Context, Structural Variability and Distinctiveness of California Towhee (Pipilo crissalis) Vocal Duets

Avian vocal duets occur when paired birds produce temporally and structurally coordinated vocalizations. Duets are given by members of many species from taxonomically distinct lineages and show great variety in form that often reflects function. By describing the structure of vocal duets we can learn about the diversity of communication signals present in nature and also gain insight into the evolution and operation of those signals. This study quantified the usage patterns and acoustic structure of California towhee duet vocalizations, and tested the distinctiveness of duets among different pairs. California towhee duets consist of a 'squeal' vocalization that is highly unlike the species-typical call note or male advertizement song. California towhees duet an average of three times per hour during the breeding season, and all duets are accompanied by an approach response that brings duetting partners into close spatial proximity. Males and females produce duet contributions with the same syllabic structure. Individual birds produce highly variable squeal vocalizations that are distinctive enough to signal identity. California towhee duet characteristics indicate that the squeal vocalization has evolved separately from other vocal traits in this species and was promoted by natural selection as a duet-specific vocalization. Duet usage patterns and structure suggest that these communication signals function in a cooperative context.     

Evidence for High Variability in Temporal Features of the Male Coda in Müller’s Bornean Gibbons (<Emphasis Type="Italic">Hylobates muelleri</Emphasis>)

Most pair-living primate species engage in duets, wherein males and females produce coordinated vocalizations. Previous analyses of male gibbon contributions to the duet have shown that calls are individually distinct. Here we investigate variation in the temporal and spectral parameters in the male contribution to the duet, also known as the coda, of wild, nonhabituated male Müller’s Bornean gibbons (Hylobates muelleri), recorded both opportunistically and as a response to playbacks at the Stability of Altered Forest Ecosystems site in Sabah, Malaysia. We used linear discriminant function analysis to estimate the intra- and interindividual variation in 13 spectral and temporal parameters of the vocalizations (N?=?337) of 31 male gibbons. To further understand how call features vary within and between individuals we used a multivariate, variance components model to investigate how variance in features was partitioned at these two levels. We could identify males with a 66% accuracy using leave-one-out cross-validation, a relatively low score compared to female Müller’s Bornean gibbons and males of other species. We found that for some features (such as maximum frequency of the notes) most of the variance occurred between males, but for others (specifically total duration of the call and duration of rest in between notes) most of the variance occurred within a single male. Overall, male Müller’s Bornean gibbon coda vocalizations showed greater variability relative to their female counterparts, raising questions about potential differences in the function of the male and female contributions to the duet in Müller’s Bornean gibbons and the gibbon taxon as a whole.     

Angry Birds Need Not Apply: A Perspective on the Flexible form and Multifunctionality of Avian Vocal Duets

Research into avian vocal duets has revealed much about the social interactions of mated pairs, including how communication signals mediate important components of fitness such as reproductive success and resource‐holding ability. Excellent reviews have been written on avian duets, but none have quantified the prevalence of different duet features and functions, or tested for potential links between structure and function. Here we have compiled a thorough list of studies at the species level and investigated whether there are trends in duet form, trends in duet function and matches between form and function. We have also provided recommendations for reporting of duet structure to help standardize future data collection and facilitate comparative analyses. We found that duet vocalizations are typically tightly coordinated (69% of 33 species), initiated by both partners (80% of 35 species), and have sex‐specific contributions (73% of 48 species). Duets vary substantially in structure both across and within species, and this flexibility in duet form may contribute to their use in multiple contexts. Among the well‐studied species (functions well supported by experimental evidence and studied post‐1983), duets are usually multifunctional, with functions including joint resource defence, signalling commitment, maintaining contact, ensuring reproductive synchrony and mate‐guarding. Levels of support for different hypotheses have changed over time as theory within the field developed, but overall, the best‐supported function of duets across avian species (81% of 26 species) has been joint resource defence. In all but one of these 26 species, duets are sometimes cooperative (meaning that duet participation benefits both duetting partners), and they often produce multiple benefits. We found associations between form and function, in that duets directed to non‐pair individuals have higher amplitude and are more likely to have sex‐specific notes than duets directed within the pair. We see the field moving away from labelling duets as representing ‘cooperation’ or ‘conflict’ and moving towards a general multifunctionality framework within which researchers test multiple hypotheses for duet function under the assumption that both duet form and function can vary moment to moment and that such flexibility can provide multiple benefits.     

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.     

