VOCAL AND OTHER BEHAVIOUR OF KENYAN BLACK-COLLARED BARBETS LYBIUS TORQUATUS

We describe 12 visual and six acoustical displays of Black-collared Barbets, all associated with interactions of these sexually monomorphic barbets in greeting ceremonies and in antiphonal duets. Certain of the displays, e. g., bill directing, smooth posture and chatter calls largely are associated with multiple-barbet interactions and greeting ceremonies involving more than two barbets, and seem to represent purely agonistic displays. Other displays, most notably the bobbing and floating flight displays, and the tyaw call, are associated with preduet greeting ceremonies involving the presumed pair. In every case duets were performed by a presumed pair, following a greeting ceremony and visual displays accompany the duetting, which usually becomes precisely synchronized after the initial few notes. Four to 17 notes of each of two types, a usually lower pitched, short and mainly double type A note that introduces most duets, and a higher pitched, usually single, clear type B note that often terminates the duet. Variation in the notes and in their tempos within duets suggests no clearly defined temporal indicators for achieving, maintaining, and concluding the duets. The A and B notes cannot yet be assigned to one or other sex and similarity in form and development of the notes, from notes of greeting ceremonies, suggests the possibility that under some circumstances a bird of one sex may be able to deliver notes of either type. The duets are delivered from singing posts within the group's (or pair's) territory and usually only one duet is uttered from a site before the birds shift to another singing post. The duetting is compared with that of other barbets and appears to be among the best synchronized and most precise in the family. Comparisons with non-barbet duetting species are difficult because of the special habits and ecology of the barbets and uncertainty as to the sexual rôle of the duetters.     

Male search behavior of the stonefly,Pteronarcella badia (hagen) (Plecoptera: Pteronarcyidae), in relation to drumming

The mating system of many northern hemisphere stoneflies involves vibrational duetting, yet searching behavior in relation to the percussive communication has remained unreported. The communication-search system of a Colorado population of the ubiquitous western stonefly Pteronarcella badiain an experimental arena entails (1) a ranging search by calling virgin or polygynous males until duet establishment with virgin females, (2) continued intermittent duetting while the male engages in a local search and the female remains stationary, and (3) a tactile-based find of the female, followed by immediate mounting and mating. The average 234-s find time for pairs engaging in strong, continuous duets was significantly shorter than those for nonduetting or anomalously duetting pairs. Males of strongly duetting pairs made significantly more turns toward the female during search than in anomalously duetting pairs, and their local search pattern appears to be a triangulation aided by resource-specific (female) vibrational cues. Males displayed a wide range of fitness, based on searching time, and an increased number of duets significantly reduced finding time. Potential female selection and possible extrapolation of these experimental results to natural field conditions are discussed.     

Context-dependent functions of avian duets revealed by microphone-array recordings and multispeaker playback

In many tropical animals, male and female breeding partners combine their songs to produce vocal duets [1-5]. Duets are often so highly coordinated that human listeners mistake them for the songs of a single animal [6]. Behavioral ecologists rank duets among the most complex vocal performances in the animal kingdom [7, 8]. Despite much research, the evolutionary significance of duets remains elusive [9], in part because many duetting animals live in tropical habitats where dense vegetation makes behavioral observation difficult or impossible. Here, we evaluate the duetting behavior of rufous-and-white wrens (Thryothorus rufalbus) in the humid forests of Costa Rica. We employ two innovative technical approaches to study duetting behavior: an eight-microphone acoustic location system capable of triangulating animals' positions on the basis of recordings of their vocalizations [10] and dual-speaker playback capable of simulating duets in a spatially realistic manner [11]. Our analyses provide the first detailed spatial information on duetting in both a natural context and during confrontations with rivals. We demonstrate that birds perform duets across highly variable distances, that birds approach their partner after performing duets, and that duets of rivals induce aggressive, sex-specific responses. We conclude that duets serve distinct functions in aggressive and nonaggressive contexts.     

Sexually Dimorphic Intrasexual Duetting in an Otherwise Monomorphic Green Lacewing (Neuroptera, Chrysopidae, Chrysoperla plorabunda): Sexual Selection or Sex Recognition?

