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.     

Testosterone increases repertoire size in an open-ended learner: An experimental study using adult male European starlings (Sturnus vulgaris)

Song in songbirds is a learned secondary sexual behavior, first acquired during a sensitive phase of juvenile development, which is affected by hormones such as testosterone (T). While the latter has received much attention, the potential involvement of T in the adult repertoire changes observed in a number of species is much less understood. Yet, this may prove essential to understand the role of song as a sexually selected trait. We therefore performed a T-implantation experiment during the non-breeding season (when T is basal), using adult male European starlings (Sturnus vulgaris), a songbird species in which song repertoire size (and composition) changes seasonally and increases with age. Repertoire size increased rapidly in T-males, but not in control males, indicating a role for T in repertoire size changes. This increase resulted from a lower proportion of dropped song types in T-males than in control males, while the proportion of added song types did not differ between both groups. Interestingly, the observed repertoire turnover (adding and removing song types from the repertoire) in both groups, suggests that elevated plasma T levels were not essential for changes in repertoire composition (contrary to repertoire size). Finally, T-males (but not control males) significantly increased their song rate, while neither group showed a significant change in their song bout length and phrase repetition rate. Taken together, our results suggest a role for T in adult song learning and provide new insights into the information content of repertoire size and song bout length as sexually selected traits.     

Testosterone increases repertoire size in an open-ended learner: An experimental study using adult male European starlings (Sturnus vulgaris)

Song in songbirds is a learned secondary sexual behavior, first acquired during a sensitive phase of juvenile development, which is affected by hormones such as testosterone (T). While the latter has received much attention, the potential involvement of T in the adult repertoire changes observed in a number of species is much less understood. Yet, this may prove essential to understand the role of song as a sexually selected trait. We therefore performed a T-implantation experiment during the non-breeding season (when T is basal), using adult male European starlings (Sturnus vulgaris), a songbird species in which song repertoire size (and composition) changes seasonally and increases with age. Repertoire size increased rapidly in T-males, but not in control males, indicating a role for T in repertoire size changes. This increase resulted from a lower proportion of dropped song types in T-males than in control males, while the proportion of added song types did not differ between both groups. Interestingly, the observed repertoire turnover (adding and removing song types from the repertoire) in both groups, suggests that elevated plasma T levels were not essential for changes in repertoire composition (contrary to repertoire size). Finally, T-males (but not control males) significantly increased their song rate, while neither group showed a significant change in their song bout length and phrase repetition rate. Taken together, our results suggest a role for T in adult song learning and provide new insights into the information content of repertoire size and song bout length as sexually selected traits.     

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.     

Sexual Differences in the Responses of Captive Great Reed Warblers (Acrocephalus arundinaceus) to Variation in Song Structure and Repertoire Size

The function and evolution of repertoires in songbirds is still not clearly understood. This study involves the great reed warbler Acrocephalus arundinaceus and playback of recorded song to both captive males and females under controlled laboratory conditions. Females were implanted with oestradiol and the copulation solicitation display used as an index of sexual response. Males were not implanted and crest-raising used as an index of aggressive response. Two types of songs were used (short and long) and two sizes of repertoire (high and low). Females only responded to long songs, and responded significantly more to high than low repertoires. It was concluded that long songs have a sexual function and that high repertoires are the result of intersexual selection through female choice. Males responded to short and long songs, but did not discriminate between high and low repertoires. Adding a live male to the experiment made no difference to female responses but males responded significantly more. When field studies are also taken into consideration the overall conclusion is that song quality and repertoire size is particularly important in female choice, and that short songs and a male presence are more important in aggressive behaviour. Integration of both field and laboratory studies on males and females should lead to more rigorous testing of hypotheses concerning the function and evolution of bird songs.     

Disentangling Relations among Repertoire Size,Song Rate,Signal Redundancy and Ambient Noise Level in European Songbird

