Vocal Plasticity – are Pied Flycatchers,Ficedula Hypoleuca,Open‐Ended Learners?

In some songbird species, large song repertoires are advantageous in female attraction, whereas song sharing with neighbours may give an advantage in male–male competition. Open‐ended learners, with the ability to memorize new song elements throughout their lives, may learn from territorial neighbours and thus benefit from increasing both repertoire size and song sharing. A distinction needs to be made between true adult song learning, i.e. memorization of novel song elements, and vocal plasticity resulting in changes in the use of previously memorized elements, such as the use of hidden repertoires or increased production of previously rare syllable types. We assessed the ability of adult pied flycatcher Ficedula hypoleuca males to learn previously unheard song elements and to change their song production in response to playback of unfamiliar, conspecific song, emulating a singing neighbour. After a 1‐week playback treatment, three out of 20 subjects had learned foreign song elements, providing evidence from the wild that pied flycatchers are true open‐ended learners. However, the syllable sharing with the playback stimulus repertoires had not changed, and the males’ repertoires had decreased rather than increased. Hence, we did not find support for increased syllable sharing with neighbours or increased repertoire size as functions of adult song learning in pied flycatchers. Because pied flycatcher song seems to serve mainly for mate attraction, copying of attractive syllable types is a possible alternative function of adult song learning in this species.     

Habituation,Recovery and the Similarity of Song Types within Repertoires in Red-winged Blackbirds (Agelaius phoeniceus) (Aves,Emberizidae)

Male red-winged blackbirds (Agelaius phoeniceus) respond to playback of conspecific song on their territories with the song spread, a graded aggressive display in which males extend their wings to expose their red epaulets while singing. We show that the intensity of song spread display declines with repeated presentation of one song type, and recovers when song types are switched. Recovery is greater for switches between song types that are acoustically dissimilar than for switches between song types that are acoustically similar. Recovery is no different for switches between two song types taken from the repertoires of different males than for switches between song types recorded from the same male. Analysis of acoustic features also indicates that song types recorded from different males are not more dissimilar than are song types from the same male. Our results do not support the idea that repertoires of red-winged blackbirds are composed of similar song types in order to facilitate individual recognition. Rather, repertoires may be constructed of dissimilar song types, so as to help maintain the response of listeners despite habituation.     

Adult Song Sparrows do not Alter their Song Repertoires

To understand fully the function of vocal learning, it is important to know when, during an individual's lifetime, learning occurs. Songbirds are generally categorized into two groups with respect to their adult song learning ability. 'Open-ended' song learners are able to learn to produce new songs in adulthood, whereas 'age-limited' song learners can only acquire songs during their first year of life. Researchers have long assumed that certain oscine species are open-ended or age-limited song learners, but the evidence to date has been inadequate to test these assumptions for most species. We tested the hypothesis that song sparrows ( Melospiza melodia ) are age-limited song learners who do not alter their song repertoires in adulthood by examining the song repertoires of 24 color-banded males who were fully recorded in two, three or four different years. We compared sonagrams of the song types produced by males in different years and looked for any changes in repertoire composition (i.e. added or dropped song types). With few exceptions, males produced song repertoires that were identical in every year they were recorded. The exceptions (four males who did not produce one of their song types during one recording session) were all cases in which we believe that we missed recording a song type that a male did indeed have, not that the males dropped a song type. The finding that adult males do not alter the composition of their song repertoires provides strong evidence that song sparrows are age-limited song learners. Although it is possible that song sparrows make subtle within-song type changes across years, such changes would not necessarily constitute new song learning.     

Effects of location in territory and distance from neighbours on the use of song repertoires by Carolina wrens

By capturing territorial Carolina wrens (Thryothorus ludovicianus) and housing them in portable cages on their territories, I could control each one's location in its territory and distance from neighbours in experiments on the use of song repertoires. Experiment 1 demonstrated that these wrens sang more songs in the centre of their territories than at the edge, but that they did not use more song types or different song types at the centre than at the edge. In experiment 2, in which I played tape-recorded songs at two distances from wrens caged in the centres of their territories, birds responded more strongly to songs at 25m, simulating an intruder, than to songs at 165 m, simulating a territorial neighbour. Birds also switched more frequently between song types and sang more song types per 100 songs in response to the nearer playback. Experiment 3 compared captive wrens 140 m, 80 m, 20 m, or 0 m apart on adjacent territories. As the distance between neighbours decreased, birds sang less, but also switched more frequently between song types, used more song types per 100 songs, and matched songs with neighbours more frequently. There were no differences in the kinds of song types sung at different distances from neighbours. A comparison of the results from experiments 1 and 3 confirms that the use of song repertoires is influenced by distance from conspecifics and not by location in the territory.     

