Song Elaboration Research Articles

Parent-offspring turn-taking dynamics influence parents’ song structure and elaboration in a singing primate

Parent-offspring interactions are essential to interpret animal social evolution and behavior, but their role in mediating acoustic communication in animals that interact vocally is still unclear. Increasing evidence shows that primate vocal communication is way more flexible than previously assumed, and research on this topic can provide further information on how the social environment shaped vocal plasticity during the evolution of the Primate order. Indris communicate through elaborated vocal emissions, usually termed songs. Songs are interactive vocal displays in which all members of the family group alternate their emissions, taking turns during chorusing events. We aimed to understand whether specific rules regulate the turn-taking of different group members and investigate the flexibility of indris’ vocal behavior when co-singing with their offspring. We found that social factors can influence the turn-taking organization in a chorus, as offspring were more likely to drop out from the parents’ duet than join in, and we speculate that overlap might signal competition by members of the same-sex. The duet between the reproductive pair was the most common type of singing organization, followed by a duet between mothers and sons and the triadic interaction between mother, father, and son. Interestingly, parents’ solo singing seems to stimulate offspring to vocalize, and we also found that mothers and fathers simplify, at least in part, song elaboration when chorusing with offspring. Our results indicate that indris can perform short-time adjustments to the number of co-emitters and their identity: our approach is advantageous in highlighting the multilevel influences on primate vocal flexibility. Moreover, it provides evidence that some aspects of our vocal plasticity were already present in the lemur lineage.

Sex role similarity and sexual selection predict male and female song elaboration and dimorphism in fairy-wrens.

Historically, bird song complexity was thought to evolve primarily through sexual selection on males; yet, in many species, both sexes sing and selection pressure on both sexes may be broader. Previous research suggests competition for mates and resources during short, synchronous breeding seasons leads to more elaborate male songs at high, temperate latitudes. Furthermore, we expect male–female song structure and elaboration to be more similar at lower, tropical latitudes, where longer breeding seasons and year‐round territoriality yield similar social selection pressures in both sexes. However, studies seldom take both types of selective pressures and sexes into account. We examined song in both sexes in 15 populations of nine‐fairy‐wren species (Maluridae), a Southern Hemisphere clade with female song. We compared song elaboration (in both sexes) and sexual song dimorphism to latitude and life‐history variables tied to sexual and social selection pressures and sex roles. Our results suggest that song elaboration evolved in part due to sexual competition in males: male songs were longer than female songs in populations with low male survival and less male provisioning. Also, female songs evolved independently of male songs: female songs were slower paced than male songs, although only in less synchronously breeding populations. We also found male and female songs were more similar when parental care was more equal and when male survival was high, which provides strong evidence that sex role similarity correlates with male–female song similarity. Contrary to Northern Hemisphere latitudinal patterns, male and female songs were more similar at higher, temperate latitudes. These results suggest that selection on song can be sex specific, with male song elaboration favored in contexts with stronger sexual selection. At the same time, selection pressures associated with sex role similarity appear to favor sex role similarity in song structure.

Urban Noise Restricts Song Frequency Bandwidth and Syllable Diversity in Bananaquits: Increasing Audibility at the Expense of Signal Quality

Anthropogenic noise can interfere with animal behavior through masking of acoustic communication. In response to masking, animals may change their acoustic signals as an apparent adjustment strategy, but this may have a drawback on signal quality. Songs and calls may show noise-dependent changes in frequency and duration, which may yield some masking avoidance, but may also constrain other acoustic parameters that might carry information about the sender. In the present study, we investigated whether noise-dependent reduction in frequency bandwidth or song duration restricted syllable diversity or song elaboration in a Neotropical songbird, the bananaquit (Coereba flaveola). We show that bananaquits sing higher frequency songs, of narrower bandwidth, in noisier territories, independent of variation in territory density, without significant variation in song duration. We also show that songs with higher minimum frequencies, narrower bandwidths, and shorter durations have on average a lower number of syllable types and higher syllable rates. This finding is in line with an acoustic restriction and may reflect a functional trade-off between audibility and signal value: higher frequencies may be more audible but less elaborate songs may weaken the message of sender quality. Consequently, noise pollution may not only alter avian communities, but also shape acoustic diversity and processes of sexual selection in urban environments.

