Vocal performance influences male receiver response in the banded wren

In a variety of songbirds the production of trilled song elements is constrained by a performance tradeoff between how fast a bird can repeat trill units (trill rate) and the range of frequencies each unit can span (frequency bandwidth). High-performance trills serve as an assessment signal for females, but little is known about the signal value of vocal performance for male receivers. We investigated the relationship between trill rate and frequency bandwidth in banded wren (Thryothorus pleurostictus) songs. Trilled song elements showed the same performance tradeoff found in other passerines and individuals differed in performance of some trill types. We tested the hypothesis that males of this species assess each other based on trill performance with a two-speaker experiment, in which territory owners were presented with alternating renditions of the same song type manipulated to differ in trill rate. Subjects were significantly more likely to approach the faster trill stimulus first. However, subjects that received trill types closer to the performance limit spent less time close to the fast speaker. Our results show that male banded wrens discriminate and respond differently to songs based on their vocal performance. Thus, performance of physically challenging songs may be important in intra- as well as inter-sexual assessment.     

Song is a reliable signal of bill morphology in Darwin's small tree finch Camarhynchus parvulus, and vocal performance predicts male pairing success

The shape and movement of the vocal tract are known to influence bird song. Current theory predicts that large bill and body size are correlated with low frequency song and slow trill rate. It is also widely accepted that song characteristics are important for mate choice by females. We investigated the relationship between bill morphology, song characteristics, and pairing success in Darwin's small tree finch Camarhynchus parvulus , on the Galapagos Islands. Contrary to predictions from a previous cross-species study on Darwin's finches, we found that individuals with larger bill size produced songs with slow trill rate, high dominant frequency, and broad frequency bandwidth, indicating that song is a reliable signal of bill morphology. Vocal performance as indicated by the deviation from an upper performance limit was higher in paired than unpaired males. Pairing was not skewed in favour of a particular bill size, and both small and large billed males that sang high performance song had high pairing success. The reliable signalling function of song has implications for female choice and territorial defence, given that both females and conspecific competitors can assess the relative size of males' bills through song, while females may use vocal performance as a signal of male quality.     

Red-winged blackbirds Ageliaus phoeniceus respond differently to song types with different performance levels

Physical constraints on animal signals can have important consequences for communication. For bird songs that include a trill, performance is physically constrained by a trade-off between the rate at which notes are repeated in the trill and the sound frequency range covered, such that a trill cannot exceed a certain frequency range for a given note repetition rate. Producing trills that are closer to this performance limit is presumably more challenging for birds than producing trills that are farther from it. Male red-winged blackbirds Ageliaus phoeniceus have repertoires of two to eight song types containing trills that span a range of performance levels. We determined the approximate trill performance maximum for a population of red-winged blackbirds, then conducted playback experiments to measure the responses of territorial males to song types with either high or low performance levels relative to this limit. Males responded significantly more strongly to songs containing low performance trills. Our results show that male red-winged blackbirds can discriminate between different song performance levels, suggesting that vocal proficiency plays a role in male-male interactions.     

CORRELATED EVOLUTION OF BEAK MORPHOLOGY AND SONG IN THE NEOTROPICAL WOODCREEPER RADIATION

Mating signals may diversify as a byproduct of morphological adaptation to different foraging niches, potentially driving speciation. Although many studies have focused on the direct influence of ecological and sexual selection on signal divergence, the role of indirect mechanisms remains poorly understood. Using phenotypic and molecular datasets, we explored the interplay between morphological and vocal evolution in an avian radiation characterized by dramatic beak variation, the Neotropical woodcreepers (Dendrocolaptinae). We found evidence of a trade-off between the rate of repetition of song syllables and frequency bandwidth: slow paced songs had either narrow or wide frequency bandwidths, and bandwidth decreased as song pace increased. This bounded phenotypic space for song structure supports the hypothesis that passerine birds face a motor constraint during song production. Diversification of acoustic characters within this bounded space was correlated with diversification of beak morphology. In particular, species with larger beaks produced slower songs with narrower frequency bandwidths, suggesting that ecological selection on beak morphology influences the diversification of woodcreeper songs. Because songs in turn mediate mate choice and species recognition in birds, these results indicate a broader role for ecology in avian diversification.     