Songs produced during heterosexual duets in a green lacewing, C. plorabunda, are sexually monomorphic. However, individuals of either sex will also engage in intrasexual duets, which can exhibit sexual dimorphism. We confined males and females together in various combinations in a small arena to study the phenotypes, behavioral interactions, and functional roles of duetting in this species. The goal was to test whether sexual selection or sex recognition provided the better explanation of song sexual dimorphism. We determined that the monomorphic form of the intrasexual duet was long and stable, and could take place either between males or between females. Such “standard” intrasexual duetting songs were acoustically indistinguishable from heterosexual songs. However, males could also engage other males in special “fast duets” that sped up and terminated abruptly. Equivalent fast duets were not part of the female repertory. Fast duetting songs between males differed significantly from other types of male or female duetting songs in every measurable characteristic, but their role in the mating system was ambiguous. Contrary to one prediction of the sexual selection hypothesis, fast duetting between males occurred less often in situations where it might be the most useful to males in securing mates, i.e., during male-male-female interactions (trios). In addition, fast songs that started, ended, both started and ended, or neither started nor ended duets were acoustically indistinguishable, making it unlikely that females were choosing males based on such variation. However, songs that “both started and ended” fast duets were associated with a significant mating advantage, indicating a possible role for fast duetting in male-male sexual competition. Because the alternative hypothesis of sex recognition was also supported by some of our results, we conclude that aggressive qualities of male-male fast duets probably mediate intrasexual selection, while their increasing tempo serves as an adaptive response to promote rapid sex recognition by truncating unproductive and potentially dangerous intrasexual duetting.     

Differentiating the evolution of female song and male–female duets in the New World blackbirds: Can tropical natural history traits explain duet evolution?

Female bird song and combined vocal duets of mated pairs are both frequently associated with tropical, monogamous, sedentary natural histories. Little is known, however, about what selects for duetting behavior versus female song. Female song likely preceded duet evolution and could drive apparent relationships between duets and these natural histories. We compared the evolution of female song and male–female duets in the New World blackbirds (Icteridae) by investigating patterns of gains and losses of both traits and their relationships with breeding latitude, mating system, nesting pattern, and migratory behavior. We found that duets evolved only in lineages in which female song was likely ancestral. Both female song and duets were correlated with tropical breeding, social monogamy, territorial nesting, and sedentary behavior when all taxa were included; however, correlations between duets and these natural history traits disappeared when comparisons were limited to taxa with female song. Also, likelihood values supported stronger relationships between the natural history traits and female song than between these traits and duets. Our results suggest that the natural histories thought to favor the evolution of duetting may in fact be associated with female song and that additional selection pressures are responsible for the evolution of duets.     

Duets defend mates in a suboscine passerine, the warbling antbird (Hypocnemis cantator)

Despite the widespread occurrence of avian duets, their adaptivesignificance is poorly understood. It is generally assumed thatthey function in the joint defense of territories, but no studyhas successfully distinguished between this hypothesis, whichinvokes cooperation between the sexes, and mate defense, whichinvokes conflict. Further, most duetting studies have focusedon oscine passerines, the songs of which are learnt and relativelycomplex. We therefore tested the mate defense hypothesis inthe warbling antbird (Hypocnemis cantator), an Amazonian suboscinethat produces simple sex-specific songs and duets. Acousticanalysis of songs showed (1) that solos were often producedby males, but rarely by females; (2) that duets consisted ofa male song and a female reply; and (3) that, although femalesong was invariable, a swift reply resulted in males producingshorter songs with fewer notes. These results suggest that duetting,and the structure of duets, is chiefly a product of female behavior,a scenario more suggestive of conflict than cooperation. Toinvestigate this idea we carried out playback experiments, whichshowed that (4) the response to solo songs was sex specific(i.e., male solos elicited a strong response from paired males,and female solos elicited a strong response from paired females);(5) males and females responded to same-sex solos more stronglythan to duets; and that (6) females answered their partner'ssongs more often, and more rapidly, in response to female solosthan male solos or duets. Although it can be argued that sex-specificresponses to solo song result from intrasexual territorial defense,we cannot use the same reasoning to explain (5) or (6). Instead,these observations imply that solitary intruders were more threateningthan paired intruders, and thus that the perceived threat wasto the partnership rather than the territory. Taken together,findings (1) to (6) suggest that females adjust their vocalbehavior in relation to the level of perceived threat to thepartnership, and duet with males in order to repel same-sexrivals. This study therefore strengthens support for the matedefense hypothesis, and suggests that conflict—ratherthan cooperation—may have played a major role in the evolutionand maintenance of avian duets.     