Noise-Related Song Variation Affects Communication: Bananaquits Adjust Vocally to Playback of Elaborate or Simple Songs

Birds communicate through acoustic variation in their songs for territorial defense and mate attraction. Noisy urban conditions often induce vocal changes that can alleviate masking problems, but that may also affect signal value. We investigated this potential for a functional compromise in a neotropical songbird: the bananaquit (Coereba flaveola). This species occurs in urban environments with variable traffic noise levels and was previously found to reduce song elaboration in concert with a noise-dependent reduction in song frequency bandwidth. Singing higher and in a narrower bandwidth may make their songs more audible in noisy conditions of low-frequency traffic. However, it was unknown whether the associated decrease in syllable diversity affected their communication. Here we show that bananaquits responded differently to experimental playback of elaborate vs. simple songs. The variation in syllable diversity did not affect general response strength, but the tested birds gave acoustically distinct song replies. Songs had fewer syllables and were lower in frequency and of wider bandwidth when individuals responded to elaborate songs compared to simple songs. This result suggests that noise-dependent vocal restrictions may change the signal value of songs and compromise their communicative function. It remains to be investigated whether there are consequences for individual fitness and how such effects may alter the diversity and density of the avian community in noisy cities.

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

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

New insights from female bird song: towards an integrated approach to studying male and female communication roles.

Historically, bird song has been regarded as a sex-specific signalling trait; males sing to attract females and females drive the evolution of signal exaggeration by preferring males with ever more complex songs. This view provides no functional role for female song. Historic geographical research biases generalized pronounced sex differences of phylogenetically derived northern temperate zone songbirds to all songbirds. However, we now know that female song is common and that both sexes probably sang in the ancestor of modern songbirds. This calls for research on adaptive explanations and mechanisms regulating female song, and a reassessment of questions and approaches to identify selection pressures driving song elaboration in both sexes and subsequent loss of female song in some clades. In this short review and perspective we highlight newly emerging questions and propose a research framework to investigate female song and song sex differences across species. We encourage experimental tests of mechanism, ontogeny, and function integrated with comparative evolutionary analyses. Moreover, we discuss the wider implications of female bird song research for our understanding of male and female communication roles.

Transcontinental latitudinal variation in song performance and complexity in house wrens (Troglodytes aedon).

There is growing interest in latitudinal effects on animal behaviour and life history. One recent focus is on birdsong, which is hypothesized to be more elaborated or complex in the north temperate zone compared with the tropics. Current evidence is mixed and based on cross-species comparisons, or single species with restricted distributions. We circumvent these limitations using a transcontinental sample of 358 songs from house wrens (Troglodytes aedon) at 281 locations spanning more than 100° of latitude (52° N-55° S) across the Americas. We found a significant latitudinal gradient in several basic elements of song performance and complexity between north temperate and tropical populations. Furthermore, we document convergence in song patterns between populations at higher latitudes in the Northern and Southern Hemispheres. Effects were strongest for the number of elements in a song, and the rate of element production, both increasing towards the poles, with similar but weaker effects for other song dimensions (e.g. number of unique elements, trills and trill rate). We consider possible causes related to variable habitats and morphology, concluding that the shorter breeding seasons at higher latitudes in both hemispheres may favour greater song elaboration to mediate territory competition and mate choice.

Female song and vocal interactions with males in a neotropical wren

Bird song is thought to function primarily in same-sex competition, mate attraction, and reproductive stimulation of a partner. However, these conclusions are based largely on studies of the song of male birds in north-temperate species. We investigate female song in a Neotropical wren, Thryophilus pleurostictus, using observations and experiments to test the function of female song. Female banded wrens sang much less often than males, their songs were shorter, and their repertoire of song types was smaller. Females did not seem to sing for same-sex competition for resources or mates: female song rate did not increase in response to simulated intrusion, and females sang in response to less than one-third of playbacks simulating territorial intrusion by either unpaired or paired females. Territorial defense is important for both survival and reproduction in species that occupy all-purpose territories year-round, but female involvement in territorial defense was limited. Females were more likely to approach simulated intruders when their partner approached more closely, and were closer to their partner during playback simulating a pair of intruders, perhaps contributing to defense jointly with their partner. Females did not appear to use song to attract males for mating: only 25% of females sang in response to playback simulating an unpaired male during the nest-building period, and they were less likely to sing shortly before laying when they were more likely to be fertile. Female song in banded wrens seemed to be used primarily for communicating with their breeding partner: female songs overlapped or began within one second of a song by their partner more often than expected by chance, and male vocal behavior changed in response to song by their partner. However, the low rate of female song in banded wrens suggests this function does not select for song elaboration, consistent with the view that same-sex competition is the main driver of female song elaboration.

