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.     

Geographical variation in the vocalizations of the suboscine Thorn‐tailed Rayadito Aphrastura spinicauda

Structural variation in acoustic signals may be related either to the factors affecting sound production such as bird morphology, or to vocal adaptations to improve sound transmission in different environments. Thus, variation in acoustic signals can influence intraspecific communication processes. This will ultimately influence divergence in allopatric populations. The study of geographical variation in vocalizations of suboscines provides an opportunity to compare acoustic signals from different populations, without additional biases caused by song learning and cultural evolution typical of oscines. The aim of this study was to compare vocalizations of distinct populations of a suboscine species, the Thorn‐tailed Rayadito. Four types of vocalizations were recorded in five populations, including all three currently accepted subspecies. Comparisons of each type of vocalization among the five populations showed that some variation existed in the repetitive trill, whereas no differences were found among alarm calls and loud trills. Variation in repetitive trills among populations and forest types suggests that sound transmission is involved in vocal differences in suboscines. Acoustic differences are also consistent with distinguishing subspecies bullocki from spinicauda and fulva, but not the two latter subspecies from each other. Our results suggest that the geographical differentiation in vocalizations observed among Thorn‐tailed Rayadito populations is likely to be a consequence of different ecological pressures. Therefore, incipient genetic isolation of these populations is suggested, based on the innate origin of suboscine vocalizations.     

Vocal communication of wild bonnet macaques (Macaca radiata)

Field observations and spectrographic analyses of sound recordings of South Indian bonnet macaques revealed a vocal repertoire of at least 25 basic patterns. The repertoire consists of well separated sound classes and acoustic categories connected by structural intergradation. Besides structural variations within and between different elements of the repertoire, the vocal system ofMacaca radiata is characterized by regular combinations of particular basic patterns. These combinations occurred not only between calls of similar structure and function but also between calls usually emitted in entirely different social contexts. According to the qualitative analysis, sex-specific asymmetries of the vocal behaviour were less pronounced than age-dependent characteristics. The comparison of clear call vocalizations ofMacaca radiata andM. fuscata revealed significant species-specific differences on the structural and the behavioural level. Evaluations of the structural features of alarm calls of various macaque species imply marked differences between members of thefascicularis group andsinica group on one hand and thesilenus group andarctoides group on the other.     

Visualizing Sound Emission of Elephant Vocalizations: Evidence for Two Rumble Production Types

Recent comparative data reveal that formant frequencies are cues to body size in animals, due to a close relationship between formant frequency spacing, vocal tract length and overall body size. Accordingly, intriguing morphological adaptations to elongate the vocal tract in order to lower formants occur in several species, with the size exaggeration hypothesis being proposed to justify most of these observations. While the elephant trunk is strongly implicated to account for the low formants of elephant rumbles, it is unknown whether elephants emit these vocalizations exclusively through the trunk, or whether the mouth is also involved in rumble production. In this study we used a sound visualization method (an acoustic camera) to record rumbles of five captive African elephants during spatial separation and subsequent bonding situations. Our results showed that the female elephants in our analysis produced two distinct types of rumble vocalizations based on vocal path differences: a nasally- and an orally-emitted rumble. Interestingly, nasal rumbles predominated during contact calling, whereas oral rumbles were mainly produced in bonding situations. In addition, nasal and oral rumbles varied considerably in their acoustic structure. In particular, the values of the first two formants reflected the estimated lengths of the vocal paths, corresponding to a vocal tract length of around 2 meters for nasal, and around 0.7 meters for oral rumbles. These results suggest that African elephants may be switching vocal paths to actively vary vocal tract length (with considerable variation in formants) according to context, and call for further research investigating the function of formant modulation in elephant vocalizations. Furthermore, by confirming the use of the elephant trunk in long distance rumble production, our findings provide an explanation for the extremely low formants in these calls, and may also indicate that formant lowering functions to increase call propagation distances in this species''.     

Individual and geographical variation in display behaviour of male harbour seals in Scotland

Studying variations in behaviour at the individual or population level enables insight into the reproductive strategies within a species. We examined individual and geographical variation in the vocal and dive behaviour of male harbour seals, Phoca vitulina, which is associated with aquatic mating. This display behaviour was recorded in the Moray Firth, Scotland, from July 1994 to 1997, and in Orkney, Scotland, during July 1998. One vocalization type was apparent in the Moray Firth and two in Orkney. Time parameters (total and pulse duration) varied between males in the population in the Moray Firth. We used both frequency and time parameters in a discriminant analysis, which showed that 73.2% of individual male vocalizations could be correctly classified; 94.6% of male vocalizations from the Moray Firth and Orkney could be correctly classified according to their geographical areas. Therefore, vocal variation was greater between geographical areas than between individuals. No individual variation was apparent between dive and surface interval durations. However, individuals varied significantly in the percentage of short surface intervals. Male harbour seals showed substantial variability in the parameters affecting their vocal and dive behaviour during the mating season. We suggest that these variations may be indicative of adaptations to varying environmental challenges influencing the reproductive strategies of discrete populations. Copyright 2000 The Association for the Study of Animal Behaviour.     

