Spectral acoustic structure of barking in roe deer (Capreolus capreolus). Sex-, age- and individual-related variations

In roe deer, barking is a loud call commonly given by males and females during inter- or intraspecific interactions. The analysis of a set of 19 spectral variables computed on 560 calls revealed significant variation between sexes, individuals, and probably age classes. Discriminant analysis predicted the sex of an individual with a 93.5% probability from a small portion of the bark frequency range. Among six males, a linear combination of six variables predicted the identity of the barking individual with a 70% probability. These sexual and individual differences provide the potential for social recognition from vocalizations. These results are consistent with the hypothesis that barking in roe deer may allow remote signalling of presence, location and identity, and play an important role in the territorial system of this species.     

Cultural transmission of vocalizations in ravens, Corvus corax

As ravens have pronounced learning capabilities, especially in the vocal domain, cultural transmission might be a relevant factor affecting the composition of their vocal repertoires. We investigated how important cultural transmission is in establishing a raven's vocal repertoire and whether there are different pathways of transmission. We studied individually known free-ranging breeding pairs whose vocalizations were recorded during interactions with a caged pair. The mean repertoire size of these vocalizations was 12 call types per individual (from 79 types recorded from 74 individuals), with no difference between the sexes. In 81% of individuals the repertoire was composed entirely of calls also used by other individuals. Analysis of the distribution of call types among individuals regarding sex, partnership and neighbourhood and aspects of the geographical distribution of calls showed that the main pathway of cultural transmission was within the sexes. In addition, transmission occurred between pair partners, as well as between neighbouring and more distant pairs. About 40% of the calls in a raven's repertoire were transmitted within a sex and 10% between pair partners. Any two given birds in the population shared on average 20% (0-77%) of their calls. Cultural transmission within the sexes led to a pronounced sexual dimorphism in vocal behaviour. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

Vocal cooperation between the sexes in Little Spotted Kiwi Apteryx owenii

Sexual call dimorphism in birds is usually associated with sexual size dimorphism. Departures from this relationship can be used to infer call function, but research into inter‐sexual call differences, as with song function in general, has been restricted by a bias towards male passerines. The nocturnal and flightless New Zealand kiwi (Apterygidae) are acoustically similar but taxonomically and ecologically very different from other birds, so provide a contrast in exploring avian call function and evolution. However, kiwi acoustic ecology is poorly understood, with the calls of only one of the five kiwi species spectrally described, and acoustic differences between the sexes virtually unknown. We conducted the first bioacoustic study of Little Spotted Kiwi Apteryx owenii, and assessed sexual call dimorphism in this species. There were significant inter‐sexual differences in call temporal and frequency characteristics that were not related to size dimorphism. Contribution to duets and variation in temporal structure with call context also differed between the sexes. We suggest that these differences indicate divergent call function, with male calls more suited for territory defence, and female calls for pair contact. There was a striking lack of overlap in the frequency spectrum distributions of male and female calls, which was also unrelated to size and was further emphasized by the presence of formants in female calls. We propose that this provides evidence for inter‐sexual acoustic cooperation in call frequency, of a type which to our knowledge has not previously been described in birds. This may result from selection for enhanced joint resource defence in kiwi.     

Can Nocturnal Flight Calls of the Migrating Songbird,American Redstart,Encode Sexual Dimorphism and Individual Identity?

Bird species often use flight calls to engage in social behavior, for instance maintain group cohesion and to signal individual identity, kin or social associations, or breeding status of the caller. Additional uses also exist, in particular among migrating songbirds for communication during nocturnal migration. However, our understanding of the information that these vocalizations convey is incomplete, especially in nocturnal scenarios. To examine whether information about signaler traits could be encoded in flight calls we quantified several acoustic characteristics from calls of a nocturnally migrating songbird, the American Redstart. We recorded calls from temporarily captured wild specimens during mist-netting at the Powdermill Avian Research Center in Rector, PA. We measured call similarity among and within individuals, genders, and age groups. Calls from the same individual were significantly more similar to one another than to the calls of other individuals, and calls were significantly more similar among individuals of the same sex than between sexes. Flight calls from hatching-year and after hatching-year individuals were not significantly different. Our results suggest that American Redstart flight calls may carry identifiers of gender and individual identity. To our knowledge, this is the first evidence of individuality or sexual dimorphism in the flight calls of a migratory songbird. Furthermore, our results suggest that flight calls may have more explicit functions beyond simple group contact and cohesion. Nocturnal migration may require coordination among numerous individuals, and the use of flight calls to transmit information among intra- and conspecifics could be advantageous. Applying approaches that account for such individual and gender information may enable more advanced research using acoustic monitoring.     

