Duet codes do not enhance neighbour recognition in two closely related species of duetting neotropical wrens

Numerous studies have shown that territorial animals exhibit less aggression in response to neighbours than to strangers, a phenomenon known as dear enemy effect. The influence of acoustic features, such as song type sharing and repertoire sizes, in neighbour recognition has been widely documented in male songbirds. However, few studies have focused on duetting species, and particularly on those where pairs have pair‐specific duet codes (consistent associations of their individual phrase types). Given that each pair in the population can have a unique repertoire of duet types, duet codes have been hypothesized to enhance discrimination. In this context, we tested for evidence of neighbour recognition and duet code discrimination in two closely related species of neotropical wrens, the riverside wren, Cantorchilus semibadius, and the canebrake wren, C. modestus zeledoni. Although both species have moderately large repertoires, riverside wrens have higher levels of phrase type and duet type sharing across the population. We compared the approach and vocal responses of focal pairs to three playback treatments: neighbours' correct duet type, neighbours' incorrect duet type and a strangers' duet type. We found that riverside wrens displayed a stronger response to the strangers' playback than to both neighbours' playbacks, whereas no differences among treatments were found in canebrake wrens. Given that both species exhibited similar levels of aggression during neighbour playbacks, regardless of whether the correct duet code was used, our findings suggest duet codes do not facilitate neighbour recognition. We conclude that the function of duet codes in these species might be more closely related to intra‐pair communication. Finally, we suggest that the level of dear enemy effect a species exhibits depends on ecological factors that influence the perceived level of threat of territory intruders.     

Repertoire sharing by the individual and the pair: Insights into duet function and development in the plain wren Thryothorus modestus

Although both repertoire sharing and individual variation have been studied widely in male solo song, few studies have investigated these aspects of song in a species that performs complex duets. This study examined song repertoires of both males and females of the plain wren (canebrake subspecies, Thryothorus modestus zeledoni ), which performs antiphonal duets, and tested for patterns of repertoire sharing at the level of the individual and of the pair. We suggest that both males and females of the plain wren develop their phrase repertoires by direct imitation from other individuals of their sex, but that males, who may exist for a period as 'floaters' after dispersal, learn their songs from a wider pool of tutors than do females. In contrast to considerable individual repertoire sharing, whole duet types were very rarely shared between pairs, despite the potential for this to occur. This suggests that duet types are developed within the pair without reference to other pairs and the implications of this for duet functions are discussed.     

Sedentary life style of Neotropical sedge wrens promotes song imitation

To what extent has the style of song development among songbirds coevolved with other life history strategies? Among Cistothorus wrens in North America, it seems that sedentary or site-faithful habits of marsh wrens, C. palustris, favour song imitation, but seminomadic habits of sedge wrens, C. platensis, favour song improvisation, whereby each male generates a large but unique song repertoire. In this study, we tested whether more sedentary populations of sedge wrens in the Neotropics would imitate songs. At our primary study site near Cartago, Costa Rica, breeding birds were colour-banded during 1995 and 1996, and follow-up surveys revealed that the birds remained at this site the year round. Extensive tape recording and analysis of songs showed that males had large song repertoires (200-300+ songs), and that many songs were shared among neighbouring males. In addition, males only 27 km distant, at La Pastora, used different songs. Furthermore, matched countersinging, in which two males answer each other with identical song types, was recorded near Brasilia, in Brazil. The sharing of songs among permanent neighbours, microgeographical variation in song, and matched countersinging can be achieved only through song imitation, thus revealing a striking difference in the style of song development among different populations of the sedge wren. In the Neotropics, having predictable neighbours throughout life appears to have favoured song imitation, so that individuals can interact using a common, learned code typical of the local population; among more mobile populations in North America, however, individuals improvise large repertoires of species-typical songs, thereby enabling singing males to communicate with any individual, no matter what the population of origin. Strategies of song development must correlate with life history features, and further surveys are needed to make sense of the great diversity of singing behaviours among songbirds. Copyright 1999 The Association for the Study of Animal Behaviour.     

SONG POPULATIONS AND DISPERSAL IN STEELBLUE AND PURPLE WIDOWFINCHES

Payne, R. B. 1985. Song populations and dispersal in Steelblue and Purple Widowfinches. Ostrich 56:135-146.