Differences in Frequency of Shared Song Types Enables Neighbour‐Stranger Discrimination in a Songbird Species with Small Song Repertoire

Acoustic Neighbour‐Stranger (N‐S) discrimination is widespread in birds and has evolved to settle territorial disputes with low costs. N‐S discrimination was found both in song‐learning oscines and non‐song‐learning bird taxa, irrespective of the repertoire sizes they have. Therefore, it seems that more than just a single mechanism enable N‐S discrimination. Species with larger repertoires, where males have unique phrases or syllables may rely on such interindividual differences. The majority of birds have rather small repertoires, which often are shared among neighbours. In this case, males are facing the problem of individual recognition when rivals produce songs, at least superficially, identical. To better understand the acoustic basis of N‐S discrimination in species with small and shared repertoires, I studied the ortolan bunting (Emberiza hortulana). Males of this small oscine species are able to N‐S discrimination based on a single song rendition when presented in a playback experiment, regardless of song‐type diversity and song‐sharing level within a particular population. It was also found that songs of the same type sung by different males differ in the frequency of the initial song phrases and these differences persist over years. Here, I tested whether males are able to discriminate among the natural songs and the artificially modified songs of their neighbours in which the frequency was experimentally changed by relatively small value in comparison with the variation range found in this population. Subjects responded significantly more aggressively to the songs with an artificially modified frequency, suggesting that males treat such songs as having come from the repertoire of a non‐neighbour. These results confirm an earlier prediction that differences in the frequency of shared song types enable N‐S discrimination. The study presents one of the possible mechanisms enabling N‐S discrimination in songbirds with small repertoires and stress the role of within‐song‐type variation, which is still understudied song characteristic.     

Song characterization in the spectacled warbler (Sylvia conspicillata): a circum-Mediterranean species with a complex song structure

The spectacled warbler (Sylvia conspicillata) is a small passerine with a patchy distribution throughout the circum-Mediterranean region, including the North Atlantic archipelagos of Madeira, Canary Islands and Cape Verde. Here we characterize the species song structure on the island of Fuerteventura, quantifying repertoire size, inter- and intra-individual spectrographic variation, to determine whether acoustic variation occurred within an island population. Male song display was organized in song bouts of a variable number of song phrases, which in turn were made up of 4–69 syllables. We classified syllable types to derive a measure of repertoire size (number of different syllables) per song bout, and then used rarefaction methods to calculate the estimated repertoire size for our population of males. Three categories of song bout length were considered in analyses: short song bouts of 10 phrases, average bouts of 19 phrases and long bouts of ≥ 29 phrases. The observed and estimated repertoire size per male (between 43 and 126 syllables per male) increased with song bout duration, although the relationship was not significant for the estimated values. To test whether songs could be individually specific, we measured 11 spectrotemporal parameters of the song. A discriminant analysis using these variables performed poorly in classifying songs to the individuals that uttered them, but we found less variation in the individual than in the population for three out of the 11 variables. These individually specific variables, involving the first or the most common syllable of the song, the trill, were the duration of the first syllable of the phrase, the duration and the dominant frequency of the trill syllable. Our study emphasizes the complexity of spectacled warbler songs, in which males continuously add novel syllables over the entire song bout. This complexity appears to be determined by individual innovation capabilities rather than by the behaviour of copying neighbour repertoires, since songs of close birds were not more similar than songs from far-away territories.     

