a rich variety of social calls that can be described as two basic call types: echolocation-like, low-frequency sweeps and long, broadband squawks. Squawks, the most common vocalization, were often noisy, i.e. exhibited a chaotic spectral structure. We further provide evidence for individual signatures and the presence of nonlinear phenomena in this species’ vocal repertoire. As the usage and acoustic structure of vocalizations is known to encode the internal state of the caller, we had predicted that the spectral structure of squawks would be affected by the caller’s aggression level. Confirming our hypothesis, we found that increased aggression positively correlated with an increase in call frequency and tonality. We hypothesize that the extreme spectral variability between and within squawks can be explained by small fluctuations in vocal control parameters (e.g. subglottal pressure) that are caused by the elevated arousal, which is in turn influenced by the aggression level.     

The influence of age and sex on the vocal repertoire of the Antillean manatee (Trichechus manatus manatus) and their responses to call playback

Limited information is available regarding the acoustic communication of Antillean manatees, however, studies have shown that other manatee taxa produce vocalizations as a method of individual recognition and communication. Here, the acoustic signals of 15 Antillean manatees in captivity were recorded, aiming to (1) describe their acoustic repertoire, (2) investigate the influence of sex and age on vocalization, and (3) examine manatee responses to call playback. Six acoustic signals ranging in mean fundamental frequencies from 0.64 kHz to 5.23 kHz were identified: squeaks and screeches were common to adult males, adult females, and juveniles; trills were common to adult males and females; whines were specific to males; creaks were specific to females; and rubbing was specific to juveniles. The structure of squeak vocalizations was significantly different between age and sex classes and screech structure was significantly different between age classes. Squeaks and screeches produced by juveniles had higher frequencies of maximum energy when compared to those produced by adult males and females. A significant increase in the vocalization rate following vocalization playbacks was found for all three age/sex groups. Our results introduce the potential of using acoustic signals in identifying and noninvasively monitoring manatees in the wild in Brazil.     

Vocal divergence between two disjunct populations of Giant Antshrike (Batara cinerea) is related to environmental conditions

Variation in the avian vocal signals emitted may have a significant impact on species evolution. Vocal divergence in suboscine species like Giant Antshrike (Batara cinerea) may be associated with selective adaptation, since learning has little influence on vocal development and variation in acoustic structure cannot be attributed to learning deviation. Consequently, tracheophone suboscine species are ideal subjects to explore vocal variation, since cultural evolution does not seem to influence vocal variation in this group. Environmental conditions may determine the selection of vocal features because acoustic transmission could be attenuated under certain conditions of temperature, humidity and vegetation cover. Here, we examined vocalizations of Giant Antshrike and assessed possible acoustic variations between two disjunct groups (Andean and Atlantic), correlating the differences to the environmental structure. Univariate and multivariate analysis show temporal and spectral differences between both groups. Andean individuals produce vocalizations with longer duration, faster trill rates, shorter syllable duration and higher frequencies. Environmental features are different between the two populations, and they are correlated to the acoustic structure of vocalizations. Temporal variations arise directly from climatic influence, while spectral divergence could be a secondary effect of morphological adaptation to habitat structure.     

A comparison of individual distinctiveness in three vocalizations of the dwarf mongoose (Helogale parvula)

Individual specificity can be found in the vocalizations of many avian and mammalian species. However, it is often difficult to determine whether these vocal cues to identity rise from “unselected” individual differences in vocal morphology or whether they have been accentuated by selection for the purposes of advertising caller identity. By comparing the level of acoustic individuality of different vocalizations within the repertoire of a single species, it is possible to ascertain whether selection for individual recognition has modified the vocal cues to identity in particular contexts. We used discriminant function analyses to determine the level of accuracy with which calls could be classified to the correct individual caller, for three dwarf mongoose (Helogale parvula) vocalizations: contact, snake, and isolation calls. These calls were similar in acoustic structure but divergent in context and function. We found that all three call types showed individual specificity but levels varied with call type (increasing from snake to contact to isolation call). The individual distinctiveness of each call type appeared to be directly related to the degree of benefit that signalers were likely to accrue from advertising their identity within that call context. We conclude that dwarf mongoose signalers have undergone selection to facilitate vocal individual recognition, particularly in relation to the species’ isolation call.     