Sex recognition in brown skuas: do acoustic signals matter?

Bird vocalisations are often essential for sex recognition, especially in species that show little morphological sex dimorphism. Brown skuas (Catharacta antarctica lonnbergi), which exhibit uniform plumage across both sexes, emit three main calls: the long call, the alarm call and the contact call. We tested the potential for sex recognition in brown skua calls of 42 genetically sexed individuals by analysing 8–12 acoustic parameters in the temporal and frequency domains of each call type. For every call type, we failed to find sex differences in any of the acoustic parameters measured. Stepwise discriminant function analysis (DFA) revealed that sexes cannot be unambiguously classified, with increasing uncertainty of correct classification from contact calls to long calls to alarm calls. Consequently, acoustic signalling is probably not the key mechanism for sex recognition in brown skuas.     

Evidence for Dialects in Three Captive Populations of Common Marmosets (<Emphasis Type="Italic">Callithrix jacchus</Emphasis>)

The vocal repertoires of nonhuman primates have long been thought to be invariable across populations and not to result from vocal learning. However, increasing evidence suggests that learning does influence vocal production in nonhuman primates, and that several species modify the structure of their calls in response to social or environmental influences. Vocal usage learning refers to the process whereby an individual learns in which circumstances to produce a certain call type, whereas vocal production learning refers to the process in which signals get modified as the result of individual experiences. Common marmosets (Callithrix jacchus) show socially mediated vocal plasticity as adults and during vocal development. This propensity to engage in simple forms of vocal production learning (accommodation) should produce population-level differences in call structure. To test this prediction, we compared the vocalizations of three captive populations of common marmosets. We analyzed the acoustic structure of 1337 phee calls, 461 trills, and 3611 food calls and compared them with a permutated discriminant function analysis. We found that all call types differed significantly between the three populations, and 76–98% of the calls were correctly classified. As physical differences in body mass and environmental differences between colonies could not explain the call differences, we conclude that vocal accommodation is the most likely explanation for the differences in call structure. This will allow us to further investigate the role and importance of vocal learning in a species increasingly used to study vocal learning and language evolution.     

人为疼痛刺激下金色中仓鼠幼鼠叫声的发育过程

金色中仓鼠幼鼠可以利用可听声和超声信号与母鼠进行交流,这些声音可以反映个体的寒冷、饥饿、疼痛等不同生理状态和需求。因疼痛诱发的叫声对于维持幼鼠的存活也有着重要意义。本实验通过录制人为疼痛刺激下,不同日龄金色中仓鼠的叫声,分析并比较了5 - 30 日龄金色中仓鼠不同发声信号特征(可听声出现频次、可听声持续时程、可听声主频率、超声出现频次、超声持续时程、超声主频率)的性别差异;同时记录了幼鼠的两种发声随日龄的变化趋势。结果发现:幼鼠两种叫声的各种参数均未表现性二型现象。幼鼠超声发生频次随日龄增加而逐渐减少。而可听声却正相反,随日龄增加而逐渐上升,并在20 日龄时达到最高峰,之后逐渐下降。可听声持续时程也随日龄而增加。综上,在人为疼痛刺激下,随着日龄增加,幼鼠更倾向于使用可听 声而非超声与母兽进行交流。     

