Receivers respond differently to chick-a-dee calls varying in note composition in Carolina chickadees, Poecile carolinensis

The chick-a-dee call of the avian genus Poecile is a structurally complex vocal system because it possesses a set of simple rules that governs how the notes of the call are ordered, and variable numbers of each of the note types strung together can generate an extraordinary number of unique calls. Whereas it has been hypothesized that chick-a-dee calls with different notes may convey different information, no experimental evidence has been offered in support of the hypothesis. Previously published studies suggested that flock members use chick-a-dee calls in the context of moving to or from a feeding site. Here, we tested Carolina chickadees' responses to playbacks of chick-a-dee calls that differed in note composition. Playbacks were conducted in the field in the context of a novel food source. Our pilot data had indicated that chick-a-dee calls with relatively large numbers of ‘C’ notes were given by birds on their first contact with a novel seed stand. In the present study, we found that chickadees flew in close to the playback speaker and subsequently took seed from a seed stand more often during playbacks of chick-a-dee calls containing C notes than chick-a-dee calls not containing C notes or than control playbacks. Vocal responses of chickadees to the playbacks also differed in relation to the particular vocal signal being played back. These results indicate that receivers respond differently to chick-a-dee calls containing different compositions of note types and represent a first step to link variation in note composition and ordering in these calls to possible meanings.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved .     

Development of Early Vocalizations and the Chick-a-dee Call in the Black-capped Chickadee,Parus atricapillus

The chick-a-dee call of the black-capped chickadee (Parus atricapillus) is composed of discrete elements, or notes, that are combined to form hundreds of different calls. To investigate the development of this complex call, 12 families of color-marked chickadees were observed and recorded in the wild. Vocalizations were monitored for 18 d in the nest and 14–18 d postfledging. Most vocalizations of nestlings and fledglings were associated with feeding. At hatching, vocalizations consisted of a structurally simple note type that became more complex and variable with age. Around 9–12 d, the development of the call occurred, when single notes became organized into a multiple-note unit. Notes within the call differentiated into higher frequency, rapidly modulated initial note types and a lower frequency, moderately modulated terminal note type, features also present in adult chick-a-dee calls. Several adult-like calls including chick-a-dee calls, fee-bee songs, and a subsong-like vocalization developed prior to fledgling dispersal. Based on resemblances of note structure and general call structure, the chick-a-dee call appeared to develop from calls of nestlings and fledglings, although not necessarily in a chronologically linear progression. Some features of the chick-a-dee call closely resembled features of older nestling and fledgling calls, while other features more closely resembled the sounds of very young nestlings. Vocal development in the chickadee is compared with song and call development in other species, and the possible significance of acoustic resemblances between chick-a-dee calls and the food-associated calls of nestlings and fledglings is discussed.     

Constraints on the Structure of Combinatorial “Chick-a-dee” Calls

“Chick-a-dee” calls of the black-capped chickadee (Parus atricapillus) constitute a combinatorial system of animal communication, apparently the only such system yet described. Four note-types, which may be omitted or repeated a variable number of times, occur in a fixed sequence (A–B–C–D) to constitute calls. Quantitative analyses determining the nature of departures from first-order transitional probabilities between successive notes revealed a variety of results that point consistently to two underlying features of calls. (1) Some constraint operates to limit the length of calls, as manifest in shortening of repetition-strings and omitting of note-types expected to follow. (2) There is an opposing tendency to include at least one or two D-notes at the end of a call, regardless of the overall length. The second feature suggests important semantic properties of D-notes, especially the ratio of D s to other note-types and the flock-specific acoustical structure of D-notes. The constraint on length of call does not appear intimately related to semantic causes, but may instead be determined by the maximum duration of continuous phonation, in parallel with the fundamental breath-group of human speech. In this respect, a “chick-a-dee” call resembles an entire “natural sentence” of spoken human language.     

