Does age matter in song bird vocal interactions? Results from interactive playback experiments

The song of oscines provides an extensively studied model of age-dependent behaviour changes. Male and female receivers might use song characteristics to obtain information about the age of a signaller, which is often related to its quality. Whereas most of the age-dependent song changes have been studied in solo singing, the role of age in vocal interactions is less well understood. We addressed this issue in a playback study with common nightingales (Luscinia megarhynchos). Previous studies showed that male nightingales had smaller repertoires in their first year than older males and males adjusted their repertoire towards the most common songs in the breeding population. We now compared vocal interaction patterns in a playback study in 12 one year old and 12 older nightingales (cross-sectional approach). Five of these males were tested both in their first and second breeding season (longitudinal approach). Song duration and latency to respond did not differ between males of different ages in either approach. In the cross-sectional approach, one year old nightingales matched song types twice as often as did older birds. Similarly, in the longitudinal approach all except one bird reduced the number of song type matches in their second season. Individuals tended to overlap songs at higher rates in their second breeding season than in their first. The higher levels of song type matches in the first year and song overlapping by birds in their second year suggest that these are communicative strategies to establish relationships with competing males and/or choosy females.     

Tuning towards tomorrow? Common nightingales Luscinia megarhynchos change and increase their song repertoires from the first to the second breeding season

In many oscines, song repertoire size correlates with male quality and female mate choice, and can vary with age. In a cross-sectional field study in common nightingales Luscinia megarhynchos , one y old birds had substantially smaller repertoires than did older ones. In laboratory experiments males can acquire new song types during this period. This longitudinal field study therefore investigates whether individual nightingales increase their repertoires from the first to the second breeding season. We report a striking repertoire turnover, with an average overall increase of 24% of the first season's repertoire, resulting from added and dropped song types (54% and 30%, respectively). The number of added song types correlated positively with the size of the first season's repertoire. These results are consistent with the notion that repertoire size in nightingales correlates with male quality, although the overlap between repertoire sizes of first and second season birds makes it impossible to discriminate age based solely on repertoire size. Comparing the number of song types an individual sang in both seasons ('permanent song types') revealed a lower overlap than reported for subsequent seasons. The frequencies with which these were sung in the first season were less predictive of how often they were sung in the second season than was the case between later years. This drastic repertoire turnover from the first to the second season may be a selective process in response to the local song types, constrained by genetic makeup and shaped by early experience.     

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.     

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.     

Age-related changes in song repertoire size and song type sharing in the Whinchat Saxicola rubetra

In many oscine passerines males’ songs, the repertoire size increases with age. At the same time it often remains unknown when and where males learn new songs. To infer the Whinchat Saxicola rubetra song learning strategy, we described and catalogued song-type repertoire, revealed age differences and examined song sharing strategies among neighbouring and distant males. We recorded song vocalizations of 40 males in a limited (104 ha) study plot during four years. Whinchats produce short and discrete songs with clear intersong pauses. In total 45 song types were allocated, the individual repertoire size averaged 23.5 ± 7.6 song types (range 9–34 song types). The males’ age significantly influenced the song-type repertoire size. The second calendar year (first breeding) males had a lower repertoire size than the older males. The majority of song types were shared by less than half of males in our sample. The Jaccard similarity indexes varied from 0.5 to 0.7. We could not find a relationship between males’ song sharing and geographic distances between their nests. We assume that Whinchat males learned new songs in the local population before territory establishment.     

Breeding Experience and not Age Modulates the Song Development of Pied Flycatchers (Ficedula hypoleuca)

In songbirds, the development of the species‐specific adult song involves a learning process that varies in extension. In species that incorporate new song elements throughout life (open‐ended learners), variation in male song composition could be the result of either age or breeding experience. Using data from 16 yr of fieldwork on pied flycatchers (Ficedula hypoleuca), we aimed to disclose the individual contribution of these two factors on the species song characteristics, as well as their relation with morphology and plumage color changes. Finally, we explored whether any of the song or physical features could predict the probability of males returning to the breeding site. We found that the song characteristics of the first‐time breeders did not differ between age classes, except for the total number of syllables per song, which was higher in the 1‐year‐old than in the 2‐year‐old males. However, we found that song variables associated with complexity (song and sample versatilities and repertoire size), increased significantly from the first to the second breeding season. Males showed delayed plumage maturation, with 1‐year‐old males being browner than the 2‐year‐old males independently of their breeding experience. Morphology, however, was not affected by age or breeding experience. The probability that males returned to the breeding site was not associated with song or physical features. Considering that some song learning occurs during the breeding period and that some males may skip the first breeding season, selective pressures may have been established for song complexity to be an honest indicator of breeding experience rather than age.     

