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.     

Vocalisations of the adult female domestic pig during a standard human approach test and their relationships with behavioural and heart rate measures

Vocal communication in the domestic pig is generally not well documented. The aim of this experiment was to categorise and ascribe the function of the vocalisations of 67 Large WhitexLandrace gilts during a standard human approach test. At testing, each group of 3-5 gilts was moved to a handling area where each individual in turn was fitted with a heart rate monitor and introduced individually to a 2.4mx2.4m test arena. After 2min familiarisation, an unfamiliar human entered the pen and stood for 3min against one wall. Behaviour and sound were recorded continuously with sound recordings transferred onto computer for analysis. Three categories of calls were initially identified: single grunts, single squeals and rapidly repeated grunts. Sixty-six gilts performed single grunts, whereas only 28 and 16 gilts performed the other two categories, respectively. Single grunts could be sub-divided into two types based on sound amplitude profile. These types differed significantly in duration. Gilts performed more short and long grunts per minute during the 3min test period than during the familiarisation period. Most short grunts observed in a subset of 15 gilts were performed with the snout close to a pen surface or the human. The rate of short grunts during the test period was negatively correlated with the time taken to make contact with the human and positively correlated with the amount of locomotor behaviour carried out, the total number of interactions with the human and the total time spent within 0.5m of the human. Most long grunts observed in a subset of 15 gilts were performed with the snout away from any surface. The rate of long grunts during the test period positively correlated with amount of locomotor behaviour and heart rate, after the effect of activity had been removed. Squeals could similarly be sub-divided into long and short types on the basis of amplitude profile. Gilts that squealed carried out more locomotor behaviour, interacted with the human more, had higher mean heart rates and lower heart rate rise when touched by the human, suggesting a higher degree of arousal. Rapidly-repeated grunts were associated with close human interaction. The results indicate that the domestic pig performs a number of distinct vocalisations during isolation. Short single grunts appear to be associated with investigatory behaviour. Long single grunts may be a form of contact call, the rate of which is related to physiological and behavioural activity. Squeals may have similar function but result from a higher level of arousal. Short, rapidly-repeated grunts appear to have either a greeting or threat function. With further research, certain pig vocalisations may be identified as providing useful additional information about an individual's welfare.     

Selection for acoustic individuality within the vocal repertoire of wild chimpanzees

Individual primates typically produce acoustically distinct calls. To investigate the factors that facilitate the evolution of individual vocal signatures, we examined two components of the call repertoire of chimpanzees: the pant hoot and pant grunt. Pant hoots are long-distance signals whose recipients can be several hundred meters away, while pant grunts are short-range calls given to conspecifics within close visual range. Given their markedly different contexts of emission, we predicted that natural selection would favor the elaboration of individually distinctive acoustic features in pant hoots compared with pant grunts. Analyses of nine acoustic features revealed that pant hoots are more stereotyped within-individuals and variable between-individuals than pant grunts. These data are consistent with the hypothesis that selection may act to encode varying degrees of individuality in different components of the vocal repertoire of a single species.     

Expression of Emotional Arousal in Two Different Piglet Call Types

Humans as well as many animal species reveal their emotional state in their voice. Vocal features show strikingly similar correlation patterns with emotional states across mammalian species, suggesting that the vocal expression of emotion follows highly conserved signalling rules. To fully understand the principles of emotional signalling in mammals it is, however, necessary to also account for any inconsistencies in the way that they are acoustically encoded. Here we investigate whether the expression of emotions differs between call types produced by the same species. We compare the acoustic structure of two common piglet calls—the scream (a distress call) and the grunt (a contact call)—across three levels of arousal in a negative situation. We find that while the central frequency of calls increases with arousal in both call types, the amplitude and tonal quality (harmonic-to-noise ratio) show contrasting patterns: as arousal increased, the intensity also increased in screams, but not in grunts, while the harmonicity increased in screams but decreased in grunts. Our results suggest that the expression of arousal depends on the function and acoustic specificity of the call type. The fact that more vocal features varied with arousal in scream calls than in grunts is consistent with the idea that distress calls have evolved to convey information about emotional arousal.     