Songs of Steelblue Widowfinches Vidua chalybeata were recorded over eight years at Lochinvar National Park, Zambia. Neighbouring males share their song-type repertoires. Individual birds changed the structural details of all song types in their repertoire from year to year, and over 4–5 years the songs accumulated changes so that a song type could scarcely be recognized as the same. A few males in addition switched their song repertoires when they dispersed from one song population to another, or when their old neighbours disappeared and new neighbouring males sang a different song repertoire. Individual marked adult birds moved from one song neighbourhood to another. Purple Widowfinches had a similar song behaviour but only one song dialect was found in the study area.

An estimate of the proportion of immigrants was made from the proportion of birds with songs unlike their neighbours, and from birds that had a mixed song repertoire or that switched their song repertoires. Song populations were sampled at several localities in the Transvaal, Botswana, Zambia, Kenya, Tanzania, Cameroon, and Nigeria. An overall minimal estimate of dispersal between song neighbourhoods was 18 %. The incidence of observed dispersal and the movements indicated by the song differences among local widowfinches were considerable and more than sufficient to prevent genetic isoloation and differentiation of local song populations.     

Song sharing and singing strategies in the winter wren Troglodytes troglodytes

The winter wren is a common forest bird living in groups of few adjacent neighbours during the breeding season. Inside each group, males vocally interact in the context of both territorial holding and sexual competition, forming a complex communication network. To study this network, we first analysed song type and syllable repertoires within and between distinct groups. We found a limited number of song types highly stereotyped in length, syntax and syllable composition, frequently shared among neighbours. Between groups, song type and syllable repertoires sharing decreased with increasing distance at a higher rate for song types than for syllables. Then, with continuous recordings, we focused on the dynamics of acoustic interactions between neighbours. We showed that male winter wrens can differentially use their song type repertoire (non-matching strategy), overlap their neighbours and modulate their singing rhythm producing longer inter-song intervals with no change in song length during acoustic interactions.     

Acoustic Monitoring Reveals Congruent Patterns of Territorial Singing Behaviour in Male and Female Tropical Wrens

Many animals defend territories against conspecific individuals using acoustic signals. In birds, male vocalizations are known to play a critical role in territory defence. Territorial acoustic signals in females have been poorly studied, perhaps because female song is uncommon in north‐temperate ecosystems. In this study, we compare male vs. female territorial singing behaviour in Neotropical rufous‐and‐white wrens Thryothorus rufalbus, a species where both sexes produce solo songs and often coordinate their songs in vocal duets. We recorded free‐living birds in Costa Rica using an eight‐microphone Acoustic Location System capable of passively triangulating the position of animals based on their vocalizations. We recorded 17 pairs of birds for 2–4 consecutive mornings and calculated the territory of each individual as a 95% fixed kernel estimate around their song posts. We compared territories calculated around male vs. female song posts, including separate analyses of solo vs. duet song posts. These spatial analyses of singing behaviour reveal that males and females use similarly sized territories with more than 60% overlap between breeding partners. Territories calculated based on solo vs. duet song posts were of similar size and similar degrees of overlap. Solos and duets were performed at similar distances from the nest for both sexes. Overall, male and female rufous‐and‐white wrens exhibit very similar spatial territorial singing behaviour, demonstrating congruent patterns of male and female territoriality.     

Neighbour recognition by resident males in the banded wren, Thryothorus pleurostictus, a tropical songbird with high song type sharing

Several studies and reviews have suggested that the ability to discriminate between neighbours and strangers decreases as neighbour song repertoire size and song type sharing increase. We tested the recognition capabilities of territorial male banded wrens by comparing the aggressive approach responses of focal birds to three playback treatments: shared song types sung by an adjacent neighbour (neighbour song), shared song types sung by unfamiliar birds (mimic song), and unshared song types sung by unfamiliar birds (unfamiliar song). All three treatments for each male were broadcast from the same location on the territorial boundary shared with the appropriate neighbour. As expected, focal males responded nonaggressively to the neighbour treatment and aggressively to the unfamiliar song treatment. The approach response to the mimic treatment was statistically indistinguishable from the unfamiliar treatment and significantly higher than the neighbour treatment, suggesting that most males were able to recognize unfamiliar singers even when the song types played were very similar to those of their neighbours. The relative strength of responses to the mimic varied: some males treated the mimic song with low aggression levels typical of responses to neighbour song. Repertoire sizes of focal and neighbour birds, the fraction of song types shared among neighbouring males, and the similarity of neighbour and mimic song types did not explain this variation. Therefore, within the short 3-min period of our playback experiments, some birds may have used repertoire composition as a recognition cue and confused the mimic with the neighbour. Copyright 2001 The Association for the Study of Animal Behaviour.     