Song repertoire and duetting behaviour of the tropical boubou, Laniarius aethiopicus

Female song and the significance of duetting in birds remain puzzling, in part because of the limited number of species studied. We investigated duetting behaviour in the tropical boubou in West Africa. Birds produced a diverse song repertoire consisting of solos, duets and trios, with duets being the most conspicuous vocalization. We identified 12 distinct duet types in which tonal and broadband notes were combined by males and females in highly synchronized and temporally precise patterns. Molecular sexing and observations of colour-banded birds revealed that duets were initiated by both sexes, with strict sex-specific roles maintained within the duet. An equal number of duet types was initiated by males and females. Seven of 12 duets were terminated by the male. Male-initiated duets were also more common (89% of total) and were repeated more often, i.e. sung more frequently. Solo singing occurred when partners did not respond and was also used by unpaired birds. Trios were produced by subadult birds joining the duet of a resident pair. The wide variety of contexts in which duets were sung suggests that they serve several functions, including territorial defence and mutual mate guarding.     

Functions of duetting in the tropical boubou, Laniarius aethiopicus: territorial defence and mutual mate guarding

Duetting, found in many animal taxa, is still a poorly understood form of signalling behaviour despite numerous hypotheses to explain its function. One contentious issue is whether duetting is a cooperative endeavour or signals conflict between the sexes. We studied the function of duetting in the tropical boubou, using interactive playback experiments. We staged encounters by presenting paired birds with four variants of each of four duet types (neighbour and stranger solos and neighbour and stranger duets). Multiple lines of evidence suggest that both joint territorial defence and mutual mate guarding are important functions of duetting, and that both cooperation and conflict between males and females have shaped duetting behaviour. Of a repertoire of 12 duet types, eight were used for joint territorial defence. Two of these eight duet types were probably also used for mutual mate guarding, suggesting that individual duets can have multiple functions. We found an unusual response for duetting birds in that females often synchronized notes with male solo playback to form precise duets. In turn, males attempted to jam the song of rivals when females joined the song of male solo playback, providing further evidence for the importance of acoustic mate guarding. Song jamming by males has not previously been described for duetting species. Finally, we suggest that the repertoire of duet types is used to fine-tune territorial encounters in a network environment of multiple interactants.     

Description of the acoustical interaction and synchronization between duetters of the Large-footed Finch (Pezopetes capitalis)

Duets are highly coordinated acoustical displays produced by two individuals. Studying the structure of duet songs and its causes is essential to understand the ecological role and evolution of this form of communication. Our goals in this study were to describe the structure of the duet of the Large-footed Finch, the temporal synchronization of each individual, and to test the effect of provoked (playback) duets. We recorded birds at Cerro de la Muerte, Costa Rica and simulated territory intrusions using playback trials. The duet of this species consisted of overlapping notes between both members of a mated pair, which were emitted either randomly or periodically. Temporal synchronization was similar between both individuals in a mated pair. Both partners adjust the silent intervals between notes in response to the duration of its partner’s note. The high frequency of the second part of the duet of this species decreased with time and the second individual, in duetting this part, not only synchronized its notes temporally with the first individual, but also synchronized the high frequency. The high frequency of the first note the second individual sung was a bit lower than the previous note sung by the first individual. Finally, birds responding to provoked duets produced duets with higher frequency and higher rate of notes compared to unprovoked duets. Our results provide information on the temporal and frequency synchronization in duets of an endemic emberizid and provide new information about a less reported vocal behaviour in bird duets, the acoustic frequency coordination.     