Directional female preference for an exaggerated male trait in canary (Serinus canaria) song

Motor constraints on vocal production impose a trade-off between trill rate and frequency bandwidth within birdsong. We tested whether domesticated canary (Serinus canaria) females, reared either in acoustic isolation or in aviary conditions, had a preference for broad bandwidth songs with artificially increased syllable rates. The copulation solicitation display (CSD) was used as an index of female preference. As predicted, both naive and experienced females were especially responsive to syllables with a broad bandwidth emitted at an artificially increased rate. Female preference for supernormal stimuli provide support for the honest-signalling hypothesis and our results are consistent with recent findings indicating that production of song phrases maximizing both bandwidth and syllable rate may be a reliable indicator of male physical or behavioural qualities. We suggest that female preference for vocal emissions, which simultaneously maximize these two parameters, could be a widespread pattern within songbirds.     

Variation in Vocal Performance in the Songs of a Wood-Warbler: Evidence for the Function of Distinct Singing Modes

Male North American wood‐warblers (family Parulidae) subdivide their song repertoires into two different categories, or modes, of singing (first and second category songs). These two modes are thought to be specialized for interacting with females and males, although the data are inconclusive. I conducted an acoustic analysis of the song types used by yellow warblers (Dendroica petechia) for type I (first category) and type II (second category) singing to ask whether there are consistent structural differences between them which could provide insight into how they might function as separate signals. I found that type I songs are performed closer to the upper boundary of a song performance limit, measured in terms of the difficulty of production, compared with type II songs. By contrast, the performance of specific song types did not depend on whether they were used for type I singing vs. type II singing by different males. In addition, type I songs had a greater amplitude increase across the first two syllables compared with type II songs. There was no relationship between the performance of type I or type II songs and male condition. These results suggest that wood‐warblers might subdivide their song repertoire into distinct categories to highlight the relative vocal performance of their songs.     

Overlapping and matching in the song contests of black-capped chickadees

During vocal contests, animals alter both the timing and the patterning of responses to their opponents. Time-specific responses (such as overlapping an opponent's song) and pattern-specific responses (such as matching the type of an opponent's song) may reveal information about interacting animals. Here we explore the consequences of overlapping and frequency matching during song contests of male black-capped chickadees, Poecile atricapillus. Using interactive playback, we engaged birds of high and low dominance status in vocal interactions with a simulated territorial intruder. The playback intruder either overlapped or avoided overlapping the subjects' songs and either matched or avoided matching the frequency of the subjects' songs. Individuals who were overlapped by the playback intruder showed higher variability in their song length and song timing than individuals who were not overlapped. Individuals who were frequency-matched by the playback intruder responded with more agitated responses (more flights and passes over the speaker and closer distances of closest approach) and spent more time farther away from the loudspeaker. We argue that the timing of song delivery and the choice of song type are distinct functional components of vocal interactions, where overlapping and matching songs are threatening signals that have separate consequences for opponent behaviour and song performance. High-ranking males responded at greater distances from the loudspeaker in all treatments and responded with lower agitation levels than low-ranking males. We demonstrate that males of different quality show different behavioural responses to territorial intruders, where males of high status appear reluctant to engage an intruder as intensely as males of low status.     

Inferring performance in the songs of dark-eyed juncos (Junco hyemalis)