SONG REPERTOIRES AND REPERTOIRE SHARING IN A LOCAL GROUP OF BLACKBIRDS

ABSTRACT

The song repertoires of a local group of blackbird Turdus merula males were determined and quantified with respect to individuality and sharing between neighbours. We focused on the long-ranging, introductory whistle part of the song and its motifs of fixed sequences of sounds. These motifs, whether they are used to start the song or placed centrally in the song, constitute the largest units of song that are repeated in the same way every time they are sung. Blackbird motifs therefore are equivalents of song types in other species. The result shows that one needs to analyse more than 200 songs to estimate a male's repertoire, which averaged 44 different motifs. Relative to other species, this is a medium to large song repertoire. The size of the repertoire of start motifs (on average 32) varied only a little between the individuals, whereas that of central motifs (on average 12) varied rather much between the same individuals, suggesting that they form a potential cue for assessment of male quality. The males within the neighbourhood showed a high degree of start motif sharing, which, together with the relatively large repertoires, could be constraining neighbour recognition. However, in most males the possession of a few individually distinctive and frequently repeated motifs could counter such an effect.     

BIOACOUSTICS IN NEW ZEALAND AND JAPAN

Although it is highly recognisable, relatively little is known about the repertoire composition and singing behaviour of the Hermit Thrush (Catharus guttatus). To address this, we recorded spontaneously singing males in two eastern populations (Halifax, NS (n = 11) and Hancock County, ME (n = 7)) and analysed the recordings with respect to repertoire size, basic song type characteristics and song syntax. Males had song type repertoires of 7–12 song types, and no song type sharing between individuals was observed within or between populations. While frequency-related structural characteristics of the song types were the same between the populations, song type duration (especially that of the introductory note) differed significantly. The song types within each repertoire could be categorised into high and low song types based on introductory note frequency, and these song type categories also differed with respect to the amount and distribution of spectral energy. In both populations, males sang with immediate variety, never repeating the same song type consecutively, and preferentially used some song-type to song-type transitions more than others. In addition, we found correlational evidence of changes in singing behaviour (e.g., the relative use of low versus high song types) over the course of the breeding season. We discuss these findings in the context of other songbird research as well as their implications for future work examining how Hermit Thrush males utilise their songs in defending territories and attracting mates.     

Duetting in space: a radio-telemetry study of the black-bellied wren

In many birds, individuals 'answer' the songs of their pair-mates to produce vocal 'duets'. One hypothesized function of song answering is that it prevents extra-pair birds from intruding into the duetting pair's territory to obtain copulations or usurp one of the pair-mates. In this capacity, answering may signal that the pair-mates are close together, and so are prepared to defend against such an intrusion. Another functional hypothesis states that answering helps pair-mates maintain contact, and so predicts that a bird is more likely to approach its mate after a duet than after a solo song. I used radio-telemetry to monitor the distance between mated black-bellied wrens (Pheugopedius fasciatoventris). I found that birds of both sexes were more likely to answer their mate's song when the mate was close, and that maximum duet length was negatively related to the distance between pair-mates. Furthermore, song answering positively affected the likelihood of one pair-mate approaching the other after a song. In a significant majority of the approaches after duet songs, the answering bird approached the initiator. I conclude that in the black-bellied wren, (i) the occurrence and duration of vocal duets covary with physical closeness and (ii) contact maintenance is a secondary function of duet participation.     

Singing the Popular Songs? Nightingales Share More Song Types with Their Breeding Population in Their Second Season than in Their First

Signals used in communication often change throughout an individual’s life course. For example, in many song bird species, males modify their song especially between their first and second breeding season. To address one possible reason of such modification, we investigated whether common nightingales Luscinia megarhynchos adjust their song type repertoires to sing the song types commonly occurring in their breeding population. We analysed nocturnal singing of six nightingales in their first and second breeding season and compared their repertoire composition and usage to the ‘typical’ repertoire and usage on the breeding ground (represented by seven reference birds). Songs that were maintained between the first and second season by the six focal birds occurred in most of the repertoires of the seven reference birds and were sung often. In contrast, song types that were dropped from the repertoires occurred less often in the reference birds’ repertoires and were sung less often. Furthermore, in the first year, each focal nightingale’s repertoire was less similar to the reference birds’ repertoires than in the second year. Thus, nightingales adjusted their singing towards the songs popular in the breeding grounds by keeping song types that were common and frequently sung by other individuals in their breeding area and by disposing of infrequently performed ones. This resulted in increased similarity with the population’s repertoire from the first to the second year. We discuss possible ontogenetic processes that may lead to such an adjustment and suggest an improved ability to match song types as possible adaptive value.     