Chick-a-dee call variation in Carolina chickadees and recruiting flockmates to food

The "chick-a-dee" call of many Paridae species (titmice, tits,and chickadees) is structurally complex and functions in socialcohesion. Studies with different Parid species suggest thatvariation in the note composition of calls relates to a widevariety of contexts. An earlier study with Carolina chickadees(Poecile carolinensis), the focal species of the present study,found that receivers responded differently to playback callsdiffering in note composition in feeding contexts. Here, weaddressed whether signalers actually produce calls differingin note composition in feeding contexts and whether those callsmight serve a recruitment function. In a first study, we foundthat the first chickadee to take seed from a feeding stationproduced calls with a greater number of D notes before the secondchickadee arrived to take seed, compared with after the secondchickadee arrived to take seed. This suggests that calls witha large number of D notes might serve a general recruitmentfunction. We tested this idea in a second study, using playbacksof calls containing a large number of D notes or a small numberof D notes at different sites. We found that the latency fora first chickadee to come into a site and take seed was shorterfor playback variants containing a large number of D notes.Thus, in Carolina chickadees, chick-a-dee calls containing alarge number of D notes may function to recruit other flockmembers to a discovered food source.     

VOCAL ONTOGENY OF NESTLING AND FLEDGLING BLACK-CAPPED CHICKADEES POECILE ATRICAPILLA IN NATURAL POPULATIONS

ABSTRACT

We studied the vocal ontogeny of black-capped chickadees Poecile atricapilla from hatching through approximately age 40 days, at which time the juveniles dispersed from their hatching area. Development of three vocalisations (chick-a-dee call, fee-bee song, gargle call) was monitored by tape recording daily. Spectrographs quantification and comparison were carried out with Sound Analysis software. Developmental changes in offspring vocalisations were compared to the parent birds, allowing a measure of acoustic similarity between adults and young throughout early life. The chick-a-dee call developed all four of its component syllables (ABCD) from a single sound of day-old nestlings, but the four syllables emerged in adult form at different times in ontogeny. The fee-bee song was produced in impressively adult form starting at about age 20 days, with no precursor sounds to indicate gradual emergence. The principal difference between adult and juvenile fee-bee songs was the persistent production of three or four notes by juveniles rather than the adults' species-typical two notes. Gargle calls appear to develop from “subsong” strings of precursor gargle syllables. Juvenile gargles began to be identifiable at about age 33–35 days but did not match any of the gargle calls of the local adult population. Post-dispersal juveniles may develop gargles that match the gargle types of local birds where they settle.     

Flexibility as a Strategy for Avoiding Call Overlap in Male Anhui Treefrogs

Male-male vocal competition is critical for mating success in anuran species; however, it remains unknown that how males regulate their competitive strategies dynamically during competition because calling is highly time-consuming, energetically demanding and likely to increase predation risks. Since different parts of calls will encode different information for vocal communication, we hypothesized that competitive strategies of male frogs may be modulated by the temporal and spectral features of different call notes. To test this hypothesis, the natural advertisement calls(OC), its modified versions with the first call note replaced by white noise(WN) or other notes and with the fifth call note replaced by WN, were played back to the Anhui tree frogs(Rhacophorus zhoukaiyae). Results showed that 1) males produced more competitive calls in response to acoustic stimuli compared to their baseline calling during silence; and 2) males emitted more non-overlapping calls compared to overlapping calls in response to the acoustic stimuli. These results are consistent with the idea that males are flexible to acoustic signals and their competition strategies are modulated dynamically by social contexts.     

Microgeographic Variation and Sharing of the Gargle Vocalization and Its Component Syllables in Black-Capped Chickadee (Aves, Paridae, Poecile atricapillus) Populations