Contexts and behavioral correlates of trill vocalizations in wild white-faced capuchin monkeys (Cebus capucinus)

Primate vocalizations that appear to occur independently of specific contexts typically are considered to be contact calls. However, results from several recent studies indicate that these calls function to facilitate social interactions. White-faced capuchin monkeys (Cebus capucinus) emit a high-frequency vocalization, termed a "trill," in social interactions and during travel. In this study, immatures emitted most trills, but adult females also trilled; by contrast, adult males rarely trilled. Infants emitted the majority of trills, and they trilled at significantly higher rates than adult females. Infants trilled most when approaching other individuals. Furthermore, infants emitted proportionately more trills than other age classes when approaching other individuals. I therefore focused on the detailed context and immediate behavioral correlates of trilling by infants. Infants that trilled when approaching others tended to interact affiliatively with them subsequently (i.e., climbing on, touching, receiving grooming, and performing food inspection) more than infants that did not trill when approaching. Therefore, infant trilling may have had an immediate effect on the recipient's behavior.     

Duetting in the Titi monkey Callicebus cupreus: structure,pair specificity and development of duets

Duets from a total of 6 female and 6 male titi monkeys (Callicebus cupreus) were analysed in order to provide an in-depth analysis of the duet structure, to investigate whether duets are pair specific and to examine whether duets of newly formed pairs differ from those of established pairs. Callicebus duets are composed of alternately uttered male and female contributions. Duets were pair specific as a result of a summation of individual attributes of the two mates rather than due to unidirectional or reciprocal song adaptation by mates. The two examined newly formed pairs produced species-typical duets from the first day on, but their duets showed greater variability in sequence durations than those of established pairs.     

Breeding season length predicts duet coordination and consistency in Neotropical wrens (Troglodytidae)

Many animals produce coordinated signals, but few are more striking than the elaborate male–female vocal duets produced by some tropical songbirds. Yet, little is known about the factors driving the extreme levels of vocal coordination between mated pairs in these taxa. We examined evolutionary patterns of duet coordination and their potential evolutionary drivers in Neotropical wrens (Troglodytidae), a songbird family well known for highly coordinated duets. Across 23 wren species, we show that the degree of coordination and precision with which pairs combine their songs into duets varies by species. This includes some species that alternate their song phrases with exceptional coordination to produce rapidly alternating duets that are highly consistent across renditions. These highly coordinated, consistent duets evolved independently in multiple wren species. Duet coordination and consistency are greatest in species with especially long breeding seasons, but neither duet coordination nor consistency are correlated with clutch size, conspecific abundance or vegetation density. These results suggest that tightly coordinated duets play an important role in mediating breeding behaviour, possibly by signalling commitment or coalition of the pair to mates and other conspecifics.     

Siberian crane duet as an individual signature of a pair: comparison of visual and statistical classification techniques

Vocal individuality varies between species and/or ontogenesis stages depending on needs in the vocal recognition, but also estimation of individual differences depends on the method of analysis. We studied pair-specific differences of duets elicited by mating pairs of Siberian crane Grus leucogeranus. We quantitatively described the duet structure and compared visual and statistical classification methods of pair identification by duet. Three methods were used: discriminant analysis, method of classification trees and visual classification of spectrogram. We found significant interpair differences. The pairs differ by duet structure that is by the ratio of male- and female-initiated duets and by the ratio of the number of male to female calls; temporal-frequency duet characteristics are pair-specific, too. All methods showed high interpair differences, which exceeded random values significantly. Discriminant analysis stepwise procedure based on 11 parameters resulted in 97.3% of correctly assigned duets. Human observers correctly assigned 80.7% of spectrograms. Our data provide a basis for remote monitoring of this endangered species with a wild population of only 3,000 birds.     

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ABSTRACT

Research on African elephant (Loxodonta africana) vocal communication has increased in recent years, yet there has been very little data collected on the vocal production of infant African elephants. Vocalizations were recorded from a group of five adult female African elephants and 3 dependent offspring (1 male and 2 female) at Disney's Animal Kingdom, Florida, U.S.A., using custom-designed audio-recording collars worn by the adult females. We measured both source and filter features of infant ‘rumble’ vocalizations made during affiliative social interactions and after cessation of nursing from adult females. Rumble vocalizations produced in the ‘nurse cessation’ context exhibited an upward shift in formant frequency locations, compared to rumbles produced during the ‘affiliation’ context. Additionally, call duration increased and fundamental frequencies decreased after nurse cessations for the male, but both females showed the opposite acoustic response. When infant rumbles accompanied nurse cessations, nursing was more likely to resume within 30 seconds compared to nurse cessations without vocalizations. These results suggest that infant rumbles associated with cessation of nursing reflect the motivational state of infants and may influence maternal responsiveness.     

