Is female preference for male repertoires due to sensory bias?

There are several possible explanations for the female preference for male repertoires in birds. These males are older, and have better territories; thus there are functional reasons for females to prefer these males. However, there is an alternative explanation; females may habituate less quickly to song repertoires than single songs. I tested whether females have a non-functional, sensory bias for male song repertoires, by testing female preference for a repertoire in zebra finches (Taenopygia guttata), a species in which males possess a single stereotyped song. Females chose between a male repertoire of four different phrases created from the song phrase of one individual and that of one of those phrases repeated four times (natural zebra finch song). Females were also given a choice between the above repertoire and a song made from the phrases of four related males (''family'' stimulus). I tested female preference by training females to press a button for presentation of a song stimulus, and counting the number of button presses. Females preferred the song repertoire to a single phrase song, and did not differentiate between the repertoire and song phrases from four males. Evidence from the Estrildidae indicates that having a single song is the ancestral state for zebra finches, so the preference is not ancestral.     

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.     

Versatility and Stereotypy of Free-Tailed Bat Songs

In mammals, complex songs are uncommon and few studies have examined song composition or the order of elements in songs, particularly with respect to regional and individual variation. In this study we examine how syllables and phrases are ordered and combined, ie “syntax”, of the song of Tadarida brasiliensis, the Brazilian free-tailed bat. Specifically, we test whether phrase and song composition differ among individuals and between two regions, we determine variability across renditions within individuals, and test whether phrases are randomly ordered and combined. We report three major findings. First, song phrases were highly stereotyped across two regions, so much so that some songs from the two colonies were almost indistinguishable. All males produced songs with the same four types of syllables and the same three types of phrases. Second, we found that although song construction was similar across regions, the number of syllables within phrases, and the number and order of phrases in songs varied greatly within and among individuals. Last, we determined that phrase order, although diverse, deviated from random models. We found broad scale phrase-order rules and certain higher order combinations that were highly preferred. We conclude that free-tailed bat songs are composed of highly stereotyped phrases hierarchically organized by a common set of syntactical rules. However, within global species-specific patterns, songs male free-tailed bats dynamically vary syllable number, phrase order, and phrase repetitions across song renditions.     

What exactly is ‘local song’ in a population of ortolan buntings with a common dialect?

Previous studies have shown that ortolan buntings (Emberiza hortulana) exhibit apparent dialect variation. Neighbouring males typically share the same final phrase of a simple two-part song. Consequently, the final phrase was considered to be the dialect cue important for discriminating between males from a local population and strangers. Recently, it was shown that in an isolated and fragmented population of the ortolan buntings in Norway there was no local dialect in the above-mentioned sense. Norwegian males often had song types with different final phrases in their repertoire, and had larger repertoires and a lower level of song type sharing than in other populations. It was experimentally revealed that only local songs (L) evoked a strong response in Norwegian males, while these birds did not respond strongly to foreign (F) or hybrid songs composed of local and foreign initial and final phrases in any composition (i.e. both FL and LF songs). These results suggest that, in the Norwegian population, the final phrase of the song is not a sufficient cue for local song dialect recognition. This paper is a further study in which we tested the response of the ortolan bunting males to L, F, FL and LF songs in a typical continuous population (in Poland) in which males share the same single final phrase, i.e. have a common dialect. We found that ortolan bunting males in Poland responded with similar strength to L, FL and LF songs. The majority of males did not respond as strongly to playback of only F songs. Our results suggest that a common final phrase for a population should not be treated as the only signal of ‘locality’. Our results show clear asymmetry in response to hybrid songs with non-local phrases in different populations.     