Throughout the year during agonistic encounters, black‐capped chickadees (Poecile atricapillus) emit a vocal signal known as the gargle call. Each bird has a repertoire of structurally differing gargle calls; some are shared with others in the local area. As a basis for understanding the cultural evolution of this social signal, we initiated a study of gargle call repertoires of birds living in a narrow belt of continuous riparian habitat occupied throughout by a resident population of chickadees. During two consecutive winter seasons, we sampled repertoires at three locations over a distance of 8.4 km to quantify micro‐geographical variation. Analyses of vocal sharing and population differentiation were carried out on whole gargle calls and on the individual acoustic units (syllables) from which the whole calls are constructed. We analysed 28 380 calls of 46 subjects in the two seasons of study. Birds averaged 7.6 different calls in their gargle repertoires. Calls were composed of about 10 syllables on average. Fifty‐six different syllables were used to construct the calls of all birds. Each study site had some gargle calls unique to the local birds and some that were shared with one or both of the other two sites. There was significantly greater sharing of both calls and syllables among birds within sample sites than between sample sites. The frequencies of the different kinds of gargles and syllables were significantly correlated across the 2 yr of the study, but the correlation was stronger (r² = 0.93) for syllables than for whole gargle calls (r² = 0.61).     

Call Learning in Black-capped Chickadees (Parus atricapillus): The Role of Experience in the Development of ‘Chick-A-Dee’ Calls

The role of learning in the development of bird vocalizations other than territorial song is not well studied. The well-known role of direct imitation in the development of territorial song potentially masks the effects of other processes in the development of vocal behaviour. The ‘chick-a-dee’ call of black-capped chickadees is a good system in which to investigate more subtle developmental processes because this call is composed of a small number of distinctive note types. These note types may be classified objectively based on a simple set of acoustic variables, allowing for a quantitative assessment of vocal learning. We raised four groups of black-capped chickadees under different degrees of social and acoustic isolation. We then used a multivariate analysis of the acoustic structure of the introductory call notes (‘A-’ ‘B-’ and ‘C-notes’) to determine how similar the notes produced by these hand-reared birds were to the notes of wild birds. Hand-reared chickadees with greater exposure to normal phonology produced notes of all three note types that were more similar to those of wild birds. Regardless of experience, however, all birds produced A-notes that fell within the normal range of those produced by wild birds. By contrast, the development of normal B- and C-notes appears to be more dependent upon experience. These data suggest that learning may play a different role in the development of different phonological units within one vocalization. Our results also illustrate the importance of considering processes other than simple imitation in the development of avian vocalizations.     

Vocal communication in a neotropical treefrog, Hyla ebraccata: Advertisement calls

We studied the vocal communication of Hyla ebraccata in central Panama. The advertisement call of this species consists of a pulsed buzz-like primary note which may be given alone or followed by 1–4 secondary click notes. Primary notes are highly stereotyped, showing little variation within or0 among individuals in dominant frequency, duration, pulse repetition rate or rise time. Males calling in isolation give mostly single-note calls. They respond to playbacks of conspecific calls by increasing calling rates and the proportion of multi-note calls, and by giving synchronized calls 140–200 ms after the stimulus begins. Responses to conspecific advertisement calls are usually given immediately after the primary note of the leading call, but the primary note of the response often overlaps with the click notes of the leading call. Experiments with synthetic signals showed that males synchronize to any type of sound of the appropriate frequency (3 kHz), regardless of the fine structure of the stimulus. Playbacks of synthetic calls of variable duration showed that males do not synchronize well to calls less than 150 ms long, but they do to longer calls (200–600 ms). The variance in response latency increased with increasing stimulus duration, but modal response times remained at around 140–200 ms. Similar results were obtained in experiments withsynthetic calls having a variable number of click notes. Males showed no tendency to increase the number of click notes in their calls in response to increasing stimulus duration or increasing number of clicks in the stimulus. Females preferred three-note to one-note calls in two-choice playback experiments, whether these were presented in alternation, or with the one-note call leading and the three-note call following. Females showed no preference for leader or follower calls when both were one-note. When two-note calls were presented with the primary note of the follower overlapping the click note of the leader, females went to calls in which click notes were not obscured. Our results indicate that male H. ebraccata respond to other males in a chorus in ways which enhance their ability to attract mates.     