Food Transfer and Development of Feeding Behavior and Food-Associated Vocalizations in Cotton-Top Tamarins

Food transfer between adults and infants is common in many marmoset and tamarin monkeys, and is often accompanied by vocalizations. We hypothesized that vocalizations by adults in a food transfer context creates an opportunity for infants to learn not only what foods are appropriate but what vocalizations are appropriate in feeding contexts. We studied the development of feeding behavior and food-associated vocalizations in 10 infant cotton-top tamarins through the first 20 wks of life. Infants obtained solid food through transfers from older group members, primarily the adult male, beginning at weeks 5–6. Both adults and infants vocalized during food transfers with adults, producing rapid sequences of the call types adults normally give when feeding. Infants were usually successful in obtaining food primarily when the adult was vocalizing. The sooner infants were active participants in food transfers, the sooner they began to feed independently. In the early weeks, infants produced a large number of vocal types during food transfers, but with increasing age there was a steady increase in the number of adult-form food calls and a reduction in other, non-food-associated calls. Infants that fed independently at an early age produced fewer non-food-associated calls by the last month of observation. Infants called at higher rates to their most preferred food. Food transfers accompanied by vocalizations may provide an opportunity for infants to learn about appropriate foods as well as the vocalizations that accompany feeding in adults, and may represent a form of 'coaching' or information donation by adults.     

Ecological and Social Factors Affecting the Vocal Behavior of Adult Female Squirrel Monkeys

Focal recordings of the vocalizations of squirrel monkeys, Saimiri sciureus, occupying an undisturbed Peruvian habitat were collected to evaluate the importance of both the ecological and social functions of female vocal behavior. The rates and sequences of six call types were examined by context: single, double, and multiple chuck, peep, tweet, and tweet-chuck. In contrast to laboratory studies, our findings emphasize the primacy of ecological functions in the wild, where calling permits females to operate as a convoy of spatially separated individuals in an arboreal habitat where visual contact is limited. Chuck calls are probably best interpreted as contact calls. The rate of chucks and tweet-chucks increased when the nearest adult female was > 5 m away. However, call production did not predict a change in the spatial separation between a female and her nearest adult female. During travel, the rate of single and double chucks was greater and the rate of multiple chucks was reduced. No direct relationship was found between foraging activities and vocal behavior. Although secondary, social factors did have a subtle effect on vocal behavior: the larger the social alliance of a female, the lower the rate and repetitiveness of her chuck vocalizations. We also consider the role of variations in vocalization rates among primates and the differing conclusions of field and captive studies of squirrel monkey vocal behavior.     

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 coordination of troop movement among white-faced capuchin monkeys,Cebus capucinus

Coordinated travel by social groups is well documented, often with evidence that cognitive spatial maps are employed. Yet the mechanisms by which movement decisions are made and implemented within social groups are poorly known. In a field study of white-faced capuchin monkeys in Costa Rica it was demonstrated that a specific call, the “trill,” is used by adults in the initiation and directing of troop movement. The trills of subadults were restricted to vocal exchanges with other subadults. Continuous vocal recordings were collected of the vocalizations of the 14 members of the study troop. A cumulative 33.7 h of continuous samples and 1,892 sonagrams were analyzed. In addition to vocalizations clearly associated with alarm, distress, or agonistic contexts, two distinct call types were identified, trills and huhs. Age-sex classes differed in the rate at which both types of calls were produced in different spatial positions within the troop. Adult females and males produced higher rates of trills when in the leading edge compared to all other spatial positions in a traveling troop. Trills at the edge of a stationary troop represented 36 “successful” and 3 “unsuccessful” start attempts; the troop usually moved in the trajectory predicted by a trilling adult's location on the troop periphery within 10 min of the initiation of trilling. Adults also altered the trajectory of traveling troops by trilling at the side and back of the troop (10 “successful” and 4 “unsuccessful” attempts). Huh vocalizations were most predictably produced when a capuchin is in a dense fruit patch. These results emphasize the role vocalizations serve in the coordination and trajectory of group movement in nonhuman primates, especially those populations that are arboreal or in which visual contact is otherwise impeded. © 1993 Wiley-Liss, Inc.

1 This article is a US Government work and, as such, is in the public domain in the United States of America.

     

A preliminary field investigation of the yellow handed titi monkey,Callicebus torquatus torquatus,in Northern Peru

A three month field study was conducted onCallicebus torquatus near the Nanay River in northern Peru. Data were collected primarily by the time sampling method. Information on daily movement, vocalizations, population density, and social behavior is presented. The titi monkeys we studied have a family unit pattern of social organization, territoriality accompanied by vocal activity, and a home range of about 20 hectares. There is a high degree of parental investment in care of the infant on the part of the adult male. A preliminary version of this paper was presented at the 73rd Annual meeting, American Anthropological Association, Mexico City, November, 1974.     