The majority of male songbirds have small repertoires and sing with eventual variety; that is, they present one song type several times before switching to the next one. Several hypotheses have been proposed to explain this phenomenon. The antiexhaustion hypothesis argues that song‐type switching prevents muscle fatigue in the syrinx. The signal redundancy hypothesis suggests that repeating the same signal increases transmission success. Here, we have studied the song behaviour of the chaffinch, Fringilla coelebs, a common Eurasian species in which the males sing a few different song types and provide eventual variety. We tested different hypotheses to explain the temporal organisation of song output (repertoire size, song rate, bout duration, etc.) as a function of ambient noise by comparing birds from the same macrogeographic region in which the birds live either in a noisy town (n = 71) or in a quieter forest habitat (n = 68). Contrary to the prediction of the signal redundancy hypothesis and the results of earlier work on chaffinches living close to noisy streams, we found no significant differences in song characteristics between the town and forest populations. Our results support the antiexhaustion hypothesis because males with larger repertoires were able to sing with a significantly higher rate due to faster switching between different song types and producing shorter bouts. Sample size or population differences between our study and earlier investigations of the same species may explain the inconsistency with previous findings. Future studies should focus on determining the relations between song organisation and the directly measured quality of males and females' choice using, preferably, a longitudinal approach.     

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.     

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.     

Song repertoire size varies with HVC volume and is indicative of male quality in song sparrows (Melospiza melodia)

Complex birdsong is a classic example of a sexually selected ornamental trait. In many species, females prefer males with large song repertoires, possibly because repertoire size is limited by the size of song control nuclei which reflect developmental success. We investigated whether song repertoire size was indicative of brain area and male quality in song sparrows (Melospiza melodia) by determining if repertoire size was related to the volume of song control nucleus HVC, as well as several morphological, immunological and genetic indices of quality. We found that males with large repertoires had larger HVCs and were in better body condition. They also had lower heterophil to lymphocyte ratios, indicating less physiological stress and a robust immune system as measured by the number of lymphocytes per red blood cell. Song repertoire size also tended to increase with neutral-locus genetic diversity, as assessed by mean d2, but was not related to internal relatedness. Our results suggest several mechanisms that might explain the finding of a recent study that song sparrows with large song repertoires have higher lifetime fitness.     

The evolution of song repertoires and immune defence in birds

Song repertoires (the number of different song types sung by a male) in birds provide males with an advantage in sexual selection because females prefer males with large repertoires, and females may benefit because offspring sired by preferred males have high viability. Furthermore, males with large repertoires suffer less from malarial parasites, indicating that a large repertoire may reflect health status. We hypothesize that sexual selection may cause a coevolutionary increase in parasite virulence and host immune defence because sexual selection increases the risk of multiple infections that select for high virulence. Alternatively, a female mate preference for healthy males will affect the coevolutionary dynamics of host-parasite interactions by selecting for increased virulence and hence high investment by hosts in immune function. In a comparative study of birds, repertoire size and relative size of the spleen, which is an important immune defence organ, were strongly, positively correlated accounting for almost half of the variance. This finding suggests that host-parasite interactions have played an important role in the evolution of song repertoires in birds.     

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.     

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.     

Song Perception in the Song Sparrow: Importance of Different Parts of the Song in Song Type Classification

Song sparrow (Melospiza melodia) males have repertoires of 6 to 12 song types, each song consisting of four or five contrasting phrases. Song types vary both within individuals (a singer varies successive renditions of a type within a singing bout) and between individuals (neighbors sing slightly different versions of a type), and in both cases this variation is concentrated in the later parts of the song. We speculated that type variation in the later parts of the song might be correlated with a perceptual tendency to classify song types primarily on the basis of the earlier parts of the song. We tested this hypothesis using a laboratory conditioning technique, supplemented by a field playback experiment. In the lab, song sparrows and white crowned sparrows were trained to discriminate between two distinct song types. In test sessions, they were presented with hybrid songs synthesized from varying proportions of the two training song types. For most birds and most song pairs, song classification was influenced by all parts of the song, with the beginning elements being most important. The field playback experiment used song type matching as the response measure. Each subject was presented a hybrid stimulus song consisting of the first half of one of his song types and the second half of another of his song types. Subjects replied with both ‘front matches’ and ‘back matches’ (60% were front matches).     

Directional Sexual Selection Cannot Explain Variation in Song Repertoire Size in the New World Blackbirds (Icterinae)

Hypotheses that song repertoire evolves by way of directional sexual selection for large repertoires were examined with a comparative study of the avian subfamily Icterinae (Passeriformes: Emberiznae). Song repertoire size varies between closely related icterine species, so phylogenetic effects cannot explain repertoire size variation. Overall, song repertoire is not associated with polygyny or with presence or absence of territoriality. Different associations of repertoire with mating system are observed in different icterine groups. Either the directional hypotheses do not apply to the evolution of song repertoire in the Icterinae, or different processes of directional sexual selection have occurred in different icterine groups.     