The first call note of the Anhui tree frog (Rhacophorus zhoukaiya) is acoustically suited for enabling individual recognition

The effective production of acoustic signals is critically important for intraspecific communication in vocal animals; however, it is also highly time-consuming, energetically demanding and likely to increase predation risks. Thus, we hypothesized that the biological significance of each component of complex acoustic signals would differ serving specific functions and that the first component of such signals would be most important for social signalling and exhibit unique acoustic characteristics because of the precedence effect. To test this hypothesis, we measured temporal and spectral acoustic parameters for each note in the advertisement calls of the Anhui tree frog (Rhacophorus zhoukaiya), a species in which males build mud burrows and call from within these nests. Multivariate analyses including hierarchical cluster analysis and multidimensional scaling were used, based on temporal and spectral acoustic parameters for each of 10 notes/call. These results show that the first call notes form one cluster while the other notes form a second cluster in multidimensional space when the parameters measured were normalized. Furthermore, the temporal and spectral sound attributes of the first call note provide sufficient information for discrimination between different individuals. Moreover, discriminant analysis showed that the fundamental frequency of the first note is sufficient to identify individuals when the data are not normalized. Taken together, these results indicate that the first call note is poised to play an important role in Anhui tree frog vocal communication insofar as the temporal and spectral features provide sufficient information for individual recognition.     

All "chick-a-dee" calls are not created equally. Part II. Mechanisms for discrimination by sympatric and allopatric chickadees

The 'chick-a-dee' call, common to all members of the genus Poecile, is used by both sexes throughout the year to putatively co-ordinate flock movements and register alarm. In some regions, two or more chickadee species occupy overlapping territories, and therefore it is essential that these sympatric species learn to discriminate between the acoustically similar calls of the species. Previous work from our laboratory has shown that black-capped (P. atricapillus) and mountain chickadees (P. gambeli) discriminate between the species' calls and treat each species' calls as belonging to separate open-ended categories. In the current set of experiments we use an operant conditioning paradigm to gain an understanding of (1) how the birds perform this discrimination and (2) whether birds with different levels of experience with heterospecific calls perform this task differently. We use natural recordings of chick-a-dee calls and perform several manipulations to test the importance of the introductory 'chick-a' portion and the terminal 'dee' portion for discriminating among the calls of the two species. Evidence suggests that birds mainly use the terminal 'dee' portion, as all groups of birds responded similarly to these probe stimuli and control chick-a-dee calls. We propose that the terminal 'dee' portion, consisting of lower frequency notes, is more likely to be resistant to degradation, and therefore a more reliable species-specific marker.     

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.     

The role of frequency modulation in controlling the response of mallard ducklings (Anas platyrhynchos) to conspecific distress calls

Three experiments examined the response of mallard ducklings to conspecific distress calls. In experiment 1, a synthetic call was constructed from a single distress note, by recording it on a digital disc and then using a software routine to regularly repeat this stored note with the average period (267 ms) of the original call. Ducklings were then tested for their tendency to inhibit their own distress vocalizations in response to either this synthetic call, the original call, or a constant-frequency tone mimic; they showed a significantly stronger inhibitory response to the synthetic call than to the tone mimic, but an even stronger response to the original call. The former result indicated that the synthetic call was an effective stimulus, and suggested that some aspect of the frequency modulation found in distress notes is required to evoke the normal duckling inhibitory response. The latter result further suggested that the acoustic variability found in the original call, but not the synthetic call, may be of importance for controlling duckling behaviour. Experiment 2 demonstrated that several other minor differences between the synthetic call and original call (number of notes per call and the digital recording of the synthetic call) could not account for the difference in duckling response to these two calls. Finally, in experiment 3, two additional synthetic stimuli were constructed from the digitized note, by first excising either the initial (front-chop) or terminal (rear-chop) frequency modulation found in each distress note, then creating amplitude envelopes for these new notes similar to that of the unaltered note, and finally repeating these notes to form calls with the same note period as the original call. The ducklings tested with the synthetic normal-note call and synthetic front-chop call showed a significantly stronger inhibitory response than the ducklings tested with the synthetic rear-chop call and the tone mimic. These results indicate that the terminal descending frequency sweep is an important feature of distress notes for triggering the response of ducklings to conspecific distress calls.     