Song, sexual selection, and a song control nucleus (HVc) in the brains of European sedge warblers

Female sedge warblers select males that have more complex songs as mates. This study tests two predictions concerning HVc, a telencephalic nucleus that is essential for song learning and production: first, that males with more complex songs will have a larger HVc, and second that males who pair successfully will have a larger HVc than unpaired males. Data on song composition and pairing status were collected from wild sedge warblers breeding in Hungary. We found significant positive correlations between three song attributes (repertoire size, song complexity, and song length) and the size of HVc. Males that paired successfully also had more complex songs (repertoire size and song complexity, though not song length) than males that did not. However, we find no direct evidence that males who paired successfully had a larger HVc than unpaired males. These findings are discussed in relation to the possible functions of HVc and also to current views on sexual selection and the evolution of the song control pathway.     

Element repertoire: change and development with age in Whitethroat Sylvia communis song

Song repertoires are often important determining factors in sexual selection. In several species, older males have larger repertoires than 1-year-old males. The development of large song repertoires by an individual is, however, poorly understood. We studied song element repertoire changes in five individual male Whitethroats Sylvia communis sampled as 1- and 2-year olds. These males increased the size of their element repertoire between their first and second year, but song length and number of different elements per song did not change. On average, 44.3% of the song elements in the first-year repertoire were also found in the second-year repertoire. Elements shared between years were found earlier in the songs and tended to occur in sequences. Sequences of shared elements also seemed to be conserved between years. The study suggests that the song element repertoire of the second year is partly based on the first-year repertoire, which may explain why large song repertoires are mainly expressed by males at least 2 years of age. It would appear, therefore, that song element repertoire size could be a reliable signal of male age.     

Changes in Performance of Shared and Unshared Songs Within and Between Years in the White‐crowned Sparrow

Song of passerine birds is one of the few animal signals that is learned and that improves with practice. Vocal practice is crucial early in life to perfect a song imitation, but it also occurs throughout life and may continue to improve aspects of song performance. Differences in song performance among males that share song types, that is sing structurally similar songs may be particularly salient to receivers. We here test the hypothesis that aspects of song performance improve in a songbird species that deletes song types from its repertoire early in the first breeding season to share their final single song type with territorial neighbours. Over 3 yrs, we recorded songs in a population of Puget Sound white‐crowned sparrows Zonotrichia leucophrys pugetensis and measured percentage peak performance and consistency thereof in all of the song types in each male's repertoire. We found that within the first year on territory, percentage peak performance was higher in shared than unshared songs but did not change from first to second recording. Contrary to the hypothesis that song performance improves with age, song performance declined from the first to the second year. Our results support the hypothesis that high‐performance singers share songs. We did not find support for song performance improving within or between years, like it does in some other songbird species.     

Limited Inter‐Annual Song Variation in the Serin (Serinus serinus)

Patterns of song plasticity in passerine birds beyond the first year are poorly studied. In general, songbirds are divided into two categories: open‐ended learners and closed‐ended learners, depending on the pattern of age‐related vocal plasticity. However, recent work based on longitudinal studies revealed a broader range of flexibility of song changes in adulthood. Serins sing very complex songs with large repertoires which are delivered in a very rigid way with little structural modification. However, there is little information on how serin song changes with age. We studied vocal plasticity in wild adult serins by recording male song over 2 years. The analyses show that male songs have only limited variation between years, with no increase in repertoire size and relatively small changes in their structural characteristics. Syllable production was very consistent within and between years with very little structural variation. New syllables represented only 8% of the repertoire, and they appeared to emerge from fusion or splitting of pre‐existing syllables. We conclude that serin song while structurally complex has a very limited age‐related plasticity after the first year. We hypothesise that this structural stability is a consequence of selection for performance consistency.     

No overlap in song repertoire size between yearling and older Starlings Sturnus vulgaris

We recorded the song of 25 male Starlings Sturnus vulgaris to investigate differences in song characteristics between yearling and older males. Older males had significantly larger repertoire sizes and sang significantly longer average song bouts than yearling males. Since older males had almost twice as large a repertoire size as yearling males and since there was no overlap in repertoire size between yearling and older males, our data indicate that the repertoire size of Starlings increases after the first year of life. The largest difference between yearling and older males occurred in the 'variable song types' which are the song types containing most of the heterospecific imitations sung in a song bout.     

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.     