Song‐type sharing in a population of Song Sparrows in the eastern United States

Male Song Sparrows (Melospiza melodia) sometimes interact with neighboring territory owners by song‐type matching or repertoire matching. In some Song Sparrow populations, levels of song sharing are high and most neighbors can interact by matching, but levels of song sharing are much lower in other populations, limiting the degree to which males can match their neighbors. One explanation for variation in sharing levels is that the importance of song‐type and repertoire matching, and therefore the extent of song sharing, varies geographically in North America, being greater in western populations than eastern populations. However, to date, two studies of eastern populations have provided conflicting evidence concerning levels of song sharing by Song Sparrows. Thus, we measured sharing of whole songs and introductory phrases of songs between males with adjacent territories in another population of Song Sparrows in the eastern United States (North Carolina), near the eastern and southern limits of the species’ breeding range. Males (N = 17) in our study shared an average of only 8.7% of their song types with neighbors, and more than half of neighbor pairs shared no whole songs. Sharing of introductory phrases was more common (mean = 22.8%). The level of whole song sharing in our study is the second lowest yet reported for any Song Sparrow population, supporting the hypothesis that sharing is generally lower in eastern than in western populations.     

Are there age-related differences in the song repertoire size of Eurasian blackbirds?

Most oscine bird species possess a repertoire of different song patterns, and repertoire size is thought to signal aspects of male quality. As age is assumed to be related to male quality in terms of experience and/or viability, repertoire size may be expected to reflect male age. Here, we investigated the relationship between repertoire size and age (yearlings or older) in Eurasian blackbirds, Turdus merula, a species with a large repertoire delivered in a highly variable manner. We found that older males tended to have larger repertoires than yearlings though the two age groups overlapped considerably. Thus, compared to other species with large repertoires, age-related differences in repertoire size seem rather small in male Eurasian blackbirds. We also compared repertoires of three individuals in two successive years (as yearlings and in the year following) and found a large element turnover. Our investigation revealed that this turnover was almost complete in the quiet terminating twitter part of the song. Such turnover may allow a young bird to adjust his repertoire to his neighbours?? repertoires, which could be useful for song matching interactions.     

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.     

Extrapair paternity in chestnut-sided warblers is correlated with consistent vocal performance

The elaborateness of many bird songs is commonly presumed tohave evolved under the influence of sexual selection by femalemate choice. Thus, aspects of acoustic diversity, such as songrepertoire size, are seen as likely targets of female choice.In many songbird species with song repertoires, however, therepertoires are small. In such species, female choice mightbe based on song features other than, or in addition to, songdiversity. To investigate this conjecture, I assessed singingand paternity in a population of chestnut-sided warblers (Dendroicapensylvanica), a species in which song repertoires are of modestsize. Twenty-two song traits were evaluated to determine whichones best predicted male extrapair reproductive success. Thecandidate traits encompassed measures of song diversity (e.g.,song repertoire size), gross-scale song performance (e.g., singingrate), and fine-scale song performance (e.g., variability amongsongs in a bout). Regression analysis revealed that the bestpredictor of extrapair success was singing with little variability.In particular, the most successful males sang with consistentpitch and timing, as well as high pitch. The greater extrapairsuccess of males with more consistent vocal performance maybe due to female preference for such performance, which couldbe an indicator of male quality.     

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.     

Age- and behavior-related variation in volumes of song control nuclei in male European starlings

There is considerable interindividual variation in the volumes of song control nuclei. Sex and physiological condition appear to contribute to these differences; however, these factors alone do not account for all of the variation. Studies have attempted to relate differences in song behavior (i.e., song repertoire size) to variation in song nucleus volume, but have met with mixed success. In this article, two studies are presented that used male European starlings (Sturnus vulgaris) to explore the relationship between song nuclei volumes and age-related differences in song behavior and interindividual variation in song behavior in adults. The results of the first study showed that song repertoire size and song bout length were significantly greater in older adult than in yearling males. In addition, the volumes of the high vocal center (HVC) and nucleus robustus archistriatalis (RA) were significantly larger in older adults than yearlings. Area X of the parolfactory lobe did not differ significantly in volume between the two age classes. In the second study, both HVC and RA volume correlated positively with song bout length but not repertoire size among adult birds. Based on these results a new hypothesis is presented that states that variation in song nuclei volumes in starlings relates more to the amount of song produced than to the number of song types stored in memory. © 1996 John Wiley & Sons, Inc.