Within bird species, songs differ in their attractiveness tofemales or effectiveness in male–male interactions. Somesongs are more difficult to sing than others, and receiversmay use a singer's performance of difficult songs as a meansfor evaluating the quality of the singer. The concept of songperformance aims at quantifying how physiologically demandingare different songs. Using variation between song types of dark-eyedjuncos, Junco hyemalis, we show that some song traits tradeoff with costly aspects of song output—short intervalsbetween syllables or loud sound amplitude—suggesting thatthose traits are difficult to sing. First, after controllingfor other traits, long syllables require longer intervals forrecovery. This supports the idea that a measure of "respiratoryperformance" could be based on the relative lengths of syllablesand intervals. Second, some syllable traits trade off stronglywith sound amplitude, suggesting that these traits may be difficultto sing at high amplitudes. The ratio of frequency bandwidthand trill rate has been used to infer performance in other birdspecies, but we found no evidence that frequency bandwidth tradesoff with any aspect of song output in the junco. The negativeassociation of bandwidth with trill rate may instead be a passiveconsequence of syllable length, with longer syllables randomlyaccumulating frequency modulation. We conclude that bird receiversmay best evaluate how well a song is performed if they integratemultiple cues and discuss how researchers may similarly devisemeasures of song performance.     

The predictive value of trill performance in a large repertoire songbird,the nightingale Luscinia megarhynchos

In animal communication, elaborate signals have been shown to be under sexual selection and often to reliably indicate a signaler's quality, condition, or motivation. For instance, the performance of physically challenging signals such as trills – i.e. rapidly repeated elements of broad frequency bandwidth – is considered to reflect signaler quality. Nightingales Luscinia megarhynchos are renowned for their outstanding song repertoire sizes, and most songs include a variety of complex trills. In the present study, we examined whether performance of trills can reliably reflect male quality. We show that vocal performance of trills predicts the age of a male. Older males sang trills that were closer to the performance limit than did younger males. Moreover, males with narrower beaks sang more consistent trills than did males with wider beaks. Vocal performance of trills, however, did not significantly predict other measures of biometric quality such as body size or body condition of the males. The findings suggest that receivers could benefit from the predictive value of physically demanding song traits in assessing age as an important quality component of potential mates or rivals. Particularly in species with high singing versatility, signaler assessment based on readily assessable structures may be adaptive, as this will allow receivers to quickly gather relevant information about the singer without attending to the full song repertoire.     

Morphological adaptation influences the evolution of a mating signal

Theory predicts that forces of natural selection can reduce the intensity of sexually selected traits. In this study, I investigate how morphological adaptation to feeding ecology influences a mating signal. In birds, changes in feeding ecology can cause rapid divergence in bill morphology. Because bills are also important for song production, feeding ecology may influence song divergence. During song, birds can rapidly change vocal tract resonance using bill movement, yet are constrained in rate and magnitude of bill movements resulting in a trade-off between trill rate and frequency bandwidth. Male swamp sparrows vary in their ability to produce rapid, broad-band trills and females prefer more physically demanding songs. Populations of swamp sparrows adapted to the feeding ecology of tidal marshes have larger bills than inland populations. Larger bills should increase the constraints of producing rapid, broad-band trills allowing for a test of how changes in feeding ecology affect a feature of song used in mate choice. I found significant differences in acoustic features of song consistent with the hypothesis that coastal males are less able to meet the physical demands of song production because of the constraints of having larger bills. As possible compensation for decreases in song performance, coastal populations exhibit an increase in song complexity. These changes support the current model of how motor constraints influence song production and suggest a mechanism by which feeding ecology can influence signal evolution.     

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.     

Male Songbird Indicates Body Size with Low-Pitched Advertising Songs

Body size is a key sexually selected trait in many animal species. If size imposes a physical limit on the production of loud low-frequency sounds, then low-pitched vocalisations could act as reliable signals of body size. However, the central prediction of this hypothesis – that the pitch of vocalisations decreases with size among competing individuals – has limited support in songbirds. One reason could be that only the lowest-frequency components of vocalisations are constrained, and this may go unnoticed when vocal ranges are large. Additionally, the constraint may only be apparent in contexts when individuals are indeed advertising their size. Here we explicitly consider signal diversity and performance limits to demonstrate that body size limits song frequency in an advertising context in a songbird. We show that in purple-crowned fairy-wrens, Malurus coronatus coronatus, larger males sing lower-pitched low-frequency advertising songs. The lower frequency bound of all advertising song types also has a significant negative relationship with body size. However, the average frequency of all their advertising songs is unrelated to body size. This comparison of different approaches to the analysis demonstrates how a negative relationship between body size and song frequency can be obscured by failing to consider signal design and the concept of performance limits. Since these considerations will be important in any complex communication system, our results imply that body size constraints on low-frequency vocalisations could be more widespread than is currently recognised.     