Migration and song elaboration in wood-warblers (Geothlypis)

ABSTRACT Although some comparative studies of oscine songbirds have found that long-distance migration is positively correlated with elaborate songs, an analysis of singing by species in the genus Geothlypis (Parulidae) found no evidence of such a correlation. The migratory species in the genus sing relatively simple songs, whereas the singing of nonmigratory species varies; some species have simple songs, and others have more elaborate songs. Elaborate songs are found in the nonmigratory species G. semiflava, G. aequinoctialis, and G. poliocephala. For example, in Costa Rican populations of these species, songs are longer, contain more notes, have greater note-type diversity, and (in G. semiflava and G. aequinoctialis) have more phrase types than the songs of the migratory Geothlypis species. However, in other nonmigratory species (G. nelsoni, G. flavovelata, G. speciosa, and G. rostrata), the duration, note count, and note-type diversity of songs are similar to those of the migratory species. Thus, ther...

Age‐dependent song changes in a closed‐ended vocal learner: elevation of song performance after song crystallization

Birdsong is a sexual signal that serves as an indicator of male quality. There is already abundant evidence that song elaboration reflects early life‐history because early developmental stress affects neural development of song control systems, and leaves irreversible adverse effects on song phenotypes. Especially in closed‐ended vocal learners, song features crystallized early in life are less subject to changes in adulthood. This is why less attention has been paid to lifelong song changes in closed‐ended learners. However, in the eyes of female birds that gain benefits from choosing mates based on male songs, not only past but also current conditions encoded in songs would be meaningful, given that even crystallized songs in closed‐ended learners would not be identical in the long term. In this study, we examine within‐individual song changes in the Java sparrow Lonchura oryzivora, with the aim of shedding light on the relationship between song and long‐term life history. Specifically, we compared song length, tempo, and song complexity measures between the point just after song crystallization and around 1 yr later, and also compared those traits between fathers and sons to clarify the effect of vocal learning. While it is not surprising that song complexity did not differ depending on age or between fathers and sons, we found that song length and tempo increased with age. Follow‐up analyses have revealed that frequency bandwidth and peak frequency of song notes also elevated with age. Our results show that song performance related to motor skills can be improved even after song crystallization. We also suggest that song performance in closed‐ended vocal learners gives a reliable clue for mate choice by reflecting male quality with aging.

Birdsong, sexual selection, and the flawed taxonomy of canaries, goldfinches and allies

Sexual signals are often diagnostic of closely related species, and their taxonomic value at higher phylogenetic levels is often discussed. Complex signals such as birdsong are potentially rich in information regarding evolutionary history. However, natural or sexual selection, if pervasive enough, might distort that information. This has seldom been evaluated. We tested how ecological effects impact on the phylogenetic information of birdsong using a real case of flawed taxonomy: the reciprocal polyphyly of the genera Serinus and Carduelis. Major axes (principal components) of between-species song variation were not related to body size or habitat type. The main axis of song variation, reflecting syllable complexity and song length, was related to ecological indexes of sexual selection (latitude and ecological breadth), resulting in a positive latitudinal gradient in song elaboration. Song significantly supported monophyly over polyphyly for these genera, despite some song data having high phylogenetic signal (relative to the correct, polyphyletic phylogeny). Thus, the presence of high phylogenetic signal did not guarantee reliable taxonomic information at the genus level. The misleading phylogenetic information appears related to the latitudinal gradient in song elaboration plus the traditional Serinus and Carduelis spp. differing in latitude. We conclude that selection, in this case sexual selection, can pervasively distort the phylogenetic information even of complex and potentially information-rich signals.