Acoustic Features of Female Chacma Baboon Barks

We studied variation in the loud barks of free-ranging female chacma baboons ( Papio cynocephalus ursinus ) with respect to context, predator type, and individuality over an 18-month period in the Moremi Game Reserve, Botswana. To examine acoustic differences in relation to these variables, we extracted a suite of acoustic parameters from digitized calls and applied discriminant function analyses. The barks constitute a graded continuum, ranging from a tonal, harmonically rich call into a call with a more noisy, harsh structure. Tonal barks are typically given when the signaler is at risk of losing contact with the group or when a mother and infant have become separated (contact barks). The harsher variants are given in response to large predators (alarm barks). However, there are also intermediate forms between the two subtypes which may occur in both situations. This finding is not due to an overlap of individuals' distinct distributions but can be replicated within individuals. Within the alarm bark category, there are significant differences between calls given in response to mammalian carnivores and those given in response to crocodiles. Again, there are intermediate variants. Both alarm call types are equally different from contact barks, indicating that the calls vary along different dimensions. Finally, there are consistent, significant differences among different individuals' calls. However, individual identity in one call type cannot directly be inferred from knowledge of the individuals' call characteristics in the other. In sum, the barks of female baboons potentially provide rich information to the recipients of these signals. The extent to which baboons discriminate between alarm and contact barks, and classify calls according to context and/or acoustic similarity will be described in a subsequent paper.     

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.     

Sex identification in King Penguins Aptenodytes patagonicus through morphological and acoustic cues

In the context of sexual selection, animals have developed a variety of cues conveying information about the sex of an individual to conspecifics. In many colonial seabird species, where females and males are monomorphic and do not show obvious differences in external morphology, acoustic cues are an important signal for individual and sex recognition. Here, we study the vocal and morphological sex dimorphism in the King Penguin Aptenodytes patagonicus, a colonial, monomorphic seabird for which our knowledge about the role of vocalizations and morphology in mate choice is very limited. Data were collected at Possession Island, Crozet Archipelago, in a breeding colony consisting of about 16 000 breeding pairs. Using measurements of six morphological features and analysing acoustic parameters of call recordings of adult individuals, we show that King Penguins can be sexed based on a single morphological measurement of the beak with an accuracy of 79%. We found a sex‐specific syntax in adult King Penguin calls that provided a 100% accurate method to distinguish between the sexes in our study population. To confirm the method at the species level, we analysed calls recorded from King Penguin adults in Kerguelen Island, 1300 km away from our study population and found the same accuracy of the sex‐specific syntax. This sex‐specific syllable arrangement is rare in non‐passerines and is a first step in understanding the mate choice process in this species. Furthermore, it offers a cost‐effective, non‐invasive technique for researchers to sex King Penguins in the field.     

Anatomical bases of sex‐ and size‐related acoustic variation in herring gull alarm calls

The hypothesis that anatomical or physiological factors can constrain the production of vocalizations is supported by an increasing number of examples from a range of taxa, where acoustic variation is related to sex, body‐size or condition. In this study, we combine anatomical and acoustic investigations in herring gulls Larus argentatus to 1) identify co‐variation between sex, body size and the dimensions of the vocal apparatus and 2) test the possible effect of this co‐variation on interindividual variation in the acoustics of alarm calls. We found that the vocal apparatus was sexually dimorphic, with males having longer trachea and bigger vibratile membranes than females. We also identified a relationship between the head–bill length – a secondary sexual trait – and the length of the trachea in males only. However, we failed to identify corresponding sex‐ and body‐size related variation in the acoustic components of alarm calls. We suggest that this absence of a relationship between anatomical and acoustic dimensions may reflect the lack of biomechanical constraints exerted during the production of alarm calls, and that such relationships are more likely to be expressed in this species’ sexual calls, whose production is characterised by more pronounced, ritualised postures that are more likely to highlight inter‐individual size variation.     

Dual Function of Chip Calls Depending on Changing Call Rate Related to Risk Level in Territorial Pairs of White‐Eared Ground‐Sparrows

Different mechanisms have been proposed for encoding information into vocalizations: variation of frequency or temporal characteristics, variation in the rate of vocalization production, and use of different vocalization types. We analyze the effect of rate variation on the dual function of chip calls (contact and alarm) produced by White‐eared Ground‐sparrows (Melozone leucotis). We conducted an acoustic playback experiment where we played back 1 min of four chip call rates (12, 36, 60, 84 calls/min). We measured the response of territorial pairs using behavioral responses, and fine structural features of calls produced in response to those playbacks. White‐eared Ground‐sparrows showed more intense behavioral responses to higher than lower call rate playbacks. Both individuals of the pair approached the source of the playback stimulus faster, produced the first vocalization faster, produced more vocalizations, and spent more time close to the stimulus in higher call rate than in lower call rate playbacks. Frequency and duration characteristics of calls (chip and tseet) were similar in response to all call rate playbacks. Our playback experiment elicited different intensity of behavioral responses, suggesting that risk‐based information is encoded in call rate. Our results suggest that variation in the rate of chip call production serves a dual function in this species; calls are used at lower rates for pair contact and at higher rates for alarm/mobbing signals.     