Structure,syntax and “small-world” organization in the complex songs of California Thrashers (Toxostoma redivivum)

We describe songs of the California Thrasher (Toxostoma redivivum), a territorial, monogamous species whose complex songs are composed of extended sequences of phonetically diverse phrases. We take a network approach, so that network nodes represent specific phrases, and links or transitions between nodes describe a subgroup structure that reveals the syntax of phrases within the songs. We found that individual birds have large and largely distinct repertoires, with limited phrase sharing between neighbours and repertoire similarity decaying between individuals with distance apart, decaying also over time within individuals. During song sequences, only a limited number of phrases (ca. 15–20) were found to be actually “in play” at any given time; these phrases can be grouped into themes within which transitions are much more common than among them, a feature contributing to a small-world structure. It appears that such “small-world themes” arise abruptly, while old themes are abandoned more gradually during extended song sequences; most individual thrashers switch among 3–4 themes over the course of several successive songs, and some small-world themes appear to have specific roles in starting or ending thrasher songs.     

Song Learning in the Anna Hummingbird (Calypte anna)

Songs of wild male Anna hummingbirds (Calypte anna) consist of syllables grouped into phrases. Nearest neighbors tend to share similar syllable types, rhythms and syntax. Songs from different localities contain different syllable types, syntax and repetition indices. A male raised by hand in isolation produced a song consisting of highly variable syllable types of a wide frequency range. The song was simple in structure, and syllables were not grouped into phrases. Three males raised by hand as a group sang songs containing two stereotyped syllable types sung in alternating sequence and without phrase structure. These three males shared syllable types and syntax. The data from our study indicate that despite its relatively simple syrinx the Anna hummingbird learns syllable types, frequency, rhythm and syntax (as do oscines with their more complex syringes) during the song development process.     

Soft Song in Song Sparrows: Acoustic Structure and Implications for Signal Function

In many species of songbirds, males sometimes produce songs at distinctly lower amplitude than in normal singing. Depending on the species, these 'soft songs' may be sung in the context of female courtship, male–male aggression, or both. In song sparrows, males produce soft songs during aggressive interactions with other males, and the amount of soft song produced is the only singing behavior that can be used to reliably predict a subsequent attack by the singer. Although soft song is clearly an important signal in this species, little is known about the acoustic structure of soft song or about how that structure compares to the structure of normal 'broadcast song'. We recorded a large sample of soft songs and broadcast songs from 10 male song sparrows, and measured song amplitudes in the field while controlling the subject's distance to a calibrated microphone. We show that song sparrow males produce songs over a wide range of amplitudes, with soft songs in the range of 55–77 dB sound pressure level and broadcast songs in the range of 78–85 dB. We present evidence for two types of soft song: 'crystallized' soft songs that are broadcast repertoire song types sung at low amplitude, and 'warbled' soft songs that are not found in the broadcast repertoire. Although highly variable, warbled soft songs produced by individual birds could be grouped into song types based on spectrographic similarity. To our knowledge, a distinct repertoire of soft song types has not been previously reported for any songbird.     

Songs of Holcostethus strictus (Fabricius): a different repertoire among landbugs (Heteroptera: Pentatomidae)

Songs emitted during mating by male and female Holcostethus strictus were recorded as substrate vibrations. Spectra of the vibrational signals have a dominant frequency peak between 100 and 260 Hz and in this respect reflect the general characteristic of the family Pentatomidae. Songs of H. strictus differ from the song repertoire of the southern green stink bug Nezara viridula (L.) (Pentatomidae) in many respects. The female calling and courtship songs differ in echeme and phrase duration. The male calling song is composed of spectrally different subunits. The male courtship song is characterised by three types of spectrally and temporally different echemes. The male copulatory song is composed of echemes of two types, which constitute a phrase of less regular temporal structure. In H. strictus, males start to sing first and female songs are less complex than in N. viridula. The female calling song is evoked by male calling and does not trigger male response. The female and male courtship song phrases are superimposed on one another and we have not observed any obvious regularity in their exchange. The possible role of different songs in H. strictus is discussed and compared with that in other pentatomide landbug species.     