Dialects in pygmy marmosets? Population variation in call structure

Population variation in primate vocal structure has been rarely observed. Here, we report significant population differences in the structure of two vocalizations in wild pygmy marmosets (Trills and J calls). We studied 14 groups of pygmy marmosets Callithrix (Cebuella) pygmaea pygmaea from five populations in northeastern Ecuador. We analyzed the acoustic structure of Trills and J calls recorded from two adult animals in each group through focal samples. Although individuals and groups within a population differed in call structure, we found consistent structural differences at a population level for Trills and J calls. Pair‐wise comparisons for the two call types point to San Pablo and Amazoonico as the populations that differed the most, whereas Hormiga and Zancudococha showed the least differences. Discriminant function analysis indicates that calls from each population could be classified accurately at rates significantly above chance. Habitat acoustics, social factors and genetic drift may explain interpopulation vocal differences. This is the first evidence of within‐subspecies vocal differences, or dialects, in wild populations of a neotropical primate species. Am. J. Primatol. 71:333–342, 2009. © 2009 Wiley‐Liss, Inc.     

Dialects in Japanese Monkeys: Vocal Learning and Cultural Transmission of Locale-specific Vocal Behavior?

Differences were detected by ear in vocalizations made during artificial feeding of Japanese monkey troops at three locations. Tape recording and sound spectrographic analysis confirmed a distinctive vocal pattern specific to each site and used only in the provisioning situation. The 3 different acoustic morphologies are variations on a shared tonal theme. Vocal learning by Macaca fuscata may have occurred separately at each site regulated by species-wide constraints on vocal production.     

Sound production in <Emphasis Type="Italic">Etheostoma oophylax</Emphasis> (Percidae) and call characteristics correlated to body size

Sound production in fishes is common in marine and freshwater species, however there are still many vocal species for which sound production has not been documented. This paper is the first account of sound production in the Guardian Darter (Etheostoma oophylax). Laboratory recordings revealed that males produced several vocalizations, including single pulse knocks, multi-pulsed purrs, and tonal drums. All vocalizations were documented during agonistic and courtship encounters, including spawning. We also investigated possible correlations between call characteristics and male size. Male standard length was found to be correlated to inter-pulse interval of purrs, as well as the slope of the drum vocalizations. Determining a link between male size and acoustic characteristics could be the first step in documenting mate or male-male assessment by acoustic communication in darters.     

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.     

Age- and Sex-Related Variations in Clear Calls of <Emphasis Type="Italic">Papio ursinus</Emphasis>

Changes in vocalizations during ontogeny can in principle be related to three factors: growth, maturation, and experience, i.e., learning. While learning is a prerequisite for the proper development of speech, it hardly appears to play a role in the development of the species-typical vocal behavior of nonhuman primates. Nonetheless, subjects of different age and sex often exhibit prominent variation in the structure of their vocalizations. We investigated ontogenetic changes and the emergence of sex-related differences in the acoustic structure of Chacma baboon (Papio ursinus) clear calls. We recorded the vocalizations emitted by individuals separated from the rest of the group or from particular individuals, in a group of baboons in the Moremi Wildlife Reserve, Botswana. We analyzed calls from 58 baboons of both sexes and all age classes. While the structure of the call appeared to be fixed from birth, call duration, the distribution of energy, and the fundamental frequency of the calls, including modulation, varied with age and sex of the caller. We discuss how body size may explain the variations. Some of the variables exhibited a different profile of variation with age between the sexes, with significant differences becoming apparent around puberty. The emergence of these sexual differences may be explained by the onset of sexual dimorphism in body size and mass. To which degree the hormonal status contributes to variation in the calling remains to be investigated.     

Preliminary observations on squirrel monkey (Saimiri oerstedi) vocalizations in Costa Rica

This study was an initial attempt to analyze quantitatively vocalizations from the Central American squirrel monkey, Saimiri oerstedi. Vocalizations were recorded in the wild at P. N. Corcovado (S. o. oerstedi). and at P. N. Manuel Antonio (S. o. citrinellus). Additional recordings were made from captive individuals of both subspecies. The acoustic structure of analyzed vocalizations resembled most closely the vocal repertoire of South American Saimiri with the “gothic arch” phenotype, in agreement with the prominent gothic arch facial pattern of S. oerstedi. New structural subtypes of the twitter (“dog-tooth twitter”) and the chuck (“bent-mast chuck”) not previously found in an extensive library of South American Saimiri sound spectrograms were documented. Calls used by older infants when socially separated and when approaching an adult male were essentially identical (sharing typical “isolation peep” structural features) but had significantly different duration and peak frequency. Analysis of recordings from an escaped captive female calling outside her compound over a 3-day period indicated the presence of numerous structural intermediates between typical chuck, twitter, and peep calls.     