Searching for the functional origins of song in eastern brown-headed cowbirds,Molothrus ater ater

Three experiments were conducted to examine the functional properties of vocal precursors to stereotyped song in socially housed captive cowbirds. Previous studies had shown that eastern male cowbirds developed different song repertoires when housed in different social contexts. This paper reports on the acoustic origins of the different vocal outcomes. In addition, a winter roost of cowbirds was studied to investigate the setting in which males naturally develop song. Analyses of the vocalizations of laboratory-housed subjects revealed acoustic differences from the earliest stages of song ontogeny. The field data indicated that free-living and laboratory-housed males shared the same structural categories of song and that the winter roost provided males with opportunities to interact with females. Taken as a whole, the studies suggest that song ontogeny is a dynamic process involving social and vocal interactions between singers and listeners.     

Evidence of repertoire sharing and stability despite a high turnover rate in a duetting neotropical wren

In songbirds, the spatial pattern of song sharing among individuals is influenced by the song learning and dispersal strategies within each species. In birds where females and males sing and create joint acoustic displays (duets), the processes defining the patterns of song sharing become more complex as there might be different selection pressures shaping the behaviour of each sex. To provide further insight into the vocal development and the dispersal strategy of duetting tropical species, we investigated the patterns of individual and pair repertoire sharing, as well as the stability of these repertoires, in a colour-marked population of riverside wrens, Cantorchilus semibadius, located in the Osa Peninsula, Costa Rica. Using data collected over a five-year period, we found considerable variation in the sharing levels of phrase and duet type repertoires among neighbouring individuals coupled with a general decline of repertoire sharing as distance increased between birds’ territories. These results are consistent with a pattern predicted in age-restricted learners that establish preferentially near their tutors. Furthermore, we found no evidence of individuals changing their phrase type repertoires over time, including after remating events. Duet type repertoires were also stable when pairs remained together. However, we observed a surprisingly high turnover rate. When individuals remated, even though the majority of the previous duet type repertoire remained, several new duet types were included. Taken together, our findings suggest that riverside wrens might create their individual repertoires by copying their same-sex parent and neighbouring individuals before dispersal. Additionally, we speculate that even though birds were able to create new duet types after changing partners, a substantial portion of their duet type repertoire might also be copied from their parents and neighbouring pairs during the initial critical period of song learning.     

Song matching in a long-lived,sedentary bird with a low song rate: The importance of song type,song duration and intrusion

Territorial songbirds often match the song features or singing patterns of rivals, commonly as an aggressive signal. Most studies of song matching have been on Northern Hemisphere species with short lifespans and high song rates, but vocal matching is predicted to be affected both by longevity and territorial stability. We studied song matching in males of the white-browed scrubwren, Sericornis frontalis, a long-lived, sedentary, territorial Australian songbird. We quantified natural song rate and diversity, and then conducted three playback experiments to test: (a) whether males match by song type; (b) how they respond physically and vocally to territorial intrusion; and (c) whether they match by song length, and use it as an agonistic signal. Males naturally had very low song rates, singing on average less than three times per hour, and moderate repertoires, with an estimated mean of 17.5 song types for individual males. Males did not engage in extended counter-singing bouts. The first experiment showed that males matched the song type of immediate neighbours almost 90% of the time, if that type was in their repertoire. The remaining experiments revealed that song-type matching was an aggressive signal; males responded more aggressively to, and were more likely to match, playback simulating a neighbour's territorial intrusion than song from their shared boundary. Males did not match songs by length, but they produced longer songs after simulated intrusion. Males also responded more aggressively to playback of longer songs that simulated intrusion, but less aggressively to longer songs from the territory boundary. Overall, we show that sedentary, long-lived songbirds with low song rates, can use song-type matching as an aggressive signal to communicate with neighbours and intruders. Song length had a different role in communication, possibly related to individual quality or territory ownership.     

Conventional and network analyses of song organization and complexity in northern House Wrens (Troglodytes aedon parkmanii)