Structural Design Principles of Complex Bird Songs: A Network-Based Approach

Bird songs are acoustic communication signals primarily used in male-male aggression and in male-female attraction. These are often monotonous patterns composed of a few phrases, yet some birds have extremely complex songs with a large phrase repertoire, organized in non-random fashion with discernible patterns. Since structure is typically associated with function, the structures of complex bird songs provide important clues to the evolution of animal communication systems. Here we propose an efficient network-based approach to explore structural design principles of complex bird songs, in which the song networks–transition relationships among different phrases and the related structural measures–are employed. We demonstrate how this approach works with an example using California Thrasher songs, which are sequences of highly varied phrases delivered in succession over several minutes. These songs display two distinct features: a large phrase repertoire with a ‘small-world’ architecture, in which subsets of phrases are highly grouped and linked with a short average path length; and a balanced transition diversity amongst phrases, in which deterministic and non-deterministic transition patterns are moderately mixed. We explore the robustness of this approach with variations in sample size and the amount of noise. Our approach enables a more quantitative study of global and local structural properties of complex bird songs than has been possible to date.     

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.     

Mechanisms of song differentiation in introduced populations of Chaffinches Fringilla coelebs in New Zealand

Chaffinches Fringilla coelebs were introduced to New Zealand from Great Britain over 100 years ago, and since then their songs have diverged from British ones in both syllable structure and basic organization of syllable sequences. The New Zealand populations seem to be unique in that their songs have undergone differentiation of trill segments by progressive elaboration of syllable morphology and hp ascending and descending changes of pitch. Published sonagrams of British song types have significantly more trill phrases and significantly fewer syllables in the end phrase than do Yew Zealand ones. Many New Zealand song types have much elaborated end phrases and concomitantly simpler trill segments, with songs quite commonly having only one trill phrase. This reversal of complexity between the trill and end phrase in New Zealand seems to have been derived by progressive reduction of the ultimate trill phrase to one syllable, and by recombination of end phrase syllables from different song types into one compound end phrase. The significance of the increased complexity in the end phrases of many song types may relate to the sound transmission properties of dense pine forests in New Zealand, in which Chaffinches are ubiquitous. Elaborate end phrases degrade much less from reverberation in pine forests because individual syllables have more dispersed temporal patterning.
Recombination of syllables to form new song types is a major mechanism of song differentiation in New Zealand. Although whole song copying is the predominant mode of replication, very few song types in a locality sample are composed of a unique set of syllables. Rather, song types in an area are interconnected by different combinations of shared syllables, suggesting that the incorporation of some local syllables in a bird's repertoire is sufficient to signal its status as a member of a neighbourhood and also allows the evolution of broadcast complexity.     

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.     

Experimental evidence that distinct song phrases in the Grey‐cheeked Fulvetta Alcippe morrisonia permit species and local dialect recognition

Birds with small song repertoires have a limited number of song types which may serve multiple functions. The Grey‐cheeked Fulvetta Alcippe morrisonia is a non‐migratory species of lowland forests in Taiwan. Its song consists of two distinct phrases: a whistled phrase and a harmonic one. Each individual usually sings only one type of whistled phrase, and the geographical patterns of songs can be distinguished by the motif of this phrase. We proposed a dual adaptation hypothesis for the functions of these two phrases. Playback experiments including six sound stimuli (familiar whistled phrases, familiar harmonic phrases familiar complete songs, foreign whistled phrases, foreign harmonic phrases and foreign complete songs) were conducted at 12 sites in the Shoushan Nature Park of Kaohsiung, Taiwan. Grey‐cheeked Fulvettas came closer to the playback speaker and increased counter‐singing more in response to the familiar whistled phrase (local dialect) than to a foreign whistled phrase (distant dialect). However, birds responded equally to the playback of harmonic stimuli from familiar and foreign sources. We suggest that in this species, whistled phrases are used for local recognition and harmonic phrases are used for species recognition in short‐range communication.     