Playbacks of food-associated calls attract chimpanzees towards known food patches in a captive setting

Food-associated calls have received much research attention due to their potential to refer to discovered food in a word-like manner. Studies have found that in many species, food-associated calls attract receivers to the food patch, suggesting these calls play roles in food sharing, cooperation and competition. Additionally, in various species, these calls play a role that has received much less attention: mediating social interactions among foragers that are already nearby or within the food patch, independently of whether they attract outside foragers. In order to increase understanding of the function of the chimpanzee (Pan troglodytes) food-associated rough grunt, we conducted captive playback studies testing whether rough grunt playbacks attract, repel or have no effect on the proximity of foragers already familiarized with the presence of food. We tested how acoustic playbacks of rough grunts (or control calls) from one of two known, identical feeding sites affected receivers’ approach and feeding behaviors. More often than expected, participants first approached the feeding site from which rough grunts, but not control calls, were broadcast. However, neither condition increased the likelihood that participants fed first from a given site. Our results support the hypothesis that rough grunts elicit an approach response in receivers, while providing no evidence that they repel. In addition, our study provides evidence that receivers may approach rough grunts even if they do not intend to feed. We discuss the information rough grunts may convey to receivers beyond information about discovered food and the potential benefits signalers may gain from this calling behavior.

     

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.     

Contact calling behaviour in the male ring‐tailed lemur (Lemur catta)

The ring‐tailed lemur (Lemur catta) is a group‐living strepsirhine primate endemic to Madagascar that has a complex vocal repertoire including affiliative moan and hmm calls. Past research has suggested that both vocalisations may be contact calls, and we assessed their usage in addition to examining the effect of individual differences and social factors on male calling behaviour. We tested three hypotheses: the group cohesion, preferred companion and individual differences hypotheses. From March to July 2010, 565 h of focal animal data were collected on 31 males aged ≥1 year at the Beza Mahafaly Special Reserve, Madagascar. Focal males were observed for 30 min, with vocalisations, allogrooming and agonistic interactions noted using one‐zero sampling, and general behaviour, nearest neighbour identity and distance noted using instantaneous point sampling, all at 2.5 min intervals. We found support for the group cohesion hypothesis for both vocalisations, and support for the preferred companion and individual differences hypotheses for the hmm but not the moan vocalisation. Moan and hmm calls maintained the distance between a male and his nearest neighbour, and were made at higher rates during behaviours which may lead to separation from conspecifics, such as travel and vigilance. Males made hmm calls at higher rates when they were low‐ranking, and/or when the nearest neighbour of a male or a preferred companion of either sex. Low‐ranking males may hmm call to maintain proximity to specific individuals within social groups who will show them increased tolerance, with the hmm signalling peaceful intent. Ring‐tailed lemurs are one of the best living models of gregarious primate ancestors, and gaining a better knowledge of the function and range of usage of their contact calls informs our understanding of the evolution of primate cognition.     

Temporal variation and characterization of grunt sounds produced by Atlantic cod Gadus morhua and pollack Pollachius pollachius during the spawning season

Fine‐scale temporal patterning in grunt production and variation in grunt attributes in Atlantic cod Gadus morhua and pollack Pollachius pollachius was examined. Pollachius pollachius produced only a single sound type, the grunt, similar to that previously described for G. morhua. Sound production and egg production were correlated in P. pollachius but not in G. morhua. Only G. morhua displayed a strongly cyclical pattern, producing more grunts at night. Finer‐scale temporal patterning in grunt production was observed in both species which produced significantly fewer grunts following a period of high grunt production. These quieter periods lasted up to 45 min for P. pollachius and up to 1 h in G. morhua. Grunts were not always produced in isolation but organized into bouts in both species. Longer bouts were more frequent during periods of increased sound activity and were linked with changes in grunt characteristics including increased grunt duration, pulse duration and repetition period of each pulse combined with decreased dominant frequency. This study provides the first evidence of acoustic signalling being used by spawning P. pollachius and presents the most detailed analysis of the complexity of gadoid sound production.     

Crepuscular Changes in Emission Rate and Parameters of the Boatwhistle Advertisement Call of the Gulf Toadfish, Opsanus Beta

We quantified crepuscular variation in the emission rate and call properties of the boatwhistle advertisement call of Gulf toadfish, Opsanus beta, from a field recording of a natural population of nesting males in the Florida Keys. Their calls are more variable and complex than previously reported. A call typically starts with a grunt followed by one to five tonal boop notes (typically two or three) and lasts for over a second. The first boop is considerably longer than later ones, and intervals between boops are relatively constant until the final interval, which approximately doubles in duration. Positions of fish are fixed and calls are sufficiently variable that we could discern individual callers in field recordings. Calling rate increases after sunset when males tend to produce shorter calls with fewer notes. Analysis by number of notes per call indicates some individuals decrease the number of initial grunts and the duration of the first note, but most of the decrease results from fewer notes. To our knowledge this sort of call plasticity has not been demonstrated before in fishes. We suggest that call shortening lowers the chances of overlapping calls of other males and that the small amount of time actually spent producing sound (total on time) is an adaptation to prevent fatigue in sonic muscles adapted for speed but not endurance.     