Advertisement calls of Guenther’s frog Hylarana guentheri (Anura: Ranidae) during the breeding season

Acoustic signalling is the most important form of communication in anuran amphibians. Here we recorded and analysed the calls of 18 male Guenther’s frogs (Hylarana guentheri) from the wild during the breeding season. The advertisement calls of H. guentheri were composed of from a single note to five notes, with three-note calls the most recorded. All individuals produced calls around 600 Hz but calls ranged from 470 to 2600 Hz. Comparing the differences between individuals calls, we found within-male coefficients of variation (CV_w) of call intensity, the fundamental frequency, the first formant, the second formant, the third formant and the fourth formant were static (less than 5% variation), whereas those of note duration, call duration, call interval, numbers of pulses and dominant frequency were dynamic (larger than 15% variation). Comparisons of the call characteristics of H. guentheri in this study with other studies from China, Singapore and Vietnam found call characteristics varied greatly between the five different locations.     

Geographic Variation in Note Composition and Use of chick‐a‐dee Calls of Carolina Chickadees (Poecile carolinensis)

The aim of this study was to determine whether geographic variation exists in the composition of note types in the chick‐a‐dee call of Carolina chickadees. This determination is of interest for two reasons: earlier studies with a related species suggested minimal geographic variation in note composition, and geographic variation in social signals may represent important developmental or selection processes shaping signal use. Carolina chickadees were recorded in a naturalistic observation study in west‐central Indiana. Chick‐a‐dee calls were analyzed and compared to calls from an eastern Tennessee population that had been described in a previously published study (Auk, 125, 2008, 896). Despite much similarity in the basic rules by which notes are organized to compose calls, there were several significant differences in how calls of the two populations were structured. Furthermore, birds from Indiana used their chick‐a‐dee calls in certain contexts in different ways compared to birds from Tennessee. These findings suggest interesting population‐level variation in this call system, and future research should be able to determine whether these differences are driven by evolutionary, ecological, or developmental factors, or some combination of these factors.     

The function of food-associated calls in white-faced capuchin monkeys, Cebus capucinus, from the perspective of the signaller

In many species, call recipients respond to food-associated calls by approaching the signaller. For this reason, most studies of food-associated calls focus on the benefits to a signaller of attracting a particular audience to a food source. Although call recipients respond as if they have been informed about the location of a food source, it is not necessarily the case that the primary function of food-associated calls is to inform others. I combined naturalistic observations and food placement experiments to investigate the environmental and social influences on call production in white-faced capuchin monkeys to assess other possible functions of food-associated calls. Individuals did not call under the circumstances predicted by an information-sharing hypothesis. The quantity of food and the age-sex composition of the audience did not influence call production, but food type did. Individuals produced more food-associated calls when they discovered fruit compared with insects or eggs. Results of observations of social interactions after food discovery indicated another possible function of food-associated calls. Individuals who called when they discovered food were less likely to be approached by others who were in visual contact than individuals who remained silent. Individuals who did not call when they discovered food were more likely to call subsequently if a higher-ranking, as opposed to a lower-ranking, animal approached them. Furthermore, individuals who called when approached by higher-ranking animals were less likely to receive aggression than individuals who did not call. Therefore, food-associated calls may function to announce food ownership, thereby decreasing aggression from other individuals.     

鬼鸮甘肃亚种繁殖期叫声研究

利用叫声回放和声谱分析对莲花山自然保护区鬼甘肃亚种(Aegolius funereus beickianus)繁殖期的叫声进行了研究。共记录了领域叫声、尖叫声等6种叫声,分析了各种叫声的特征及与行为的联系。发现不同地点录制的领域叫声存在差异,但同一个体的叫声也有变化。甘肃亚种的领域叫声和欧洲的指名亚种A.f.fu-nereus及北美的亚种A.f.richardsoni相比较,单音数量少,单音长度小,频率高,但差异不大。     