Uncorrelated evolution between vocal and plumage coloration traits in the trogons: a comparative study

The costs of bird song incurred in a diversity of ways may result in trade-offs in the production and maintenance of elaborate plumage ornaments. In this paper, we examine evolutionary trade-offs between acoustic and visual signalling in trogon birds (Trogonidae). Using multiple regressions with phylogenetically independent contrasts, we found that interspecific variation in male plumage coloration was not significantly predicted by song traits (reduced by PCA) or altitude. Although plumage coloration is expected to decrease with increases in song elaboration, both groups of variables were not related. Given that song and plumage coloration traits are likely targets of sexual selection, we also examined their relationships with sexual plumage dimorphism. We found that male carotenoid-derived coloration was positively related to sexual plumage dimorphism, suggesting that sexual selection on male carotenoid-derived coloration may be stronger than on melanin- or structurally based coloration, or than on acoustic traits. Comparative studies on other bird families accounting for the effects of phylogeny as well as environmental covariates are required to test the generality of our findings in trogons.

Ecological gradient of sexual selection: elevation and song elaboration in finches

Ecological gradients in natural and sexual selection often result in evolutionary diversification of morphological, life history, and behavioral traits. In particular, elevational changes in habitat structure and climate not only covary with intensity of sexual selection in many taxa, but may also influence evolution of mating signals. Here we examined variation in courtship song in relation to elevation of breeding across cardueline finches-a subfamily of birds that occupies the widest elevational range of extant birds and shows extensive variation in life histories and sexual selection along this range. We predicted that decrease in sexual selection intensity with elevation of breeding documented in this clade would result in a corresponding evolutionary reduction in elaboration of courtship songs. We controlled for the effects of phylogeny, morphology, and habitat structure to uncover a predicted elevational decline in courtship song elaboration; species breeding at lower elevations sang more elaborated and louder songs compared to their sister species breeding at higher elevations. In addition, lower elevation species had longer songs with more notes, whereas frequency components of song did not vary with elevation. We suggest that changes in sexual selection account for the observed patterns of song variation and discuss how elevational gradient in sexual selection may facilitate divergence in mating signals potentially reinforcing or promoting speciation.

Relations between song repertoire size and the volume of brain nuclei related to song: comparative evolutionary analyses amongst oscine birds

Song and brain structure are compared amongst 41 species of oscine birds by using the method of independent evolutionary contrasts. We find a significant correlation between the relative volume of the song control centre, the high vocal centre (HVC), and the number of song types typically found in the repertoire. Relative HVC volume is not correlated with the number of different syllable types per song bout. The relative volume of a second song nucleus, area X, is not significantly correlated with either measure. Relative HVC volume is uncorrelated with relative volume of the hippocampus, a brain area involved in other forms of memory. This is the first evidence for repeated independent evolution of an association between complexity of learned song and the relative volume of one of the song control nuclei though to be involved in song learning.

Sexual selection and the evolution of bird song: A test of the Hamilton-Zuk hypothesis

Hamilton and Zuk (1982) suggested that secondary sexual characters evolve because they allow females to assess a potential mate's ability to resist parasites. A prediction of this theory is that the degree of elaboration of secondary sexual characters should be positively correlated with parasite load across species. In support of their hypothesis, Hamilton and Zuk reported a correlation across North American passerine species between haematozoa prevalence and both brightness and song “complexity and variety”, scored on a subjective six point scale. Here we show that this relationship is confounded by phylogenetic associations. We use quantitative data on song duration, inter-song interval, song continuity, song rate, song versatility, and song and syllable repertoire size for 131 species of European and North American passerines to test the Hamilton-Zuk hypothesis. Across species, there are significant negative relationships between haematozoa prevalence and song continuity, contrary to the direction predicted by Hamilton and Zuk. In accordance with their prediction, there is a positive correlation with song versatility. However, these relationships come about through taxonomic associations: within taxa there are no consistent relationships between any of the song variables and haematozoa prevalence. None of the other song variables correlate with haematozoa prevalence. We conclude that there is no evidence of an association between song elaboration and parasites.