Vocal behaviour and neighbour spatial arrangement during vocal displays in eagle owls (Bubo bubo)

We quantified temporal and spatial patterns of adult eagle owl Bubo bubo vocal behaviour throughout an entire year. The duration of adult eagle owl vocal displays showed significant differences during different periods of the year: there was one major peak in the pre-laying period, when duets of mates were also more frequent. The daily distribution of adult vocalizations showed a similar pattern among the different periods, with vocal activity being most intense at sunset and sunrise. Analyses of the characteristics of call posts showed that the choice of such focal points was guided by the trade-off between the need to defend the territory and within-pair communication inside the core areas, as well as efficient communication with neighbours.     

Vocal negotiation over parental care? Acoustic communication at the nest predicts partners' incubation share

In species with biparental care, individuals adjust their workload to that of their partner to either compensate or match its investment. Communication within a pair might be crucial for achieving this adjustment. Zebra finches, Taeniopygia guttata, form life‐long monogamous pair bonds, in which partners are highly coordinated and both incubate the eggs. When relieving each other during incubation, partners perform a structured call duet at the nest. If this duet functions to coordinate incubation workload, disrupting the pair's usual nest‐relief pattern by delaying the male's return to the nest should affect the structure of the duet. Using domesticated birds breeding in a large aviary, we found that delaying the male's return induced shorter duets with higher call rates. In addition, we tracked the location of individuals with a transponder at the nest and the feeder, and showed that these accelerated duets were associated with an increased haste of the partners to take turns incubating and foraging. Females also spent less time incubating during their subsequent shift, and females' time off‐nest was best predicted by their mate's calling behaviour in the previous duet. Taken together, these results suggest that duets may function as ‘vocal negotiation’ over parental care.     

Convergence and divergence in Diana monkey vocalizations

Individually distinct vocalizations are widespread among social animals, presumably caused by variation in vocal tract anatomy. A less-explored source of individual variation is due to learned movement patterns of the vocal tract, which can lead to vocal convergence or divergence in social groups. We studied patterns of acoustic similarity in a social call produced by 14 female Diana monkeys (Cercopithecus diana) in two free-ranging groups. Calls showed variability in fundamental frequency contours owing to individual identity and external context. Vocal divergence increased significantly between females during poor visibility and tended to increase in the presence of neighbours. In contrast, vocal convergence increased significantly between females during vocal interactions, because females matched the frequency contour of their own call with another female's preceding call. Our findings demonstrate that these primates have some control over the acoustic fine structure of their most important social vocalization. Vocal convergence and divergence are two opposing processes that enable callers to ensure spatial proximity and social cohesion with other group members.     

The Responses of Duetting Antbirds to Stereo Duet Playback Provide Support for the Joint Territory Defence Hypothesis

In multiple animal taxa, including many birds and primates, members of mated pairs produce coordinated acoustic displays known as duets. By observing the behaviour of territorial animals as they respond to playback‐simulated duets of rivals, we can gain insight into the behavioural significance of vocal duets. Playback experiments, however, have been conducted across a very narrow range of duetting animals. Furthermore, many studies have been conducted with single‐speaker playback, whereas stereo‐speaker playback offers more spatially realistic simulation of duets. Moreover, by evaluating the reactions of animals to separate loudspeakers broadcasting male and female duet contributions, we can study the interactions of both males and females with same‐sex vs. opposite‐sex rivals. We used a paired experimental design to broadcast duet stimuli through a single‐speaker and a stereo‐speaker apparatus to 30 pairs of duetting barred antshrikes Thamnophilus doliatus in Costa Rica. Our goals were (1) to evaluate whether territorial antbirds respond more aggressively to male vs. female duet components and (2) to assess aggressive responses of antbirds towards single‐speaker vs. stereo‐speaker playback. Neither males nor females differentiated between the loudspeaker simulating the male vs. female duet contribution during stereo‐speaker playback trials. Barred antshrikes displayed significantly stronger responses to stereo‐speaker playback compared with single‐speaker playback. Males displayed stronger playback responses than females with closer, quicker and more vocal responses. These results provide evidence for a joint resource defence function of antbird duets given that pairs responded together with equivalent intensity to male and female simulated intruders. This is the first study to show that although duetting is an aggressive territorial signal, birds do not necessarily respond to sex‐specific components of duets. Our results support the idea that spatially realistic stereo presentation of duet stimuli is critical for experimental duet research.     

Antiphonal duetting in Steere’s liocichla (Liocichla steerii): male song individuality and correlation between habitat and duetting behavior

We studied duet song and vocal duetting behavior in an endemic Taiwanese passerine, Steeres liocichla (Liocichla steerii). We found that the leading male song in duets was highly individualistic. Also, we found duetting behavior varied significantly across different habitat types. Females were more likely to answer male songs in densely vegetated, steep forest habitat compared to open agricultural habitat. These findings provide quantitative evidence for vocal individuality for a duet song and provide tentative support for the idea that females are duetting to reveal their location to their mates.