Geographic Variation in the Long Calls of Male Orangutans (Pongo spp.)

Relatively few data exist on population differences in the vocal behavior of mammals. Geographic variation in calls is of special interest because of the implications for resolving evolutionary and behavioral questions. For example, information on geographic variation in vocalizations complements morphological and molecular data used to infer phylogenetic relationships and provides evidence for the mechanisms underlying call development. A quantitative acoustic analysis of orangutan long calls was undertaken, comparing flanged adult males from four geographically distinct sites across Borneo and Sumatra, revealing consistent differences among the calls of individuals. Long calls produced by orangutans from the four sites in Borneo and Sumatra differ in quantitative acoustic measures. Discriminant function analysis reveals that acoustic variables can be used in combination to assign calls to the correct individual, site and island at rates higher than that expected by chance. Specifically, four acoustic parameters proved reliable for distinguishing among the individuals, between sites, and across the two islands that arguably represent populations from separate species or subspecies. Although Bornean and Sumatran long calls share a repetitive structure and show similar call rates (0.100–0.500 LCs/h) and maximum frequency bands (0.400–1.500 kHz), they differ significantly in the number of pulses per call, call speed, call duration, bandwidth, pulse duration, and dominant frequency. Strong consistency in these acoustic parameters is also seen among males within sites and the observed variation may allow for individual recognition. Individual identification by call structure presumably benefits dispersed orangutans, where individuals characteristically forage independently and both encounters and interactions with signaling males are highly variable and largely dependent on context. Acoustic recognition of callers facilitates the choice of which males to join or avoid, thus allowing receivers to manipulate potential costs and benefits of association.     

Production and perception of sex differences in vocalizations of Wied's black‐tufted‐ear marmosets (Callithrix kuhlii)

Males and females from many species produce distinct acoustic variations of functionally identical call types. Social behavior may be primed by sex‐specific variation in acoustic features of calls. We present a series of acoustic analyses and playback experiments as methods for investigating this subject. Acoustic parameters of phee calls produced by Wied's black‐tufted‐ear marmosets (Callithrix kuhlii) were analyzed for sex differences. Discriminant function analyses showed that calls contained sufficient acoustic variation to predict the sex of the caller. Several frequency variables differed significantly between the sexes. Natural and synthesized calls were presented to male–female pairs. Calls elicited differential behavioral responses based on the sex of the caller. Marmosets became significantly more vigilant following the playback of male phee calls (both natural and synthetic) than following female phee calls. In a second playback experiment, synthesized calls were modified by independently manipulating three parameters that were known to differ between the sexes (low‐, peak‐, and end‐frequency). When end‐frequency‐modified calls were presented, responsiveness was differentiable by sex of caller but did not differ from responses to natural calls. This suggests that marmosets did not use end‐frequency to determine the sex of the caller. Manipulation of peak‐and low‐frequency parameters eliminated the discrete behavioral responses to male and female calls. Together, these parameters may be important features that encode for the sex‐specific signal. Recognition of sex by acoustic cues seems to be a multivariate process that depends on the congruency of acoustic features. Am. J. Primatol. 71:324–332, 2009. © 2008 Wiley‐Liss, Inc.     