Testosterone Induces Sexual Release Quality in the Song of Female Canaries

We tested the sexual responsiveness of female canaries, Serinus canaria, to two sets of different types of female conspecific songs versus an adult male conspecific song. Female songs were either spontaneously emitted (‘S-songs’) or were testosterone-induced (‘T-songs’). Copulation-solicitation displays (CSD) were used as an index of female sexual response. Playbacks were performed several days before and during egg laying, a period of natural sexual responsiveness of the females to song. We demonstrated that the weaker sexual displays of female canaries were recorded to S-songs, thus suggesting that these types of female songs do not contain fully functional sexual releasers. Three T-songs elicited high levels of sexual displays, thus demonstrating that testosterone treatment may induce sexual release quality in the female songs. Study of the phonology of these three T-songs strongly suggested that special song phrases may be good candidates as powerful sexual releasers. To test the sexual value of these female song phrases, we carried out a third experiment, using hybrid songs where each of these special T-song phrase types was included in a well-known heterospecific context. Two phrases elicited high levels of sexual responses in females. Essential features of the male full song, such as broadband rapid frequency modulations and high repetition rate, are retrieved in both female song phrases. Taken together, these data demonstrate that testosterone treatment not only induces a male-like structure in the songs of females, but also induces functionally ‘male-like’ songs. This result allows features of the vocal control network of testosterone-treated females to be compared with those of adult males singing full songs, to distinguish neural correlates of testosterone-dependent full songs. However, because testosterone does not induce functionally male-like songs in all the females, neuroanatomical structure-function correlations need detailed behavioural analysis.     

Preferences and predispositions of female canaries (Serinus canaria) for loud intensity of male sexy phrases

In the present study, we tested the effect of song amplitude on intersexual relationships. To evaluate preferences and predispositions of female canaries for amplitude levels of male song, we conducted two experiments using both females raised in acoustic isolation and females raised in an aviary under 'normal' acoustic conditions. The songs used in both experiments followed the same pattern: one reactive phrase surrounded by two nonreactive phrases. The first experiment consisted of testing female preferences for weak, normal, or loud amplitudes of the reactive phrase (i.e. these reactive phrases provoke sexual stimulation in females). In a second experiment, we tested female preferences for weak, normal, or loud amplitudes of the nonreactive phrases. These two experiments allowed us to evaluate female preferences for intensity levels of reactive and nonreactive phrases. In the first experiment, all females significantly prefered loud and normal reactive phrases, whereas the second experiment showed that loud nonreactive phrases do not necessarily provoke more responses than other nonreactive phrases. Female canaries were, thus, more stimulated by the intensity level of a particular part of the song. Both females raised in acoustic isolation and in normal acoustic conditions responded in the same way. We can therefore suppose that learning, through acoustic experience, has little influence on preference development for amplitude levels.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 808–814.     

The Role of Song Phonology and Syntax in Vocal Learning Preferences in the Song Sparrow,Melospiza melodia

Are young songbirds ready to learn virtually any song, or are they predisposed to learn songs of their own species? To explore this question tests were conducted on the equipotentiality of auditory song learning stimuli in the song sparrow. 23 males reared as nestlings were exposed to tape recordings of their own and other species songs in early life and subsequent song production was analyzed for imitations. Birds exposed to natural song sparrow songs, including their fathers', and equal numbers of swamp sparrow songs, strongly preferred conspecific songs. They neither favored nor eschewed paternal songs despite having had access to them for 6–10 days as nestlings. In three other experiments synthetic songs were used in which some properties were held constant and others were systematically varied. Birds were exposed to 1–4 segmented songs varying in phrase order, tempo and syllable number, each synthesized in two versions, one from conspecific and the other from heterospecific (swamp sparrow) song syllables. With one-segmented songs (alien syntax) subjects favored conspecific over heterospecific syllable songs. Heterospecific syllables were rendered more acceptable by incorporation into two-segmented trilled songs (more song sparrow-like syntax). Heterogeneous summation of phonological and syntactical cues appeared to occur. There was also evidence of interaction between phonology and syntax. When another phrase type, the note complex, was added, in three- and four-segmented songs, a preference for conspecific syllables reappeared. Heterospecific syllables may be more readily accepted as a trilled sequence than without repetition, as in a note-complex. When phrase structure within four-segmented songs was varied, birds favored patterns most like normal conspecific song. We conclude that there are innate learning preferences in the song sparrow, based on note and syllabic structure (phonology and syllabic syntax), and temporal organization of phrases (segmental syntax), differing from those of the closely related swamp sparrow, Melospiza georgiana, in which song syntax plays no role in learning preferences.     