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.     

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.     

Song duration mediates responses of territory owner in a songbird species with a small song repertoire

Song is a sexually selected trait that is involved in mate attraction and territory defence in birds. Songs may convey information about different male quality components. They are flexible in terms of frequency, amplitude, and duration. Although changes in song duration are common, the function of this behaviour has been studied less strongly. It is known that song duration changes within a singing bout and may provide information about aggressive motivation. We tested whether the elongation and shortening of songs affects the responses of territorial ortolan bunting males to neighbour song playback. If changing song duration signals level of aggressiveness, then songs differing solely in duration may elicit behavioural responses of different strength. We performed two tests with different males assigned to two experimental groups and measured approaching and vocal response. In Experiment 1, we tested 18 males, which responded to the playbacks of elongated and normal (control) neighbour songs. In Experiment 2, we tested 17 males, which responded to the playbacks of shortened and normal (control) neighbour songs. Males responded significantly stronger to longer songs in both experiments as measured by the approach variable. Vocal response was not affected by treatment, but it was affected by the order of playback presentation. Our results indicate that song duration might be used for signalling current aggressive motivation during close interactions with rivals.     

Syllable repertoire and the size of the song control system in captive canaries (Serinus canaria)

In songbirds, there is considerable interest in relationships between song structure and the size of the song control system in the forebrain. In male canaries, earlier studies have reported that repertoire size increased with age, and positive correlations were obtained between repertoire size and the volume of song control nuclei such as high vocal center (HVC). Here we investigate whether age has an effect upon both the song structure and the morphology of two song control nuclei [HVC and robustus archistriatalis (RA)] that are important in song production. We recorded songs from an aviary population of 1- and 2-year-old male domesticated canaries. We found that repertoire size, number of sexually attractive (sexy) syllables, and size of song nuclei did not differ between 1- and 2-year-old males. Neither did we find significant correlations between syllable repertoire size and the size of the song control nuclei. However, HVC size was positively correlated with the proportion of sexy syllables in the repertoires of 2-year-old males. Some older males may enhance vocal performance by modifying the control of syllables rather than by increasing repertoire size or neural space.     

Manipulations of sensory experiences during development reveal mechanisms underlying vocal learning biases in zebra finches

Biological predispositions in learning can bias and constrain the cultural evolution of social and communicative behaviors (e.g., speech and birdsong), and lead to the emergence of behavioral and cultural “universals.” For example, surveys of laboratory and wild populations of zebra finches (Taeniopygia guttata) document consistent patterning of vocal elements (“syllables”) with respect to their acoustic properties (e.g., duration, mean frequency). Furthermore, such universal patterns are also produced by birds that are experimentally tutored with songs containing randomly sequenced syllables (“tutored birds”). Despite extensive demonstrations of learning biases, much remains to be uncovered about the nature of biological predispositions that bias song learning and production in songbirds. Here, we examined the degree to which “innate” auditory templates and/or biases in vocal motor production contribute to vocal learning biases and production in zebra finches. Such contributions can be revealed by examining acoustic patterns in the songs of birds raised without sensory exposure to song (“untutored birds”) or of birds that are unable to hear from early in development (“early‐deafened birds”). We observed that untutored zebra finches and early‐deafened zebra finches produce songs with positional variation in some acoustic features (e.g., mean frequency) that resemble universal patterns observed in tutored birds. Similar to tutored birds, early‐deafened birds also produced song motifs with alternation in acoustic features across adjacent syllables. That universal acoustic patterns are observed in the songs of both untutored and early‐deafened birds highlights the contribution motor production biases to the emergence of universals in culturally transmitted behaviors.     