Vocal Repertoire of Golden-backed Uakaris (<Emphasis Type="Italic">Cacajao melanocephalus</Emphasis>): Call Structure and Context

The study of vocal behavior can reveal important aspects of how and why a species communicates in relation to ecological and social challenges. We here focus on vocal communication in golden-backed uakaris (Cacajao melanocephalus), diurnal, pitheciine monkeys that exhibit fission-fusion social organization and typically inhabit dense forests that limit the potential for visual communication. Moreover, the species spends little time engaged in tactile or olfactory communication, e.g., social grooming and scent marking, respectively. Hence, vocalizations may be very important for the coordination of social organization in these monkeys. We 1) categorized golden-backed uakari vocalizations, 2) ascertained their behavioral context, and 3) investigated whether golden-backed uakari calls can encode information about the signaler. We observed the monkeys during 2 wet seasons in the flooded igapó forest of Jaú National Park, Brazil. We showed that golden-backed uakaris have 9 call types in their vocal repertoire, all distinguishable by ear and from analysis of spectrograms. Some calls, e.g., play-specific calls, were used only in particular behavioral contexts, and by individuals of specific age, whereas others were emitted under a range of situations. The structure of the loud tchó call varied among individuals, and according to behavioral context, i.e., whether individuals were foraging/feeding, traveling, or performing agonistic interactions. This knowledge of the species’ vocal repertoire is valuable for surveying the monkeys acoustically in habitats where visual surveys are difficult.     

Variability in spider monkeys' vocalizations may provide basis for individual recognition

Analysis of long-range vocalizations given by spider monkeys revealed consistent acoustic differences among the calls of individuals. Of seven acoustic measurements, four exhibited significant variation between individuals. A discriminant analysis demonstrated that two of these variables allowed correct identification of the caller 44% of the time. Including the remainder of the variables increased the percentage correctly identified to 50%. Individual identification by call structure could benefit spider monkeys, where individuals forage separately in subgroups and the interactions between specific pairs of individuals is highly variable. Acoustic recognition of callers would facilitate the choice of which subgroups to join, thus allowing individuals to manipulate the size and composition of their subgroups. In addition, the calls of mothers and offspring appeared to be similar in acoustic properties.     

Vocal behavior of captive Sichuan golden monkeys (Rhinopithecus r. roxellana)

Vocal behavior of four pairs of adult Sichuan golden monkeys was studied at the San Diego, Seattle, Los Angeles, and San Francisco zoos between November 1984 and October 1986. Four call types made up 86% (1,357/ 1,578) of vocalizations that occurred during 45 h of tape-recording. All four major vocalization types were made by both sexes. There was, however, significant sexual asymmetry in their production: 82% to 99% of each type was produced by one sex or the other. Frequently, duets developed when an individual male or female responded to the vocalizations of its cagemate. Males vocalized more than did females in each of the four pairs, and males responded vocally to female calls more than twice as frequently.     

Female bonobos use copulation calls as social signals

During mating events, females of many primate species produce loud and distinct vocalizations known as 'copulation calls'. The adaptive significance of these signals is considered to be in promoting the caller's direct reproductive success. Here, we investigated copulation calling in bonobos (Pan paniscus), a species in which females produce these vocalizations during sexual interactions with partners of both sexes. Females were more likely to call when mating with males than with females. We also observed a positive relationship between the likelihood of calling and partner rank, regardless of partner sex. Sexual activity generally increased with swelling size (an indicator of reproductive state) and, during their peak swelling, females called more with male than with female partners. Female bonobos are unusual among the non-human primates in terms of their heightened socio-sexuality. Our results suggest that in this species, copulation calls have undergone an evolutionary transition from a purely reproductive to a more general social function, reflecting the intrinsic evolutionary links between vocal behaviour and social cognition.     

Vocal signature in capybara,Hydrochoerus hydrochaeris

Social groups of capybaras are stable and cohesive. The species’ vocal communication is complex and mediates social interaction. The click call is emitted in a variety of contexts by animals from all age groups, but differs among groups; its attributed function is to keep contact among animals. To evaluate the presence of individual characteristics in the click call of capybaras, we recorded the vocalizations emitted spontaneously by six adults kept either solitary or in groups. We selected and measured the acoustic parameters of 300 click call phrases, 50 per individual. The parameters were submitted to a discriminant function analysis that revealed a classification accuracy of 76.8 %. A General Linear Model analysis revealed significant differences among the six individuals, and post hoc results showed that differences between a given pair were different from those of any other pair. The acoustic parameters that most contributed to discriminate the individual calls were click interval duration and click duration, suggesting that temporal parameters are more important than frequency parameters for individuals’ discrimination. The findings of individual characteristics in the click calls indicate that these vocalizations can be used as vocal signatures during social interactions.