Age-Dependent Expression of Song in the Collared Flycatcher, Ficedula albicollis

Theory suggests that male ornaments should be reliable signals of age, with more elaborated ornaments reflecting superior quality in terms of experience and/or viability. Bird song is immensely involved in sexual selection, thus not‐surprisingly, it usually shows age‐dependent variation. Although the collared flycatcher Ficedula albicollis has been intensively studied for its sexual traits, and female preference for male age is expected to be strong, there is no quantitative information on age‐dependent expression of song in this species. Here, we fill this gap and, based on phenotypic correlations, we report the relationship between age and several song features. Repertoire size was consistently smaller in yearlings than in older males, but it also tended to increase after the second year of breeding. In a meta‐analysis of effect sizes using data from the literature, we found that the strength of the relationship between age and repertoire size in the collared flycatcher is similar to patterns observed in other passerines. Song rate was inversely related to age, as yearlings sang at higher rates than older males. Generally, effect sizes for the relationship between age and other song traits were of medium level on average, and had broad confidence intervals. Song traits covaried with survival in a direction suggesting that differences in song between age categories are unlikely to result from phenotype dependent survival. Our results generally support the hypothesis that song holds the potential to function as a reliable indicator of male age in the collared flycatcher. However, it seems that not all song traits are unambiguously more expressed in older males than in yearlings.     

Long-time storage of song types in birds: evidence from interactive playbacks

In studies of birdsong learning, imitation-based assays of stimulus memorization do not take into account that tutored song types may have been stored, but were not retrieved from memory. Such a 'silent' reservoir of song material could be used later in the bird's life, e.g. during vocal interactions. We examined this possibility in hand-reared nightingales during their second year. The males had been exposed to songs, both as fledglings and later, during their first full song period in an interactive playback design. Our design allowed us to compare the performance of imitations from the following categories: (i) songs only experienced during the early tutoring; (ii) songs experienced both during early tutoring and interactive playbacks; and (iii) novel songs experienced only during the simulated interactions. In their second year, birds imitated song types from each category, including those from categories (i) and (ii) which they had failed to imitate before. In addition, the performance of these song types was different (category (ii) > category (i)) and more pronounced than for category (iii) songs. Our results demonstrate 'silent' song storage in nightingales and point to a graded influence of the time and the social context of experience on subsequent vocal imitation.     

Buzzwords in Females’ Ears? The Use of Buzz Songs in the Communication of Nightingales (Luscinia megarhynchos)

Differences in individual male birds’ singing may serve as honest indicators of male quality in male-male competition and female mate choice. This has been shown e.g. for overall song output and repertoire size in many bird species. More recently, differences in structural song characteristics such as the performance of physically challenging song components were analysed in this regard. Here we show that buzz elements in the song of nightingales (Luscinia megarhynchos) hold the potential to serve as indicators of male quality and may therefore serve a communicative function. Buzzes were produced with considerable differences between males. The body weight of the males was correlated with one measure of these buzzes, namely the repetition rate of the buzz subunits, and individuals with larger repertoires sang buzzes at higher subunit-rates. A model of buzz performance constraints suggested that buzzes were sung with different proficiencies. In playback experiments, female nightingales showed more active behaviour when hearing buzz songs. The results support the idea that performance differences in the acoustic fine structure of song components are used in the communication of a large repertoire species such as the nightingale.     

Territory tenure in song sparrows is related to song sharing with neighbours, but not to repertoire size

Song repertoires may be a product of sexual selection and several studies have reported correlations of repertoire size and reproductive success in male songbirds. This hypothesis and the reported correlations, however, are not sufficient to explain the observation that most species have small song repertoire sizes (usually fewer than 10, often fewer than five song types). We examined a second important aspect of a male's song repertoire, the extent to which he shares songs with his neighbours. Song sharing has not been measured in previous studies and it may be partially confounded with repertoire size. We hypothesized that in song sparrows, Melospiza melodia, song sharing rather than repertoire size per se is crucial for male territorial success. Our longitudinal study of 45 song sparrows followed from their first year on territory showed that the number of songs a bird shares with his neighbourhood group is a better predictor of lifetime territory tenure than is his repertoire size. We also found that song sharing increases with repertoire size up to but not beyond eight to nine song types, which are the most common repertoire sizes in the population (range in our sample 5-13). This partial confound of song sharing and repertoire size may account for some earlier findings of territory tenure-repertoire size correlations in this species and other species having small- or medium-sized repertoires. Copyright 2000 The Association for the Study of Animal Behaviour.     