Song sharing by neighbourhood groups of territorial male Blackeyed Bulbuls

Lloyd, P., Hulley, P.E. & Craig, A.J.F.K. 1999. Song sharing by neighbourhood groups of territorial male Blackeyed Bulbuls. Ostrich 70 (3&4): 208–213

The territorial song of the Blackeyed Bulbul Pycnonotus barbatus is described for a population followed over two breeding seasons in suburban Grahamstown, South Africa. The Blackeyed Bulbul has a discrete and highly structured system of phrase sharing by neighbourhood groups of males. The most common repertoire size is five phrases (mean 5.5; range 2–9). In a temporally-stable song population, individual phrases are shared independently by discrete clusters of males. Neighbours show a strong tendency to share phrase types (but rarely whole repertoires), with overall song similarity decreasing rapidly with increasing distance between birds. This species is an ideal candidate for testing hypotheses concerning the adaptive value of song sharing by neighbourhood groups.     

Species Recognition by Song in the Veery (Catharus fuscescens: Aves)

Territorial male veeries (Catharus fuscescens) were presented with three series of reorganised songs to determine the functional properties of song structure. Songs consist of three parts: a low frequency initial phrase, a higher frequency middle phrase, and again a lower frequency end phrase. The latter two consist of vibrato syllables. Three series of experiments were performed, using various combinations of song syllables. In Series I, songs consisted of repetitions of a single syllable, one from each part of the song. Veeries responded to syllables from the middle and end phrases as to normal songs, both in localisation and in vocal responses. However, they showed little localisation and vocalisation responses to the introductory syllables. In Series II and III, veeries responded significantly more to songs in which syllables from the middle and end phrases were presented in proper sequence, rather than not. The minimum requirements for intraspecific recognition were determined in relation to the number and order of syllables in the song.     

Male Song Stimulation of Female Reproduction in Canaries: Features Relevant to Sexual Displays are not Relevant to Nest-building or Egg-laying

Recent studies conducted in our laboratory have demonstrated that a special type of song phrase (‘sexy’ phrases) containing bipartite syllables composed of abrupt frequency falls and short silences stimulate female canaries to solicit for copulation. The study was undertaken to determine whether sexy phrases also facilitated other aspects of the reproductive activity of the female canary, namely, nest-building and egg-laying. During the first experiment, we studied the effect of sexy and non-sexy songs on copulation solicitation displays in 1-year-old females without reproductive experience and in mature females with previous reproductive experience. We confirmed that sexy songs elicited more sexual responses than did non-sexy songs in yearlings and in mature females. During the second experiment, we studied the effect of male songs on nest-building activities and egg-laying in 1-year-old inexperienced females, and in mature, experienced females. The songs of conspecific males significantly triggered and increased nest-building behaviour in female canaries whatever their age or reproductive experience. In contrast, song effects on egg-laying were only found in young females. One-year-old inexperienced females exposed to sexy or non-sexy songs laid more eggs and laid earlier than did 1-year-old inexperienced controls; no such differences were observed in mature, experienced females. The efficiency of songs in promoting nest-building or egg-laying appeared to be unrelated to their efficiency in eliciting sexual responses. No difference was found between females exposed to sexy songs and females exposed to non-sexy songs; differences were only found between control and the two groups of song-exposed females. This result demands further experiments in order to determine whether other song phrase types may account for the stimulating effects of male song on female nest-building and egg-laying.