Female marsh wrens do not provide evidence of anatomical specializations of song nuclei for perception of male song

In songbirds the forebrain nuclei HVC (high vocal center) and RA (robust nucleus of the archistriatum) are larger in individuals or species that produce larger song repertoires, but the extent to which the size of these nuclei reflects a need for either producing or perceiving large repertoires is unknown. We, therefore, tested the hypothesis that species differences in the size of song nuclei reflect a commitment of “brain space” to the perceptual processing of conspecific song. The two species of marsh wren (Cistothorus palustris western and eastern) provide a good test case. Western males produce larger song repertoires, and have larger HVC and RA than do eastern males. Female marsh wrens do not sing, and if they use their song nuclei to assess conspecific male song repertoires, then we predicted that measurable cellular and nuclear parameters of HVC and RA would be greater in western than eastern female wrens. For males we confirmed that the volumes of HVC and RA, and cellular parameters of HVC, are greater in western than in eastern birds. These nuclei were also considerably larger in males than in conspecific females. Western and eastern female wrens, however, did not differ in any measured parameters of HVC or RA. Females of these wren species thus do not provide any direct evidence of anatomical specializations of song nuclei for the perceptual processing of conspecific male song. 1994 John Wiley & Sons, Inc.     

Male and female rufous‐and‐white wrens do not match song types with same‐sex rivals during simulated territorial intrusions

In birds with song repertoires, song‐type matching occurs when an individual responds to another individual's song by producing the same song type. Song‐type matching has been described in multiple bird species and a growing body of evidence suggests that song‐type matching may serve as a conventional signal of aggression, particularly in male birds in the temperate zone. Few studies have investigated song‐type matching in tropical birds or female birds, in spite of the fact that avian biodiversity is highest in the tropics, that female song is widespread in the tropics, and that female song is the ancestral state among songbirds. In this study of rufous‐and‐white wrens Thryophilus rufalbus, a resident neotropical songbird where both sexes sing, we presented territorial males and females with playback that simulated a territorial rival producing shared and unshared songs. In response, both males and females sang matched song types at levels statistically equal to levels expected by chance. Furthermore, males and females exhibited similar levels of aggression and similar vocal behaviours in response to playback of both shared and unshared songs. These results indicate that rufous‐and‐white wrens do not use song‐type matching in territorial conflicts as a conventional signal of aggression. We discuss alternative hypotheses for the function of song‐type sharing in tropical birds. In particular, we point out that shared songs may play an important role in intra‐pair communication, especially for birds where males and females combine their songs in vocal duets, and this may supersede the function of song‐type matching in some tropical birds.     

Trill consistency is an age-related assessment signal in banded wrens

Older males tend to have a competitive advantage over younger males in sexual selection. Therefore, it is expected that signals used in sexual selection change with age. Although song repertoire size in songbirds is often mentioned as an age-related trait, many species, including the banded wren (Thryothorus pleurostictus), do not increase their repertoires after the first year. Here, we show that banded wrens reproduce the trill notes in their songs with less variability between them (i.e. more consistently) when they grow older. In a playback experiment, we also show that banded wrens discriminate between younger and older birds based on structural aspects of their song. In a second experiment, banded wrens also respond differentially to natural songs versus songs with artificially enhanced consistency. We argue that consistency in trill note reproduction may be achieved through practice. Sexual selection in the form of male–male competition may therefore operate on a phenotypic trait, the expression of which is enhanced by practice.     