Vocal duetting by a mated pair of coral-billed ground-cuckoos (Carpococcyx renauldi) at the Metro Toronto Zoo

During a behavioral study, I recorded vocal duetting by a mated pair of Coral-billed Ground-cuckoos (Carpococcyx renauldi) at the Metro Toronto Zoo. The birds produced three different call types that were combined into four distinct simultaneous and antiphonal duetted sequences. Duetting bouts (n = 12) were always initiated by the male and lasted 1–15 min, although they were generally 2–5 min in length. During longer bouts, both male and female duet elements changed frequently. The loud, clear antiphonal duets were likely associated with long distance communication and could function in territorial defense. These calls were most frequently given as the cuckoos moved around the exhibit, and were often associated with interspecific aggression. Quiet duet sequences were generally uttered when the cuckoos were in close proximity to one another, often from an exposed perch. Close perching and concurrent duetting by this mated pair may also function in spatial defense since it could advertise a coordinated alliance in territorial defense to potential intruders. Zoo Biol 16:179–186, 1997. © 1997 Wiley-Liss, Inc.     

MINUTES OF THE FORTY-SEVENTH ANNUAL GENERAL MEETING OF THE SOUTH AFRICAN ORNITHOLOGICAL SOCIETY HELD AT JOHANNESBURG AT 2.00 P.M. ON SATURDAY 19 JUNE 1976

Many duetting species are monomorphic and the suggestion has recently been made that duets may be female-led more often than has been supposed in the past. Observations were carried out on several marked individuals of two monomorphic duetting species, the Moho, Hypergerus atriceps, and Yellow-crowned Gonolek, Laniarius barbarus, in The Gambia. DNA analysis from blood samples revealed that in each case the male was the pair member that normally led the duet. It is suggested that duets of this sort may arise where it benefits a male to attract more than one female, but it is to the female's disadvantage that he should do so.     

Function of pair duets in the eastern whipbird: cooperative defense or sexual conflict?

Paired male and female eastern whipbirds, Psophodes olivaceus,sing precisely coordinated, male-led duets. Four broad explanationshave been proposed for the function of duets: 1) cooperativeresource defense, 2) prevention of partner usurpation, 3) defenseof an individual's own position within the partnership, or 4)mate identification and localization. These 4 hypotheses makedifferent predictions about how male and female residents shouldrespond to simulated intrusion by other pairs or individuals.We compared the behavioral and vocal responses of 20 pairs ofeastern whipbirds to simulated territorial intrusions by: 1)a solitary singing male, 2) a solitary singing female, and 3)a duetting pair. Males and females did not coordinate theirapproach to the playback speaker and showed sex-specific responsesto playback. Males did not respond differently to duetting versussolo singing intruders. By contrast, females approached moreclosely during solo female song than during solo male song orduet playback. Females also produced specific vocalizationsonly in response to duet and solo female playback. Both sexesapproached the speaker more closely and quickly during playbackof same-sex solo songs than opposite-sex solo songs. Finally,females answered more of their mate's songs during simulatedintrusion by a lone female than during simulated intrusion bya lone male. Our results suggest that duets in this speciesprimarily function to allow females to defend their exclusiveposition in a partnership. Mate defense by females is unusualin birds but may be promoted in eastern whipbirds by a female-biasedsex ratio and the need for exclusive access to male care. Thus,duets result from independent and conflicting strategies ofmate and territory defense in males and females.     

Migration and the evolution of duetting in songbirds

Many groups of animals defend shared resources with coordinated signals. The best-studied of these signals are the vocal duets produced by mated pairs of birds. Duets are believed to be more common among tropical-breeding species, but a comprehensive test of this hypothesis is lacking, and the mechanisms that generate latitudinal patterns in duetting are not known. We used a stratified sample of 372 songbird species to conduct the first broad-scale, phylogenetically explicit analysis of duet evolution. We found that duetting evolves in association with the absence of migration, but not with sexual monochromatism or tropical breeding. We conclude that the evolution of migration exerts a major influence on the evolution of duetting. The perceived association between tropical breeding and duetting may be a by-product of the migration–duetting relationship. Migration reduces the average duration of partnerships, potentially reducing the benefits of cooperative behaviour, including duetting. Ultimately, the evolution of coordinated resource-defence signals in songbirds may be driven by ecological conditions that favour sedentary lifestyles and social stability.     