Among songbirds with large song-type repertoires, there may be functional variation in how individuals organize and display song-type diversity over time. Past studies focusing on conventional measures of song production have been extremely productive. However, network analysis offers a novel set of tools to quantify additional, previously unstudied elements of song organization and display. We studied protracted bouts of singing by 10 male House Wrens (Troglodytes aedon) to (1) test functional hypotheses of variation in song diversity in this species, and (2) evaluate the utility of network metrics in such research. Our analysis included a variety of conventional measures of song production and several standard metrics from network theory to quantify how variably the many song types in a male’s repertoire could be connected to one another and the limitations or diversity of their song sequences. Analysis of conventional variables showed that males produced more and longer songs, at shorter intervals, containing more syllables and more different syllable types, and also more different song types, prior to than after pairing and early in the morning regardless of breeding stage. These results are consistent with the hypothesis that song diversity functions in mate attraction and possibly in territory signaling. In contrast, analyses of network metrics revealed variety in song sequencing by males, but comparatively few and weak effects associated with either breeding stage or time of day. Overall, most song types connected to only a few others and a relatively small proportion of all possible song-type transitions actually occurred. Hence, much of the variety in song sequencing that was possible with the large song repertoires of males was not realized. The latter outcomes, brought to light via network analyses, highlight an important paradox for future research on this and related species with large song repertoires.     

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.     

Relationship between Song Repertoire and Age in the Canary,Serinus canarius

We recorded the song of six male canaries during their first and second year of life to see if we could detect systematic age-related changes. Soundspectrographic analysis revealed that many syllables from the previous year were omitted and many new ones were added. There was a significant increase in syllable repertoire during the second year. The proportion of single-element syllables increased between year 1 and year 2. These results indicate that neural pathways responsible for song learning in canaries remain plastic in adulthood. We suggest that the larger song repertoires of older ♂♂ confer a reproductive advantage by acting as a more potent stimulus on ♀♀.     

Interspecific Differences in the Response of Female Birds to Song Repertoires

The functional significance of the wide variation between bird species in the sizes of individual song repertoires is not understood. We have studied the effects of song repertoires on females. Song triggers copulation solicitation display in female sparrows treated with estradiol. Song sparrow males (Melospiza melodia) have repertoires of about 10 song types, and female song sparrows display significantly more to presentations of large repertoires than to a single song type. By contrast, male field sparrows (Spizella pusilla) and white-throated sparrows (Zonotrichia albicollis) each have only one song type, and their females show no significant increment in responsiveness to repertoires of several song types over a single song type. Swamp sparrows (Melospiza georgiana) fall in between, with male repertoires of three song types. Female swamp sparrows behave in intermediate fashion, responding more to several song types than to one, but the response increment is less than in song sparrows. Thus species differences that males exhibit in song repertoires are paralleled by differences in female responsiveness to multiple song types, implicating variations in female reactivity in the evolution of song repertoires. Female song sparrows respond preferentially to repertoires programmed in eventual variety rather than immediate variety, while field sparrows and white-throated sparrows show no discrimination.     

Song convergence in multiple urban populations of silvereyes (Zosterops lateralis)

Recent studies have revealed differences between urban and rural vocalizations of numerous bird species. These differences include frequency shifts, amplitude shifts, altered song speed, and selective meme use. If particular memes sung by urban populations are adapted to the urban soundscape, "urban-typical" calls, memes, or repertoires should be consistently used in multiple urban populations of the same species, regardless of geographic location. We tested whether songs or contact calls of silvereyes (Zosterops lateralis) might be subject to such convergent cultural evolution by comparing syllable repertoires of geographically dispersed urban and rural population pairs throughout southeastern Australia. Despite frequency and tempo differences between urban and rural calls, call repertoires were similar between habitat types. However, certain song syllables were used more frequently by birds from urban than rural populations. Partial redundancy analysis revealed that both geographic location and habitat characteristics were important predictors of syllable repertoire composition. These findings suggest convergent cultural evolution: urban populations modify both song and call syllables from their local repertoire in response to noise.     

Rufous‐capped Warblers Basileuterus rufifrons show seasonal,temporal and annual variation in song use