沼水蛙繁殖期鸣声特征及鸣叫节律

鸣叫对无尾两栖类动物的生存与繁殖起重要作用。蛙类的鸣叫行为受到环境因素影响表现出一定的节律性。2016年8和9月,采用录音机和指向性话筒,在野外录制了57只沼水蛙（Hylarana guentheri）的鸣声并对其鸣声特征进行分析;通过悬挂录音笔和自动温湿度记录仪研究了沼水蛙鸣叫节律（17 d）及其与环境温度、相对湿度的关系。结果显示,沼水蛙的鸣声由1 ~ 4个音节组成,不同类型鸣声间的音节主频、音节时长存在显著差异（P < 0.05）。该物种全天具有鸣叫行为,13:00 ~ 14:00时为鸣叫高峰期。白天单音节鸣声、双音节鸣声、三音节鸣声、总鸣声和总音节的数量较夜晚显著增加（P < 0.01）。鸣声数量和音节数量均与环境温度呈正相关（P < 0.01）。结果表明,沼水蛙通过改变音节数量、音节主频和音节时长改变鸣叫策略。沼水蛙的鸣叫行为具有昼夜节律性且受环境温度的影响。     

The versatility of graded acoustic measures in classification of predation threats by the tufted titmouse Baeolophus bicolor: exploring a mixed framework for threat communication

Many mammal and bird species respond to predator encounters with alarm vocalizations that generate risk‐appropriate responses in listeners. Two conceptual frameworks are typically applied to the information encoded in alarm calls and to associated anti‐predator behaviors. ‘Functionally referential’ alarm systems encode nominal classes or categories of risk in distinct call types that refer to distinct predation‐risk situations. ‘Risk‐based’ alarms encode graded or ranked threat‐levels by varying the production patterns of the same call types as the urgency of predation threat changes. Recent work suggests that viewing alarm‐response interactions as either referential or risk‐based may oversimplify how animals use information in decision‐making. Specifically, we explore whether graded alarm cues may be useful in classifying risks, supporting a referential decision‐making framework. We presented predator (hawk, owl, cat, snake) and control treatments to captive adult tufted titmice Baeolophus bicolor and recorded their vocalizations, which included ‘chick‐a‐dee’ mobbing calls (composed of chick and D notes), ‘seet’ notes, two types of contact notes (‘chip’, ‘chink’), and song. No single call type was uniquely associated with any treatment and the majority of acoustic measures varied significantly among treatments (46 of 60). The strongest models (ANOVA and classification tree analysis) grouped hawk with cat and owl, and control with snake, and were based on the number or proportion of a) chick and D notes per chick‐a‐dee call, b) chip versus chink notes produced following treatment exposure, and c) the frequency metrics of other note types. We conclude that (1) the predation‐threat information available in complex titmouse alarm calls was largely encoded in graded acoustic measures that were (2) numerous and variable across treatments and (3) could be used singly or in combinations for either ranking or classification of threats. We call attention to the potential use of mixed threat identification strategies, where risk‐based signal information may be used in referential decision‐making contexts.     

Sex Differences in the Song of <Emphasis Type="Italic">Indri indri</Emphasis>

In some primate species, males and females within a social group emit loud calls in a coordinated manner or chorus. Indri indri emits a very conspicuous loud call that elicits the loud calls of neighboring groups. Previous investigations have hypothesized that the main functions of the indri chorus are related to territorial announcement, intergroup avoidance, and group cohesion. We investigated sex differences in indri song. We recorded and analysed songs given by 10 different groups over 160 d. Overall singing duration did not vary between the sexes. However, males emitted significantly fewer but longer notes. Adult males and females of each group participated in the song with sex-specific repertoires. Females had a song repertoire of 8 note types; males shared all of their 6 notes with females. Apart from the initial roars, in all note types shared by both sexes, male notes were significantly longer than female ones, whereas variations in frequency parameters differed according to the note type. These findings suggest that indri song may provide cues to conspecifics, such as group size and sex composition, which could influence interactions between groups.