Repertoire Sharing and Song Similarity between Great Tit Males Decline with Distance between Forest Fragments

Birdsong can play a critical role in establishing a territory and finding a mate among individuals from local and foreign populations. Variation in birdsong among populations can be influenced by habitat fragmentation and might affect successful dispersal among habitat fragments. We studied variation in great tit song in a long‐term study population distributed over nine forest fragments. All individual males recorded had a known dispersal history within the fragmented forest habitat. We found spatial structure of declining song‐type sharing with distance, with a marked drop from an individual’s own forest fragment to another across a habitat gap. We also found decreasing song similarity among increasingly distant fragments in terms of temporal and spectral characteristics of shared song types. The change in acoustic structure was more gradual and seemed less affected by habitat discontinuity but also showed a tight correlation with dispersal index among forest fragments. Immigrant birds shared fewer song types with neighbouring birds that were born within the same forest fragment, but not less compared to birds born in another forest fragment within the study area. Our data provide detailed insight into the relationship between song differentiation and male dispersal and contribute to our understanding of the potential role of song in reproductive exchange and avian speciation. The fact that birds in small forest fragments shared more songs than birds in larger forest fragments confirms that song analysis has potential as a tool for conservation in rare species.     

Songbird cheaters pay a retaliation cost: evidence for auditory conventional signals

Conventional signals impose costs on senders through receiver retaliation rather than through investment in signal production. While several visual conventional signals have been described (mainly 'badges of status'), acoustic examples are rare; however, several aspects of repertoire use in songbirds are potential candidates. We performed interactive playback experiments to determine whether song-type matches (responding to a song with the same song type), repertoire matches (responding to a song with a different song type, but one in the repertoires of both singers) and unshared song types serve as conventional signals during male-male territorial interactions in banded wrens, Thryothorus pleurostictus. Our results demonstrate that these three signals incite varying levels of receiver aggression: song-type matches induce faster approach than do repertoire matches, and repertoire matches induce faster approach than do unshared song types. Production costs do not differ, while the receiver response does. Because territorial banded wrens approach opponents who signal aggressively, such opponents risk attack. This system will punish and prevent cheaters, as weak males signalling aggression will be subject to escalation by stronger or more-motivated opponents.     

A test for repertoire matching in eastern song sparrows

Repertoire matching occurs when one songbird replies to another with a song type that the two birds share. Repertoire matching has previously been demonstrated to occur at well above chance levels in a western population of song sparrows, where it is hypothesized to serve as a low level threat in a hierarchy of aggressive signals. Here we test for repertoire matching in an eastern population of song sparrows. Previous work indicates that this eastern population differs from the western one in having lower levels of song sharing between neighboring males and in showing no association between song sharing and territory tenure. Here we confirm that males in this eastern population on average share few whole songs with their neighbors. The eastern males are familiar with their neighbors’ repertoires, as evidenced by a stronger singing response to stranger song than to neighbor song. Males in the eastern population did not repertoire match: when played an unshared song type from a specific neighbor, they did not reply with a song type shared with that neighbor more often than expected by chance or more often than in response to playback of a control song (an unshared stranger song). The results thus demonstrate a qualitative difference in vocal signaling strategies between two populations of the same species.     

Song Sharing in Neighboring and Non-Neighboring Thrush Nightingales (Luscinia luscinia) and its Implications for Communication

In many passerines, males have repertoires of different songs of which some songs are often shared with other males. Sharing of song repertoires among males can provide insights into the context in which songs were acquired and on the role of song repertoires in inter- and intrasexual communication. Here we studied repertoire sharing in male territorial thrush nightingales ( Luscinia luscinia ). We compared male vocal repertoires of the basic song components, full songs, and the sequencing of songs in a bout. The results show that males differed significantly in the size of their song repertoires but not in the size of the repertoire of basic song components. Moreover, they shared almost all (80%) the repertoire of song components but only 30% of their song types. Neighboring males shared significantly more song types than did non-neighboring males but interestingly they did not share more basic song components than non-neighboring males. These results show that the repertoire of basic song components is under much less sexual selection than the size of song repertoires. Sharing of song repertoires among neighbors presumably results from repertoire conversion over time and from males returning to their territories in the following season. Repertoire sharing then can be an indicator of territory tenure and thus it can be important in repelling rivals and in female choice.     

Vocal mimicry in male bowerbirds: who learns from whom?

Vocal mimicry is one of the more striking aspects of avian vocalization and is widespread across songbirds. However, little is known about how mimics acquire heterospecific and environmental sounds. We investigated geographical and individual variation in the mimetic repertoires of males of a proficient mimic, the spotted bowerbird Ptilonorhynchus maculatus. Male bower owners shared more of their mimetic repertoires with neighbouring bower owners than with more distant males. However, interbower distance did not explain variation in the highly repeatable renditions given by bower owners of two commonly mimicked species. From the similarity between model and mimic vocalizations and the patterns of repertoire sharing among males, we suggest that the bowerbirds are learning their mimetic repertoire from heterospecifics and not from each other.