Acoustic mimicry and disruptive alternative calling tactics in an Australian bushcricket (Caedicia; Phaneropterinae; Tettigoniidae; Orthoptera): does mating influence male calling tactic?

Abstract.  Male calling and searching tactics are described for a duetting Australian bushcricket, Caedicia sp. 12 (Phaneropterinae; Tettigoniidae; Orthoptera). The repertoire of Caedicia sp. 12 consists of the calling song and, by nonduetting males, a series of calling tactics that include short-click calling, disruptive over-singing and a call mimicking the entire duet. Nonduetting males respond to the production of a duet by another male and a female with short-click calls that mimic the female call at the conclusion of a duet. By manipulating the male's mating history, it is found that this form of calling behaviour is more likely to occur within the male's 6-day postmating refractory period; the low cost tactic allows males to re-mate during spermatophore replenishment. Males also produce disruptive calls in response to a duet, where the male may over-sing the duetting male's signal or produce a call that appears to mimic the entire duet; the male produces a calling song followed by a short signal that has the same latency as the female's reply within a duet. Males also over-sing crucial elements of the duetting-male's song that are normally critical for the female's conspecific recognition. There is no evidence that females search for the duetting male partner, but males unable to enter a duet will search for the call of a responding female. Searching by males is more common when these males are producing disruptive calls. Alternative male calling tactics are discussed as a set of conditional strategies for securing unmated females.     

Partners coordinate territorial defense against simulated intruders in a duetting ovenbird

Duets in breeding pairs may reflect a situation of conflict, whereby an individual answers its partner's song as a form of unilateral acoustic mate guarding or, alternatively, it may reflect cooperation, when individuals share in territory defense or safeguard the partnership. The degree of coordination between the sexes when responding to solo versus paired intruders may elucidate the function of songs in duets. We examined this issue in a study with rufous horneros (Furnarius rufus), a duetting, socially monogamous Neotropical species with low levels of extrapair paternity. We exposed social pairs during the nonbreeding season to playbacks of duets, male solos, female solos, and control heterospecific songs. Partners approached all conspecific stimuli together and responded by singing quickly, at higher rates and by coordinating ~80% of their songs into duets. For both sexes, most response variables (seven of nine) did not vary across conspecific treatments. These results suggest that partners duet and coordinate behaviors to cooperatively defend common territories. However, females spent more time in territorial vigilance, and partners were highly coordinated (correlated responses) in response to duets and female solos in comparison with male solos. This indicates that female intrusions (paired or solo) might be more threatening than male intrusions in the nonbreeding season, especially for territorial females, and that females are less cooperative with their partners in territory defense against male intruders.     

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.     

DUETTING AND VOCAL RECOGNITION BY ALDABRA WHITE-THROATED RAILS DRYOLIMNAS CUVIERI ALDABRANUS

Sonagrams of duets and solo song of the Aldabra White-throated Rail are presented and their behavioural contexts and functions noted. Visual inspection of sonagrams suggests that the structure of elements of high intensity song is individually characteristic. Elements of low intensity song are characteristic for some but not all individuals. The elements of medium intensity song are the most variable within individuals, and it is least likely that these could permit individual recognition. However, it is noted that the temporal pattern of delivery of song elements may provide cues for vocal recognition.
Both male and female Aldabra Rails respond to the playback of their mate's song by either approaching the sound source or vocalizing. A greater number of positive responses was given to playbacks of the mate's song than to playbacks of a song of a strange bird of the same sex as the mate. It is suggested that this difference in responsiveness may in part result from a more rapid habituation to unfamiliar song.
The function of duetting in the Aldabra Rail and other tropical rails is discussed. It is considered that the major function of duetting in the Aldabra Rail may be co-operative territorial defence. The extent and precision of duetting in the Rallidae is considered. It is concluded that precise duetting has not yet been demonstrated in any rail species and that the rates of calling in duets may be internally derived.     

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.     

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.     

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.