Learned Vocal Variation Is Associated with Abrupt Cryptic Genetic Change in a Parrot Species Complex

Contact zones between subspecies or closely related species offer valuable insights into speciation processes. A typical feature of such zones is the presence of clinal variation in multiple traits. The nature of these traits and the concordance among clines are expected to influence whether and how quickly speciation will proceed. Learned signals, such as vocalizations in species having vocal learning (e.g. humans, many birds, bats and cetaceans), can exhibit rapid change and may accelerate reproductive isolation between populations. Therefore, particularly strong concordance among clines in learned signals and population genetic structure may be expected, even among continuous populations in the early stages of speciation. However, empirical evidence for this pattern is often limited because differences in vocalisations between populations are driven by habitat differences or have evolved in allopatry. We tested for this pattern in a unique system where we may be able to separate effects of habitat and evolutionary history. We studied geographic variation in the vocalizations of the crimson rosella (Platycercus elegans) parrot species complex. Parrots are well known for their life-long vocal learning and cognitive abilities. We analysed contact calls across a ca 1300 km transect encompassing populations that differed in neutral genetic markers and plumage colour. We found steep clinal changes in two acoustic variables (fundamental frequency and peak frequency position). The positions of the two clines in vocal traits were concordant with a steep cline in microsatellite-based genetic variation, but were discordant with the steep clines in mtDNA, plumage and habitat. Our study provides new evidence that vocal variation, in a species with vocal learning, can coincide with areas of restricted gene flow across geographically continuous populations. Our results suggest that traits that evolve culturally can be strongly associated with reduced gene flow between populations, and therefore may promote speciation, even in the absence of other barriers.     

Sources of Acoustic Variation in Rhesus Macaque (Macaca mulatta))Vocalizations

All species in the genus Macaca produce a set of harmonically rich vocalizations known as “coos”. Extensive acoustic variation occurs within this call type, a large proportion of which is thought to be associated with different social contexts such as mother-infant separation and the discovery of food. Prior studies of these calls have not taken into account the potential contributions of individual differences and changes in emotional or motivational state. To understand the function of a call and the perceptual salience of different acoustic features, however, it is important to determine the different sources of acoustic variation. I present data on the rhesus macaques' (M. mulatta) coo vocalization and attempt to establish some of the causes of acoustic variation. A large proportion of the variation observed was due to differences between individuals and to putative changes in arousal, not to differences in social context. Specifically, results from a discriminant-function analysis indicated that coo exemplars were accurately assigned to the appropriate individual, but vocal “signatures” were more variable in some contexts than in others. Moreover, vocal signatures may not always be reliable cues to caller identity because closely related individuals sound alike. Rhesus macaque coos evidently provide sufficient acoustic information for individual recognition and possibly kin recognition, but are unlikely to provide sufficient information about an external referent.     

The gradual vocal responses to human-provoked discomfort in farmed silver foxes

Vocal indicators of welfare have proven their use for many farmed and zoo animals and may be applied to farmed silver foxes as these animals display high vocal activity toward humans. Farmed silver foxes were selected mainly for fur, size, and litter sizes, but not for attitudes to people, so they are fearful of humans and have short-term welfare problems in their proximity. With a human approach test, we designed here the steady increase and decrease of fox–human distance and registered vocal responses of 25 farmed silver foxes. We analyzed the features of vocalizations produced by the foxes at different fox–human distances, assuming that changes in vocal responses reflect the degrees of human-related discomfort. For revealing the discomfort-related vocal traits in farmed silver foxes, we proposed and tested the algorithm of “joint calls,” equally applicable for analysis of all calls independently on their structure, either tonal or noisy. We discuss that the increase in proportion of time spent vocalizing and the shift of call energy toward higher frequencies may be integral vocal characteristics of short-term welfare problems in farmed silver foxes and probably in other captive mammals.     