In the majority of songbird species, males have repertoires of multiple song types used for mate attraction and territory defence. The wood‐warblers (family Parulidae) are a diverse family of songbirds in which males of many migratory species use different song types or patterns of song delivery (known as ‘singing modes’) depending on context. The vocal behaviour of most tropical resident warblers remains undescribed, although these species differ ecologically and behaviourally from migratory species, and may therefore differ in their vocal behaviour. We test whether male Rufous‐capped Warblers Basileuterus rufifrons use distinct singing modes by examining song structure and context‐dependent variation in their songs. We recorded multiple song bouts from 50 male Warblers in a Costa Rican population over 3 years to describe seasonal, diel and annual variation in song structure and vocal behaviour. We found that Rufous‐capped Warbler songs are complex, with many syllable types shared both within and between males’ repertoires. Males varied their song output depending on context: they sang long songs at a high rate at dawn and during the breeding season, and shortened songs in the presence of a vocalizing female mate. Unlike many migratory species, Rufous‐capped Warblers do not appear to have different singing modes; they did not change the song variants used or the pattern of song delivery according to time of day, season or female vocal activity. Our research provides the first detailed vocal analysis of any Basileuterus warbler species, and enhances our understanding of the evolution of repertoire specialization in tropical resident songbirds.     

Species‐level repertoire size predicts a correlation between individual song elaboration and reproductive success

Birdsong has long been considered a sexually selected trait that relays honest information about male quality, and laboratory studies generally suggest that female songbirds prefer larger repertoires. However, analysis of field studies across species surprisingly revealed a weak correlation between song elaboration and reproductive success, and it remains unknown why only certain species show this correlation in nature. Taken together, these studies suggest that females in numerous species can detect and prefer larger repertoires in a laboratory setting, but larger individual repertoires correlate with reproductive success only in a subset of these species. This prompts the question: Do the species that show a stronger correlation between reproductive success and larger individual repertoires in nature have anything in common? In this study, we test whether between‐species differences in two song‐related variables—species average syllable repertoire size and adult song stability over time—can be used to predict the importance of individual song elaboration in reproductive success within a species. Our cross‐species meta‐analysis of field studies revealed that species with larger average syllable repertoire sizes exhibited a stronger correlation between individual elaboration and reproductive success than species with smaller syllable repertoires. Song stability versus plasticity in adulthood provided little predictive power on its own, suggesting that the putative correlation between repertoire size and age in open‐ended learners does not explain the association between song elaboration and reproductive success.     

Relative threat and recognition ability in the responses of tropical mockingbirds to song playback

It has been suggested that individual recognition based on song may be constrained by repertoire size in songbirds with very large song repertoires. This hypothesis has been difficult to test because there are few studies on species with very large repertoires and because traditional experiments based on the dear enemy effect do not provide evidence against recognition. The tropical mockingbird, Mimus gilvus, is a cooperative breeder with very large song repertoires and stable territorial neighbourhoods. The social system of this species allowed us to test individual recognition based on song independently from the dear enemy effect by evaluating male response to playback of strangers, neighbours (from shared and unshared boundaries), co-males (i.e. other males in the same social group) and own songs. Although subjects did not show a dear enemy effect, they were less aggressive to co-males than to all other singers. Our results suggest that recognition in tropical mockingbirds (1) does not simply distinguish between familiar and unfamiliar singers, (2) requires a small sample of both songs and song types, (3) does not rely on individual-specific sequences of song types and (4) is not likely to rely on group-specific vocal signatures potentially available in cooperatively breeding groups. We conclude that this is a case of true recognition and suggest that the lack of a dear enemy effect in this and other species with large repertoires may relate to the role of song in mate attraction and the perception of neighbours as a threat to future paternity.     

A Previously Undescribed Method of Song Matching in a Species with a Single Song “Type”, the Kentucky Warbler (Oporornis formosus)

The song of male Kentucky warblers consists of several repetitions of syllables containing 2 to 7 elements within a frequency range from 1.8 to 6.0 kHz. Each male has a single, individually distinctive song type that spans part or all of this frequency range. Song responses to playbacks of songs artificially elevated or lowered in frequency by 200 Hz show that a bird is able to alter its song in two ways. It may lower or raise the frequency range of the song or put more energy into the lower or higher frequencies within the song's elements to more closely match the energy distribution of the playback song. The syllable or element structure of the song is kept constant with only the song's energy distribution among frequencies changing. Thus, Kentucky warblers are able to alter their single song type to “match countersing” in a way similar to that of species with song repertoires who switch song types to match those of rivals. The Kentucky warbler may have a single songtype because of its ability to change its song's energy distribution. This ability may perform the same functions as song matching does in species with larger repertoires. This finding is discussed in relation to ideas on the evolution of song repertoires in intrasexual competition.     

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.