Individual Acoustic Variation in Fallow Deer (Dama dama) Common and Harsh Groans: A Source-Filter Theory Perspective

Mammals are able to distinguish conspecifics based on vocal cues, and the acoustic structure of mammal vocalizations is directly affected by the anatomy and action of the vocal apparatus. However, most studies investigating individual patterns in acoustic signals do not consider a vocal production‐based perspective. In this study, we used the source‐filter model of vocal production as a basis for investigating the acoustic variability of fallow deer groans. Using this approach, we quantified the potential of each acoustic component to carry information about individual identity. We also investigated if cues to individual identity carry over among the two groan types we describe: common and harsh groans. Using discriminant function analysis, we found that variables related to the fundamental frequency contour and the minimum frequencies of the highest formants contributed most to the identification of a given common groan. Common groans were individually distinctive with 36.6% (53.6% with stepwise procedure) of groans assigned to the correct individual. This level of discrimination is approximately six times higher than that predicted by chance. In addition, univariate anova s showed significant inter‐individual variation in the minimum formant frequencies when common and harsh groans were combined, suggesting that some information about individuality is shared between groan types. Our results suggest that the sound source and the vocal tract resonances act together to determine groan individuality and that enough variation exists to potentially allow individual recognition based on groans.     

The behavioral repertoire of the black-and-white ruffed lemur, Varecia variegata variegata (Primates: Lemuridae)

A stable social group of 7 semifree-ranging black-and-white ruffed lemurs (Varecia variegata variegata) was studied for 4 months to catalog the behavioral repertoire of this species. Observations focussed on particular aspects of behavior were conducted before and after this 4-month period to supplement information gathered. Behavior in 11 major categories is detailed: postures, terrestrial locomotion, arboreal locomotion, feeding behavior, vocalizations, scent-marking, affinitive social behavior, agonistic social behavior, play behavior, sexual behavior, and parental behavior. Ruffed lemurs frequently used body positions and locomotor patterns unusual among lemurids, including bipedal hanging and long-descent leaps. These behaviors reinforce dental evidence that Varecia are among the most frugivorous of the Malagasy lemurs. Low intragroup cohesion, infrequent social interaction, and antiphonal use of several long-distance vocalizations suggest that ruffed lemurs naturally exhibit fission-fusion sociality. Social structure based on interindividual familiarity probably extends across foraging parties for several of the diurnally active lemurs; however, thus far only Varecia seems likely to exhibit fission-fusion sociality analogous to that seen in spider monkeys and chimpanzees.     

Use of Vocal Fingerprinting for Specific Discrimination of Gray (Microcebus murinus) and Rufous Mouse Lemurs (Microcebus rufus)

Advertisement calls are often important noninvasive tools for discriminating cryptic species and for assessing specific diversity and speciation patterns in nature. We investigated the contribution of these calls to uncover specific diversity in nocturnal Malagasy lemurs. We compared sexual advertisement and predator advertisement calls of two mouse lemur species, western gray and eastern rufous mouse lemurs (Microcebus murinus and M. rufus, respectively) living in two contrasting habitats (dry deciduous vs. rain forest), and analyzed them statistically. Both species emitted several highly variable whistle calls in the context of predator-avoidance. Intrapopulation variation was high and overlapped interspecific variation. Sexual advertisement calls, given in the mating context, displayed a totally distinct, species-specific acoustic structure. Whereas gray mouse lemurs produced rapidly multifrequency modulated, long trill calls, rufous mouse lemurs gave slowly frequency-modulated short chirp calls. Our results suggest specific status for gray and rufous mouse lemurs and indicate the importance of predation and social needs in shaping vocal communication.