Neighbor-stranger discrimination by song in a suboscine bird, the alder flycatcher, Empidonax alnorum

Bird song and its functions have been studied extensively formore than 50 years, but almost entirely in oscine passerines.Few studies have investigated any aspect of song in suboscinepasserines. This is significant because song development andthe extent of individual variation in song differs greatly betweenthese groups. Learning and auditory feedback play major rolesin song development in all oscines studied, but apparently nopart in song ontogeny in suboscines. The ability of territorialoscine males to discriminate between songs of neighbors andstrangers has received considerable attention, but this phenomenonis virtually unstudied in suboscines. We tested whether a suboscinebird, the alder flycatcher (Empidonax alnorum), was able todiscriminate between songs of neighbors and strangers despitelimited individual variation in song. We performed playbackexperiments to measure responses of males to songs of neighborsand strangers broadcast from the territory boundary shared bythe subject and the neighbor. Subjects responded more aggressivelyto songs of strangers than to songs of neighbors. These resultsfurther our understanding of the evolution of song and its functionsin suboscines by demonstrating that, similar to their oscinerelatives, they can discriminate between the songs of neighborsand strangers.     

Cross fostering experiments suggest that mice songs are innate

Background

Vocal learning is a central functional constituent of human speech, and recent studies showing that adult male mice emit ultrasonic sound sequences characterized as “songs” have suggested that the ultrasonic courtship sounds of mice provide a mammalian model of vocal learning.

Objectives

We tested whether mouse songs are learned, by examining the relative role of rearing environment in a cross-fostering experiment.

Methods and Findings

We found that C57BL/6 and BALB/c males emit a clearly different pattern of songs with different frequency and syllable compositions; C57BL/6 males showed a higher peak frequency of syllables, shorter intervals between syllables, and more upward frequency modulations with jumps, whereas BALB/c males produced more “chevron” and “harmonics” syllables. To establish the degree of environmental influences in mouse song development, sons of these two strains were cross-fostered to another strain of parents. Songs were recorded when these cross-fostered pups were fully developed and their songs were compared with those of male mice reared by the genetic parents. The cross-fostered animals sang songs with acoustic characteristics - including syllable interval, peak frequency, and modulation patterns - similar to those of their genetic parents. In addition their song elements retained sequential characteristics similar to those of their genetic parents'' songs.

Conclusion

These results do not support the hypothesis that mouse “song” is learned; we found no evidence for vocal learning of any sort under the conditions of this experiment. Our observation that the strain-specific character of the song profile persisted even after changing the developmental auditory environment suggests that the structure of these courtship sound sequences is under strong genetic control. Thus, the usefulness of mouse “song” as a model of mammalian vocal learning is limited, but mouse song has the potential to be an indispensable model to study genetic mechanisms for vocal patterning and behavioral sequences.     

Song characteristics track bill morphology along a gradient of urbanization in house finches (<Emphasis Type="Italic">Haemorhous mexicanus</Emphasis>)

Introduction

Urbanization can considerably impact animal ecology, evolution, and behavior. Among the new conditions that animals experience in cities is anthropogenic noise, which can limit the sound space available for animals to communicate using acoustic signals. Some urban bird species increase their song frequencies so that they can be heard above low-frequency background city noise. However, the ability to make such song modifications may be constrained by several morphological factors, including bill gape, size, and shape, thereby limiting the degree to which certain species can vocally adapt to urban settings. We examined the relationship between song characteristics and bill morphology in a species (the house finch, Haemorhous mexicanus) where both vocal performance and bill size are known to differ between city and rural animals.

Results

We found that bills were longer and narrower in more disturbed, urban areas. We observed an increase in minimum song frequency of urban birds, and we also found that the upper frequency limit of songs decreased in direct relation to bill morphology.

Conclusions

These findings are consistent with the hypothesis that birds with longer beaks and therefore longer vocal tracts sing songs with lower maximum frequencies because longer tubes have lower-frequency resonances. Thus, for the first time, we reveal dual constraints (one biotic, one abiotic) on the song frequency range of urban animals. Urban foraging pressures may additionally interact with the acoustic environment to shape bill traits and vocal performance.