Hatching late in the season requires flexibility in the timing of song learning

Most songbirds learn their songs from adult tutors, who can be their father or other male conspecifics. However, the variables that control song learning in a natural social context are largely unknown. We investigated whether the time of hatching of male domesticated canaries has an impact on their song development and on the neuroendocrine parameters of the song control system. Average age difference between early- and late-hatched males was 50 days with a maximum of 90 days. Song activity of adult tutor males decreased significantly during the breeding season. While early-hatched males were exposed to tutor songs for on average the first 99 days, late-hatched peers heard adult song only during the first 48 days of life. Remarkably, although hatching late in the season negatively affected body condition, no differences between both groups of males were found in song characteristics either in autumn or in the following spring. Similarly, hatching date had no effect on song nucleus size and circulating testosterone levels. Our data suggest that late-hatched males must have undergone accelerated song development. Furthermore, the limited tutor song exposure did not affect adult song organization and song performance.     

Complex modulation of singing behavior by testosterone in an open-ended learner, the European Starling

In many temperate zone songbird species males only produce song during the breeding season, when plasma testosterone (T) levels are high. Males of some species sing throughout the year, even when T levels are low, indicating a dissociation between high T levels and song rate. Given that few studies have taken advantage of these species, we compare here song traits expressed under high versus low T concentrations and we study the role of testosterone in adult song learning in the European Starling, an open-ended learner in which repertoire size dramatically increases with age. We performed a detailed comparison of song complexity and song rate between fall and spring in 6-year-old intact male European starlings. In parallel, we investigated whether potential seasonal changes were regulated by the gonadally induced increase in plasma T, by comparing seasonal changes in intact and castrated males of the same age (castrated as juveniles during their first fall) and by subsequently experimentally elevating T in half of the castrated males. While song rate and stereotypy did not differ between intacts and castrates or between fall and spring, both groups increased their average song bout length from fall to spring, but only intact males increased their repertoire size, indicating that effects of seasonal T changes differ between song traits. Intact males overall displayed a larger song repertoire and a longer bout length than the castrates, and implantation with T caused a turnover in repertoire composition in castrates. However, as the castrates had never experienced high T levels and yet displayed a markedly higher repertoire size than that of typical yearling males, this suggests that the progressive increase of song repertoire with age in male starlings is not dependent on gonadal T, although it may be T-enhanced.     

Complex modulation of singing behavior by testosterone in an open-ended learner,the European Starling

In many temperate zone songbird species males only produce song during the breeding season, when plasma testosterone (T) levels are high. Males of some species sing throughout the year, even when T levels are low, indicating a dissociation between high T levels and song rate. Given that few studies have taken advantage of these species, we compare here song traits expressed under high versus low T concentrations and we study the role of testosterone in adult song learning in the European Starling, an open-ended learner in which repertoire size dramatically increases with age. We performed a detailed comparison of song complexity and song rate between fall and spring in 6-year-old intact male European starlings. In parallel, we investigated whether potential seasonal changes were regulated by the gonadally induced increase in plasma T, by comparing seasonal changes in intact and castrated males of the same age (castrated as juveniles during their first fall) and by subsequently experimentally elevating T in half of the castrated males. While song rate and stereotypy did not differ between intacts and castrates or between fall and spring, both groups increased their average song bout length from fall to spring, but only intact males increased their repertoire size, indicating that effects of seasonal T changes differ between song traits. Intact males overall displayed a larger song repertoire and a longer bout length than the castrates, and implantation with T caused a turnover in repertoire composition in castrates. However, as the castrates had never experienced high T levels and yet displayed a markedly higher repertoire size than that of typical yearling males, this suggests that the progressive increase of song repertoire with age in male starlings is not dependent on gonadal T, although it may be T-enhanced.     

Seasonality in song behaviour revisited: seasonal and annual variants and invariants in the song of the domesticated canary (Serinus canaria)

The song of the domesticated canary (Serinus canaria) is one of the most widely used models to study the neural correlates of behavioural plasticity and the mechanisms of female mate choice. However, only few studies have described the song behaviour in detail and monitored their changes throughout the year, and these data are restricted to the “Waterslager” strain. Here, we studied the song characteristics of the male common domesticated canary at different times of the year, the spring breeding and autumnal non-breeding season, and monitored the birds' songs up to the following breeding season. During breeding, males have increased plasma levels of testosterone, and songs are on average longer and consist of fewer non-repeated syllable types compared to the non-breeding season. When subsequent seasons are compared, song duration and the proportion of non-repeated syllable types change seasonally but not across years. Repertoire size remains constant throughout seasons although syllable types are exchanged. Syllable carry-over is significantly higher from one breeding season to the next than between the breeding and non-breeding season. Further, the repertoire of the breeding season contains more potentially sexually attractive syllable types than that of the non-breeding season. These data show that overall song structure is retained throughout the year while seasonality occurs in the temporal pattern and in repertoire composition.