Collective decision-making in white-faced capuchin monkeys

In group-living animals, collective movements are a widespread phenomenon and occur through consensus decision. When one animal proposes a direction for group movement, the others decide to follow or not and hence take part in the decision-making process. This paper examines the temporal spread of individual responses after the departure of a first individual (the initiator) in a semi-free ranging group of white-faced capuchins (Cebus capucinus). We analysed 294 start attempts, 111 succeeding and 183 failing. Using a modelling approach, we have demonstrated that consensus decision-making for group movements is based on two complementary phenomena in this species: firstly, the joining together of group members thanks to a mimetic process; and secondly, a modulation of this phenomenon through the propensity of the initiator to give up (i.e. cancellation rate). This cancellation rate seems to be directly dependent upon the number of followers: the greater this number is, the lower the cancellation rate is seen to be. The coupling between joining and cancellation rates leads to a quorum: when three individuals join the initiator, the group collectively moves. If the initiator abandons the movement, this influences the joining behaviour of the other group members, which in return influences the initiator''s behaviour. This study demonstrates the synergy between the initiator''s behaviour and the self-organized mechanisms underlying group movements.     

Wild Female Olive Baboons Adapt their Grunt Vocalizations to Environmental Conditions

Sound propagates differently and visibility varies according to the habitat type. Animals should therefore adapt the acoustic structure and the usage of their vocal signals to the environment. In the present study, we examined the influence of the habitat on the vocal behaviour of wild olive baboons ( Papio hamadryas anubis ) in two populations: one living in Gashaka-Gumti National Park, Nigeria, and the other in Budongo Forest, Uganda. We investigated whether female baboons modified the acoustic structure of their grunts and their rate of grunting when they wandered between closed and open habitat types. As an adaptation to the environmental conditions, baboons might utter calls with a longer duration, a lower fundamental frequency and/or energy concentrated in lower frequencies in a closed habitat like forest than in an open habitat. Baboons should also grunt more frequently in the closed habitat. Analyses showed that in both populations grunts uttered in forest were significantly longer than in open habitat. Additionally, baboons from Uganda showed a significantly higher grunt rate in forest than in open habitat. These results revealed a certain degree of plasticity in vocal production and call usage with regard to the habitat type. However, results in Nigeria suggested that, besides habitat structure, other proximate factors like the context of calling and the proximity between group members could also have an influence on the actual communication patterns.     

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.     

Spawning sounds in meagre Argyrosomus regius recorded in the Gironde estuary, France

During their spawning period (June to mid-July) in the Gironde estuary, meagre Argyrosomus regius produce two distinct sounds: regular long grunts, the most common calls, and sometimes also short grunts. It is suggested that long grunts serve the formation of spawning aggregations and short grunts announce the beginning of courtship behaviour. The meagre's long grunts include long series of 30–112 closely spaced pulses placed into call units. Each pulse produces multiple and rapidly decaying swimbladder vibrations with a dominant frequency varying between 336 and 444 Hz.     

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.     

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.     

Ready to Fight: Reliable Predictors of Attack in a Cooperatively Breeding,Non‐Passerine Bird

Signal reliability is a major focus of animal communication research. Aggressive signals are ideal for measuring signal reliability because the signal referent – attack or no attack – can be measured unambiguously. Signals of aggressive intent occur at elevated rates in aggressive contexts, predict subsequent aggression by the signaler, and elicit appropriate responses from receivers. We tested the ‘predictive criterion’ in smooth‐billed anis, Crotophaga ani, by broadcasting one of two playback types (‘ahnee’ calls only or ‘ahnee + hoot’ calls), presenting a taxidermic mount, and observing the animals’ behavior. Based on the hypotheses that ‘hoot’ calls and ‘throat‐inflation’ displays signal aggressive intent, we predicted that they would be associated with attack, and that signaling rate would increase over the time period leading up to an attack. Indeed, both hoots and throat‐inflation displays reliably predicted attack. The second prediction, that signaling rate increases in the time leading up to attack, was strongly supported for throat‐inflation displays, which increased over the pre‐attack period in both treatments. Hoots increased over the pre‐attack period in ahnee playbacks but not in ahnee + hoot playbacks. Hierarchical signaling systems are characterized by early, less‐reliable predictors of attack, and later, more reliable predictors of attack. During both natural and simulated interactions, the more‐reliable throat‐inflation display tended to precede the less‐reliable hoot call, suggesting that this signaling system is not hierarchical. In a comparison of 11 putative signals of aggressive intent in birds, the throat‐inflation display had the second highest mutual information (reduction in uncertainty) among visual signals and non‐passerine signals while hoots had below‐average mutual information. Natural observations indicate that both hoots and throat‐inflation displays occur in the context of aggressive between‐group encounters, and hoots also occur during within‐group interactions. Throat‐inflation displays appear to be reliable indicators of aggressive intent, but the function of hoot calls is less clear.     