Predicting Achievable Fundamental Frequency Ranges in Vocalization Across Species

Vocal folds are used as sound sources in various species, but it is unknown how vocal fold morphologies are optimized for different acoustic objectives. Here we identify two main variables affecting range of vocal fold vibration frequency, namely vocal fold elongation and tissue fiber stress. A simple vibrating string model is used to predict fundamental frequency ranges across species of different vocal fold sizes. While average fundamental frequency is predominantly determined by vocal fold length (larynx size), range of fundamental frequency is facilitated by (1) laryngeal muscles that control elongation and by (2) nonlinearity in tissue fiber tension. One adaptation that would increase fundamental frequency range is greater freedom in joint rotation or gliding of two cartilages (thyroid and cricoid), so that vocal fold length change is maximized. Alternatively, tissue layers can develop to bear a disproportionate fiber tension (i.e., a ligament with high density collagen fibers), increasing the fundamental frequency range and thereby vocal versatility. The range of fundamental frequency across species is thus not simply one-dimensional, but can be conceptualized as the dependent variable in a multi-dimensional morphospace. In humans, this could allow for variations that could be clinically important for voice therapy and vocal fold repair. Alternative solutions could also have importance in vocal training for singing and other highly-skilled vocalizations.     

Vocal Repertoire of Cebus capucinus: Acoustic Structure, Context, and Usage

Researchers studying nonhuman primate vocal repertoires suggest that convergent environmental, social, and motivational factors account for intra- and interspecific vocal variation. We provide a detailed overview of the vocal repertoire of white-faced capuchins, including acoustic analyses and contextual information of vocal production and vocal usage by different age-sex classes in social interactions. The repertoire is a mixture of graded and discrete vocalizations. In addition, there is general support for structural variation in vocalizations with changes in arousal level. We also identified several combined vocalizations, which might represent variable underlying motivations. Lastly, by including data on the social contexts and production of vocalizations by different age-sex classes, we provide preliminary information about the function of vocalizations in social interactions for individuals of different rank, age, and sex. Future studies are necessary to explore the function of combined vocalizations and how the social function of vocalizations relate to their acoustic structure, because social use of vocalizations may play an important role in shaping vocal evolution.     

Vocal Production by a Language-Competent Pan paniscus

Human spoken language and nonhuman primate vocalization systems have traditionally been regarded as qualitatively different from one another with respect to their semanticity and the way in which individuals acquire and utilize these signals. However, recent studies of the vocal behaviors of both captive and free-ranging monkeys and apes suggest that this dichotomy may not be unequivocal. We examined the vocalizations produced by a linguistically-competent adult male bonobo (Pan paniscus) named Kanzi. We analyzed his vocalizations during communicative interactions with humans in order to determine whether they vary systematically according to the semantic context in which they are produced. We determined semantic contexts based upon a vocalization's co-occurrence with predefined behavioral correlates. Spectrographic and statistical analyses revealed that acoustic structure is similar among the vocalizations that occurred within a specific semantic context and structural differences are evident between the vocalizations produced in different contexts. The results provide evidence that, during communicative interactions with humans, Kanzi modulates his vocal output on both the temporal and spectral levels.     

Etude du répertoire vocal du Gibbon a favoris blancs (Hylobates concolor leucogenys): Analyse structurale des vocalisations

The basic vocal repertoire of the Crested Gibbon (Hylobates concolor leucogenys) can be defined from qualitative and quantitative analysis of the vocalizations. From sonagraphic analysis of calls of 9 juveniles, one adult ♂, one sub-adult ♂ and one sub-adult ♀ arranged in three semi-free living groups, one can describe six vocal types on a morphological basis. The sound repertoire of juveniles is composed of graded signals which seem to become more discrete and fewer as the individual matures. These features are compared with other Cercopithecides and Pongides on the basis of eco-ethological and ontogenetic implications.     

Killer whales are capable of vocal learning

The production learning of vocalizations by manipulation of the sound production organs to alter the physical structure of sound has been demonstrated in only a few mammals. In this natural experiment, we document the vocal behaviour of two juvenile killer whales, Orcinus orca, separated from their natal pods, which are the only cases of dispersal seen during the three decades of observation of their populations. We find mimicry of California sea lion (Zalophus californianus) barks, demonstrating the vocal production learning ability for one of the calves. We also find differences in call usage (compared to the natal pod) that may reflect the absence of a repertoire model from tutors or some unknown effect related to isolation or context.     

A field study on the vocalizations of Goeldi's monkeys (Callimico goeldii)

The vocalizations ofCallimico goeldii inhabiting North Bolivia were recorded and analyzed. A total vocal repertoire of 40, including two ultrasonic vocalizations, was recognized. The situation in which each vocalization was emitted is described. The phonetic structure of the longdistance location calls varied precisely dependent on the interindividual distances, while distinct individual variations were seen in the short-distance location calls. Every agonistic call could grade into another, from that reflecting the most aggressive tendency to that of fear emotion. The repertoire and characteristics of the infantile calls are also described, and their ontogeny is discussed.