Calling by Concluding Sentinels: Coordinating Cooperation or Revealing Risk?

Efficient cooperation requires effective coordination of individual contributions to the cooperative behaviour. Most social birds and mammals involved in cooperation produce a range of vocalisations, which may be important in regulating both individual contributions and the combined group effort. Here we investigate the role of a specific call in regulating cooperative sentinel behaviour in pied babblers (Turdoides bicolor). 'Fast-rate chuck' calls are often given by sentinels as they finish guard bouts and may potentially coordinate the rotation of individuals as sentinels, minimising time without a sentinel, or may signal the presence or absence of predators, regulating the onset of the subsequent sentinel bout. We ask (i) when fast-rate chuck calls are given and (ii) what effect they have on the interval between sentinel bouts. Contrary to expectation, we find little evidence that these calls are involved in regulating the pied babbler sentinel system: observations revealed that their utterance is influenced only marginally by wind conditions and not at all by habitat, while observations and experimental playback showed that the giving of these calls has no effect on inter-bout interval. We conclude that pied babblers do not seem to call at the end of a sentinel bout to maximise the efficiency of this cooperative act, but may use vocalisations at this stage to influence more individually driven behaviours.     

Functionally referential communication in a chimpanzee

The evolutionary origins of the use of speech signals to refer to events or objects in the world have remained obscure. Although functionally referential calls have been described in some monkey species, studies with our closest living relatives, the great apes, have not generated comparable findings. These negative results have been taken to suggest that ape vocalizations are not the product of their otherwise sophisticated mentality and that ape gestural communication is more informative for theories of language evolution. We tested whether chimpanzee rough grunts, which are produced during feeding contexts, functioned as referential signals. Individuals produced acoustically distinct types of "rough grunts" when encountering different foods. In a naturalistic playback experiment, a focal subject was able to use the information conveyed by these calls produced by several group mates to guide his search for food, demonstrating that the different grunt types were meaningful to him. This study provides experimental evidence that our closest living relatives can produce and understand functionally referential calls as part of their natural communication. We suggest that these findings give support to the vocal rather than gestural theories of language evolution.     

The Organization of Collective Group Movements in Wild Barbary Macaques (Macaca sylvanus): Social Structure Drives Processes of Group Coordination in Macaques

Social animals have to coordinate activities and collective movements to benefit from the advantages of group living. Animals in large groups maintain cohesion by self-organization processes whereas in smaller groups consensus decisions can be reached. Where consensus decisions are relevant leadership may emerge. Variation in the organization of collective movements has been linked to variation in female social tolerance among macaque species ranging from despotic to egalitarian. Here we investigated the processes underlying group movements in a wild macaque species characterized by a degree of social tolerance intermediate to previously studied congeneric species. We focused on processes before, during and after the departure of the first individual. To this end, we observed one group of wild Barbary macaques (Macaca sylvanus) in the Middle Atlas, Morocco using all-occurrence behaviour sampling of 199 collective movements. We found that initiators of a collective movement usually chose the direction in which more individuals displayed pre-departure behavior. Dominant individuals contributed to group movements more than subordinates, especially juveniles, measured as frequencies of successful initiations and pre-departure behaviour. Joining was determined by affiliative relationships and the number of individuals that already joined the movement (mimetism). Thus, in our study group partially shared consensus decisions mediated by selective mimetism seemed to be prevalent, overall supporting the suggestion that a species’ social style affects the organization of group movements. As only the most tolerant species show equally shared consensus decisions whereas in others the decision is partially shared with a bias to dominant individuals the type of consensus decisions seems to follow a stepwise relation. Joining order may also follow a stepwise, however opposite, relationship, because dominance only determined joining in highly despotic, but not in intermediate and tolerant species.     

Vocalisations of wild common marmosets are influenced by diurnal and ontogenetic factors

The vocalisations of wild common marmosets, Callithrix jacchus, were recorded to investigate whether call rate by individuals is affected by time of day, age, sex or dominance rank within a group. We also investigated how vocalisation pitch was affected by age, focussing on a single common call, the trill call. Adults vocalised more than juveniles or infants during the majority of daylight hours. Only the call rate of juveniles varied significantly over the day. No differences were found between either sex or dominance rank with respect to rate of vocalisations. The trill calls emitted by young wild common marmosets were of a higher pitch than those emitted by adults. We conclude that the auditory communication of wild common marmosets is related to the age of the animals, both in terms of call rate and the physical characteristics of their vocalisations.